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Chapter 3: Synapses
Postsynaptic Potentials (animation)
Release of Neurotransmitter (animation)
Cholinergic (animation)
Release of ACh (animation)
AChE Inactivates ACh (animation)
AChE Inhibitors (animation)
Opiate Narcotics
Heres another way to make learning an interactive Chapter 6: Vision Anatomy of the Eye
experience. Get prepared for class exams with chapter- The Retina (animation)
by-chapter online quizzes, a final test thats great for Virtual Reality Eye (virtual reality)
Blind Spot (Try It Yourself)
practice, web links, flashcards, a pronunciation glossary,
Color Blindness in Visual Periphery (Try It Yourself)
Try It Yourself experiential exercises (different from the Brightness Contrast (Try It Yourself)
ones in the text), and more. Visit today! Motion Aftereffect (Try It Yourself)
Visuo-Motor Control (Try It Yourself) New!
Due to contractual reasons certain ancillaries are available only in higher
education or U.S. domestic markets. Minimum purchases may apply to receive
the ancillaries at no charge. For more information, please contact your local
Thomson sales representative.
to provide multimedia support for each chapter!
Animations, Videos, and Activities
3 Synapses 51
6 Vision 151
8 Movement 231
v
Contents
Module 1.3
Summary 25
Module 1.1 Answers to Stop & Check Questions 25
The MindBrain Relationship 2
Chapter Ending Key Terms and Activities
Biological Explanations of Behavior 3
Terms 26
The Brain and Conscious Experience 5
Suggestions for Further Reading 26
Research Approaches 7
Websites to Explore 26
Career Opportunities 8
Exploring Biological Psychology CD 27
In Closing: Your Brain and Your Experience 10
ThomsonNOW 27
Summary 10
Answers to Stop & Check Questions 10
Thought Questions 11
Authors Answer About Machine
Consciousness 11 2
Module 1.2
The Genetics of Behavior 12
Nerve Cells and
Mendelian Genetics 12
Chromosomes and Crossing Over 13
Nerve Impulses 29
Sex-Linked and Sex-Limited Genes 13
Sources of Variation 14
Module 2.1
The Cells of the Nervous System 30
Heredity and Environment 14
Possible Complications 14 Anatomy of Neurons and Glia 30
Environmental Modification 15 EXTENSIONS AND APPLICATIONS Santiago
How Genes Affect Behavior 16 Ramn y Cajal, a Pioneer of Neuroscience 30
The Evolution of Behavior 16 The Structures of an Animal Cell 31
Common Misunderstandings About Evolution 16 The Structure of a Neuron 32
Evolutionary Psychology 19 Variations Among Neurons 34
Glia 35
In Closing: Genes and Behavior 20
The Blood-Brain Barrier 36
Summary 21 Why We Need a Blood-Brain Barrier 36
Answers to Stop & Check Questions 21 How the Blood-Brain Barrier Works 36
Thought Questions 21 The Nourishment of Vertebrate Neurons 37
vi
In Closing: Neurons 37 Relationship Among EPSP, IPSP,
and Action Potential 56
Summary 38
In Closing: The Neuron as Decision Maker 56
Answers to Stop & Check Questions 38
Summary 56
Module 2.2 Answers to Stop & Check Questions 56
The Nerve Impulse 39 Thought Questions 57
The Resting Potential of the Neuron 39
Forces Acting on Sodium and Potassium Ions 40 Module 3.2
Why a Resting Potential? 41 Chemical Events at the Synapse 58
The Action Potential 42 The Discovery of Chemical Transmission
The Molecular Basis of the Action Potential 43 at Synapses 58
The All-or-None Law 44
The Sequence of Chemical Events
The Refractory Period 44
at a Synapse 59
Propagation of the Action Potential 45 Types of Neurotransmitters 59
The Myelin Sheath and Saltatory Conduction 46 Synthesis of Transmitters 60
Local Neurons 47 Transport of Transmitters 61
Graded Potentials 47 Release and Diffusion of Transmitters 61
Activation of Receptors of the Postsynaptic Cell 62
EXTENSIONS AND APPLICATIONS Small Neurons
Inactivation and Reuptake of Neurotransmitters 66
and Big Misconceptions 47
Negative Feedback from the Postsynaptic Cell 67
In Closing: Neural Messages 47 Synapses and Personality 67
Summary 48 In Closing: Neurotransmitters and Behavior 68
Answers to Stop & Check Questions 48 Summary 68
Thought Questions 48 Answers to Stop & Check Questions 68
Chapter Ending Key Terms and Activities Thought Questions 69
Terms 49
Suggestions for Further Reading 49 Module 3.3
Websites to Explore 49 Drugs and Synapses 70
Exploring Biological Psychology CD 49 Drug Mechanisms 71
ThomsonNOW 50 Common Drugs and Their Synaptic Effects 71
Stimulant Drugs 71
Nicotine 73
Opiates 74
Marijuana 74
3 Hallucinogenic Drugs 75
In Closing: Drugs and Behavior 76
Summary 76
Answers to Stop & Check Questions 77
Synapses 51 Thought Question 77
Module 3.1 Chapter Ending Key Terms and Activities
The Concept of the Synapse 52 Terms 78
The Properties of Synapses 52 Suggestions for Further Reading 78
Speed of a Reflex and Delayed Transmission Websites to Explore 78
at the Synapse 53
Temporal Summation 53 Exploring Biological Psychology CD 78
Spatial Summation 53 ThomsonNOW 79
Inhibitory Synapses 54
Contents vii
Module 4.3
Research Methods 105
4 Correlating Brain Anatomy with Behavior
Recording Brain Activity 107
105
viii Contents
In Closing: Brain Development 135 The Eye and Its Connections to the Brain 153
The Route Within the Retina 153
Summary 135
Fovea and Periphery of the Retina 155
Answers to Stop & Check Questions 136
Visual Receptors: Rods and Cones 156
Thought Questions 136
Color Vision 157
The Trichromatic (Young-Helmholtz) Theory 158
Module 5.2 The Opponent-Process Theory 159
Plasticity After Brain Damage 137 The Retinex Theory 161
Brain Damage and Short-Term Recovery 137 Color Vision Deficiency 163
Visual Coding and the Retinal Receptors 152 Thought Question 183
General Principles of Perception 152
From Neuronal Activity to Perception 152 Module 6.3
Law of Specific Nerve Energies 152 Development of Vision 184
Infant Vision 184
Attention to Faces and Face Recognition 184
Visual Attention and Motor Control 184
Contents ix
Early Experience and Visual Development 185
Early Lack of Stimulation of One Eye 185 Module 7.2
Early Lack of Stimulation of Both Eyes 186 The Mechanical Senses 205
Uncorrelated Stimulation in the Two Eyes 186 Vestibular Sensation 205
Restoration of Response After Early Deprivation
of Vision 187
Somatosensation 206
Somatosensory Receptors 206
Early Exposure to a Limited Array of Patterns 187
People with Vision Restored After EXTENSIONS AND APPLICATIONS Tickle 206
Early Deprivation 188 Input to the Spinal Cord and the Brain 208
In Closing: The Nature and Nurture of Vision 190 Pain 209
Pain Stimuli and the Pain Pathways 209
Summary 190
Ways of Relieving Pain 210
Answers to Stop & Check Questions 191 Sensitization of Pain 212
Thought Questions 191 Itch 213
Chapter Ending Key Terms and Activities In Closing: The Mechanical Senses 213
Terms 192 Summary 213
Suggestions for Further Reading 192 Answers to Stop & Check Questions 214
Websites to Explore 192 Thought Question 214
Exploring Biological Psychology CD 193
ThomsonNOW 193 Module 7.3
The Chemical Senses 215
General Issues About Chemical Coding 215
Taste 216
x Contents
In Closing: Heredity and Environment in
Movement Disorders 261
8 Summary 261
Answers to Stop & Check Questions 261
Thought Questions 262
Chapter Ending Key Terms and Activities
Movement 231 Terms 262
Module 8.1 Suggestions for Further Reading 263
The Control of Movement 232 Websites to Explore 263
Muscles and Their Movements 232 Exploring Biological Psychology CD 263
Fast and Slow Muscles 234 ThomsonNOW 263
Muscle Control by Proprioceptors 235
Units of Movement 236
Voluntary and Involuntary Movements 236
EXTENSIONS AND APPLICATIONS
Infant Reflexes 236
Movements with Different Sensitivity
to Feedback 238
9
Sequences of Behaviors 238
In Closing: Categories of Movement 239 Wakefulness and Sleep 265
Summary 239
Module 9.1
Answers to Stop & Check Questions 239
Rhythms of Waking and Sleeping 266
Thought Question 239
Endogenous Cycles 266
Duration of the Human Circadian Rhythm 268
Module 8.2
Mechanisms of the Biological Clock 269
Brain Mechanisms of Movement 240 The Suprachiasmatic Nucleus (SCN) 269
The Cerebral Cortex 241 The Biochemistry of the Circadian Rhythm 270
Connections from the Brain to the Spinal Cord 243 Melatonin 271
Areas Near the Primary Motor Cortex 244 Setting and Resetting the Biological Clock 271
Conscious Decisions and Movements 245 Jet Lag 272
The Cerebellum 247 Shift Work 272
Evidence of a Broad Role 248 How Light Resets the SCN 273
Cellular Organization 249 In Closing: SleepWake Cycles 273
The Basal Ganglia 250
Summary 274
Brain Areas and Motor Learning 251
Answers to Stop & Check Questions 274
In Closing: Movement Control and Cognition 251
Thought Questions 274
Summary 252
Answers to Stop & Check Questions 252 Module 9.2
Thought Question 253 Stages of Sleep and Brain Mechanisms 275
The Stages of Sleep 275
Module 8.3 Paradoxical or REM Sleep 276
Disorders of Movement 254 Brain Mechanisms of Wakefulness
Parkinsons Disease 254 and Arousal 277
Possible Causes 254 Brain Structures of Arousal and Attention 278
L-Dopa Treatment 257 Getting to Sleep 280
Therapies Other Than L-Dopa 257 Brain Function in REM Sleep 281
Huntingtons Disease 258
Heredity and Presymptomatic Testing 259
Contents xi
Sleep Disorders 282 EXTENSIONS AND APPLICATIONS Surviving in
Sleep Apnea 283 Extreme Cold 298
Narcolepsy 283 The Advantages of Constant High
Periodic Limb Movement Disorder 284 Body Temperature 299
REM Behavior Disorder 284 Brain Mechanisms 300
Night Terrors, Sleep Talking, and Sleepwalking 284 Fever 300
xii Contents
Summary 320 Gender Identity and Gender-Differentiated
Answers to Stop & Check Questions 321 Behaviors 340
Intersexes 341
Thought Question 322
Interests and Preferences of CAH Girls 342
Chapter Ending Key Terms and Activities Testicular Feminization 343
Terms 322 Issues of Gender Assignment and Rearing 343
Discrepancies of Sexual Appearance 344
Suggestions for Further Reading 323
Possible Biological Bases of
Websites to Explore 323
Sexual Orientation 345
Exploring Biological Psychology CD 323 Genetics 345
ThomsonNOW 323 Hormones 346
Prenatal Events 347
Brain Anatomy 348
In Closing: We Are Not All the Same 349
11 Summary 349
Answers to Stop & Check Questions
Thought Questions 350
350
12
331
Rodents 331
Humans 331
EXTENSIONS AND APPLICATIONS
Premenstrual Syndrome 334
Parental Behavior 335 Emotional Behaviors 353
In Closing: Reproductive Behaviors and
Motivations 336 Module 12.1
What Is Emotion? 354
Summary 337
Emotions, Autonomic Response, and the
Answers to Stop & Check Questions 337
James-Lange Theory 354
Thought Questions 338 Is Physiological Arousal Necessary
for Emotions? 355
Module 11.2 Is Physiological Arousal Sufficient
Variations in Sexual Behavior 339 for Emotions? 355
Brain Areas Associated with Emotion 356
Evolutionary Interpretations of
Attempts to Localize Specific Emotions 357
Mating Behavior 339
Contributions of the Left and Right
Interest in Multiple Mates 339
Hemispheres 358
What Men and Women Seek in Their Mates 339
Differences in Jealousy 340 The Functions of Emotions 359
Evolved or Learned? 340 In Closing: Emotions and the
Conclusions 340 Nervous System 360
Contents xiii
Summary 360
Answers to Stop & Check Questions 360
Thought Question 360
Module 12.2
13
Attack and Escape Behaviors 361
Attack Behaviors 361 The Biology of Learning
Heredity and Environment in Violence 361
Hormones 363
and Memory 383
Brain Abnormalities and Violence 363
Module 13.1
Serotonin Synapses and Aggressive Behavior 364
Learning, Memory, Amnesia, and
Escape, Fear, and Anxiety 366 Brain Functioning 384
Fear, Anxiety, and the Amygdala 366
Studies of Rodents 366 Localized Representations of Memory 384
Studies of Monkeys 367 Lashleys Search for the Engram 384
Activation of the Human Amygdala 368 The Modern Search for the Engram 386
Damage to the Human Amygdala 369 Types of Memory 387
METHODS 12.1 Microdialysis 371 Short- and Long-Term Memory 387
Anxiety-Reducing Drugs 371 Working Memory 389
Thought Questions 375 Other Types of Brain Damage and Amnesia 398
Korsakoffs Syndrome and Other
Prefrontal Damage 398
Module 12.3 Alzheimers Disease 399
Stress and Health 376 What Patients with Amnesia Teach Us 401
Concepts of Stress 376 In Closing: Different Types of Memory 402
Stress and the Hypothalamus-Pituitary-Adrenal Summary 402
Cortex Axis 376
The Immune System 377 Answers to Stop & Check Questions 402
xiv Contents
In Closing: The Physiology of Memory 411 In Closing: One Brain, Two Hemispheres 427
14
Does Language Learning Have a
Critical Period? 434
Brain Damage and Language 435
Brocas Aphasia (Nonfluent Aphasia) 435
Wernickes Aphasia (Fluent Aphasia) 437
Cognitive Functions 415 Dyslexia 438
In Closing: Language and the Brain 440
Module 14.1
Lateralization of Function 416 Summary 440
Handedness and Its Genetics 416 Answers to Stop & Check Questions 441
Contents xv
Module 15.3
Schizophrenia 470
15 Characteristics 470
Behavioral Symptoms 470
EXTENSIONS AND APPLICATIONS Differential
Diagnosis of Schizophrenia 471
Psychological Disorders 451 Demographic Data 471
Genetics 472
Module 15.1
Twin Studies 472
Substance Abuse and Addictions 452 Adopted Children Who Develop Schizophrenia 472
Synapses, Reinforcement, and Addiction 452 Efforts to Locate a Gene 472
Reinforcement and the Nucleus Accumbens 452 The Neurodevelopmental Hypothesis 473
Addiction as Increased Wanting 452 Prenatal and Neonatal Environment 473
Sensitization of the Nucleus Accumbens 453 Mild Brain Abnormalities 474
Alcohol and Alcoholism 454 METHODS 15.1 The Wisconsin Card Sorting Task 475
Genetics 454 Early Development and Later Psychopathology 476
Risk Factors 455
Neurotransmitters and Drugs 477
Medications to Combat Substance Abuse 456 Antipsychotic Drugs and Dopamine 477
Antabuse 456 Role of Glutamate 478
Methadone 456 New Drugs 479
In Closing: Addictions 457 In Closing: The Fascination of Schizophrenia 480
Summary 457 Summary 480
Answers to Stop & Check Questions 458 Answers to Stop & Check Questions 481
Thought Question 458 Thought Questions 481
Chapter Ending Key Terms and Activities
Module 15.2
Mood Disorders 459 Terms 482
Suggestions for Further Reading 482
Major Depressive Disorder 459
Genetics and Life Events 459 Websites to Explore 483
Hormones 460 Exploring Biological Psychology CD 483
Abnormalities of Hemispheric Dominance 461 ThomsonNOW 483
Viruses 461
Antidepressant Drugs 462
EXTENSIONS AND APPLICATIONS Accidental
A Brief, Basic Chemistry 485
Discoveries of Psychiatric Drugs 462
Other Therapies 464
B Society for Neuroscience
Bipolar Disorder 466 Policies on the Use of Animals
Genetics 467
and Human Subjects in
Treatments 467
Neuroscience Research 491
Seasonal Affective Disorder (SAD) 467
In Closing: The Biology of Mood Swings 468 References 494
Summary 468 Name Index 550
Answers to Stop & Check Questions 469
Subject Index/Glossary 565
Thought Question 469
xvi Contents
Preface
xvii
New explanation of binding with an improved Try- Reorganized section on theories of the need for sleep.
It-Yourself activity. New examples of sleep specializations in other spe-
New section discussing the research on the relation- cies: dolphins, migratory birds, European swifts (who
ship between brain size and intelligence. Discussion sleep while flying).
of species differences in brain anatomy moved here Added information about sleep in astronauts, sub-
from Chapter 5. marine sailors, and people working in Antarctica.
Chapter 5 Chapter 10
Revised order of topics in both modules. Several new examples of seemingly odd animal
Experiment on reorganization of the infant ferret behaviors that make sense in terms of temperature
cortex revised and moved here from Chapter 6. regulation.
New discussion of brain changes that result from A completely rewritten section on brain mecha-
lifelong blindness. nisms of feeding.
Discussion of evidence suggesting that consum-
Chapter 6 ing high-fructose corn syrup increases the risk of
New examples of species differences in vision. obesity.
Updated discussion of blindsight, face recognition,
motion blindness, and visual attention. Chapter 11
Added several new studies of the development of Revised discussion of hormonal effects on intellec-
vision, including people who had vision restored tual performance.
in adulthood after having had little or none since New study included that shows that one gene con-
early childhood. trolling vasopressin can alter social behaviors, caus-
ing male meadow voles to establish pair bonds with
Chapter 7 females and help them rear babiesa behavior never
Much expanded discussion of the auditory cortex, previously seen in males of this species.
including parallels between the auditory and visual Much revised discussion of gender identity and
systems. gender-differentiated behaviors in people with con-
Neuropsychological studies of a patient who can- genital adrenal hyperplasia.
not integrate vestibular sensation with other senses Several updates about homosexuality including:
and therefore has out of body experiences. the probability of homosexuality is increased among
Neuropsychological studies of a patient who has no boys with older brothers; and men with a homosex-
conscious touch sensation but nevertheless feels ual orientation have female relatives who have a
pleasure when touched. greater than average number of childrena possi-
Reorganized discussion of pain. ble explanation for maintenance of a gene promot-
New research added showing that chronic pain de- ing homosexuality.
pends on a mechanism related to learning.
New section added on synesthesia, the tendency of Chapter 12
certain people to experience one sense in response Substantial updating and revision throughout this
to stimulation of a different sense. chapter.
Clarification of the James-Lange theory and evidence
Chapter 8 relevant to it.
Added mirror neurons in the motor cortex that Monkeys with low serotonin turnover become ag-
respond both to ones own movements and the sight gressive and are likely to die young, but if they sur-
of other people doing the same movements. vive, they are likely to achieve dominant status.
New section on the relationship between conscious The human amygdala responds most strongly to
decisions and movements. emotional stimuli that are sufficiently ambiguous
to require processing.
Chapter 9 People with amygdala damage fail to identify fear
New material added on the differences between in photographs because they focus their vision al-
morning people and evening people. most entirely on the nose and mouth.
New research included on the role of orexin in main- Genetic variance in the amygdala probably con-
taining wakefulness. tributes to variance in predisposition to anxiety
GABA release during sleep does not decrease neu- disorders.
ronal activity, but decreases the spread of excitation Stress module: Deleted the discussion of psycho-
at synapses. Neuronal activity continues, although somatic illness and expanded discussion of stress
much of it is not conscious. and the immune system.
xviii Preface
New evidence indicates that people with smaller comprehensive final exam. The test items are also avail-
than average hippocampus are predisposed to PTSD. able electronically on Examview. The Study Guide,
written by Elaine M. Hull of Florida State University,
Chapter 13 may be purchased by students. Also available is the
New studies on patient H.M.: If tested carefully, he Multimedia Manager Instructors Resource CD-ROM,
shows slight evidence of new declarative memo- written by Chris Hayashi, Southwestern College.
ries since his operation, although no new episodic I am grateful for the excellent work of Darin Der-
memories. stine, Cynthia Crawford, JeffreyStowell, Elaine Hull,
Brief new discussion of individual differences in and Chris Hayashi.
the hippocampus and their relationship to differ- In addition, it is possible to use technology in a
ences in memory. variety of ways in your course with the following new
Reorganized discussion of consolidation of memory. products:
Updates added on Alzheimers disease, including
some new prospects for treatment. JoinIn on TurningPoint
Exclusive from Thomson for colleges and universi-
Chapter 14 ties . . . turn your lecture into an interactive experi-
New section on the genetics of handedness. ence for your students, using clickers.
Revised module on attention.
WebTutor Advantage
Chapter 15 Save time managing your course, posting materials,
The first module now deals with substance abuse incorporating multimedia, and tracking progress with
and addiction, but not the mechanisms of drugs in this engaging, text-specific e-learning tool. Visit http://
general. That section is now in Chapter 3. webtutor.thomsonlearning.com.
Greatly revised discussion of addiction, with more
explanation of the distinction between wanting and ThomsonNow
liking. A powerful, assignable, personalized online learning
Evidence now says depression relates more to lack companion that assesses individual study needs and
of happiness than to increased sadness. builds focused Personalized Learning Plans that rein-
New evidence relates depression to an interaction force key concepts with interactive animations, text
between a gene and a series of stressful experiences. art, and more.
New evidence on the genetics of schizophrenia.
New evidence suggests a parasitic infection in child-
hood can predispose someone to schizophrenia later.
Reorganized discussion of antipsychotic drugs and
Acknowledgments
their relationship to neurotransmitters. Let me tell you something about researchers in this
field: As a rule, they are amazingly cooperative with
textbook authors. Many of my colleagues sent me com-
Supplements ments, ideas, articles, and photos. I thank especially the
following:
The CD-ROM that accompanies this text includes ani-
Greg Allen, University of Texas Southwestern
mations, film clips, Try It Yourself activities, quizzes,
Medical Center
and other supplements to the text. Darin Derstine took
Ralph Adolphs, University of Iowa
responsibility for coordinating the CD, working with
Danny Benbassat, Ohio Northern University
Rob Stufflebeam, University of New Orleans, and me
Stephen L. Black, Bishops University
on the new online Try It Yourself activities. Those
Martin Elton, University of Amsterdam
who adopt the book may also obtain from the pub-
Jane Flinn, George Mason University
lisher a copy of the Instructors Manual, written by
Ronnie Halperin, SUNY-Purchase
Cynthia Crawford, California State University at San
Julio Ramirez, Davidson College
Bernardino. The manual contains chapter outlines,
Sarah L. Pallas, Georgia State University
class demonstrations and projects, a list of video re-
Alex Pouget, University of Rochester
sources, additional Websites, InfoTrac Virtual Reader,
Robert Provine, University of Maryland,
and the authors answers to the Thought Questions.
Baltimore County
A separate print Testbank lists multiple-choice and
Roberto Refinetti, University of South Carolina
truefalse items written and assembled by Jeffrey
Stowell, Eastern Illinois University. Note the test bank I have received an enormous number of letters
includes special files of questions for a midterm and a and e-mail messages from students. Many included
Preface xix
helpful suggestions; some managed to catch errors or In preparing this text I have been most fortunate
inconsistencies that everyone else had overlooked. I to work with Vicki Knight, a wise, patient, and very
thank especially the following: supportive acquisitions editor/publisher. She was es-
pecially helpful in setting priorities and planning the
Jacqueline Counotte, Leiden University,
major thrust of this text. Kirk Bomont, my develop-
Netherlands
mental editor, reads manuscripts with extraordinary
Terry Fidler, University of Victoria, British
care, noticing discrepancies, unclear points, and ideas
Columbia
that need more explanation. His work helped me enor-
Paul Kim, N. C. State University
mously in the preparation of this edition. Karol Ju-
Florian van Leeuwen, University of Groningen,
rado, Content Project Manager, did a stellar job in co-
Netherlands
ordinating the production process and working closely
Elizabeth Rose Murphy, North Carolina State
with all of the players, including Nancy Shammas at
University
New Leaf Publishing Services who provided the ser-
Steve Williams, Massey University, New Zealand
vice for the book and undertook the management of
I appreciate the helpful comments and suggestions all of the talented people who contributed to the pro-
provided by the following reviewers who commented duction of this booka major task for a book with
on the 8th edition and provided suggestions for the such a large art and photo program. As art editor, Lisa
9th edition, and/or who reviewed the revised manu- Torris considerable artistic abilities helped to com-
script for the 9th edition: pensate for my complete lack. And once again, Preci-
sion Graphics did an outstanding job with modifica-
Joseph Porter, Virginia Commonwealth University
tions on the art and new renderings. Joohee Lee
Marjorie Battaglia, George Mason University
handled all of the permissions, no small task for a
Anne Marie Brady, St. Marys College of Maryland
book like this. Eric Schrader was the photo researcher;
Linda James, Georgian Court University
I hope you enjoy the new photos in this text as much
Mary Clare Kante, University of Illinois at
as I do. Jennifer Wilkinson oversaw the development
Chicago Circle
of supplements, such as the Instructors Manual and
Frank Scalzo, Bard College
test item file. I thank Vernon Boes, who managed the
Nancy Woolf, University of California, Los Angeles
interior design and the cover, Tani Hasegawa for the
Joseph Dien, University of Kansas
outstanding changes to the interior design, Frank Hu-
Derek Hamilton, University of New Mexico
bert for the copyediting, Linda Dane for the proofread-
Alexander Kusnecov, Rutgers University
ing, and Do Mi Stauber for the indexes. All of these
Ronald Baenninger, College of St. Benedict/
people have been splendid colleagues.
St. Johns University
I thank my wife, Jo Ellen, for keeping my spirits
Christine Wagner, SUNY, Albany
high, and my department head, David Martin, for his
Amira Rezec, Saddleback College
support and encouragement. I especially thank my son
Brian Kelley, Bridgewater College
Sam for many discussions and many insightful ideas.
Lisa Baker, Western Michigan University
Sam Kalat, coming from a background of biochemistry
Steven Brown, Rockhurst University
and computer science, has more original and insight-
Chris Bloom, University of Southern Indiana
ful ideas about brain functioning than anyone else
Anthony Risser, University of Houston
I know.
Douglas Grimsley, University of North Carolina,
I welcome correspondence from both students
Charlotte
and faculty. Write: James W. Kalat, Department of Psy-
Yuan B. Peng, University of Texas at Arlington
chology, Box 7650, North Carolina State University,
Carlota Ocampo, Trinity University
Raleigh, NC 276957801, USA. E-mail: james_kalat
Ron Salazar, San Juan College
@ncsu.edu
James W. Kalat
xx Preface
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Sincerely yours,
James W. Kalat
Biological
Psychology
1
The Major Issues
Frontal
lobe of Longitudinal
cerebral fissure
Frontal lobe cortex
Olfactory
Precentral gyrus bulbs
Temporal
Central sulcus Anterior lobe of
cerebral Optic
Postcentral gyrus cortex nerves
Parietal lobe
Posterior Medulla
Dr. Dana Copeland
Figure 1.1 A dorsal view (from above) and a ventral view (from below) of the
human brain
The brain has an enormous number of divisions and subareas; the labels point to a few
of the main ones on the surface of the brain.
Dorr/Premium Stock/PictureQuest
verse exists at all: Why is there something instead of
nothing? And if there is something, why is it this par-
ticular kind of something? Biological psychologists
ask: Given the existence of this universe composed of
matter and energy, why is there consciousness? Is it a
necessary function of the brain or an accident? How
Researchers continue to debate exactly what good yawning
does the brain produce consciousness and why?
does. Yawning is a behavior that even people do without
Researchers also ask more specific questions such
knowing its purpose.
as: What genes, prenatal environment, or other factors
predispose some people to psychological disorders?
Is there any hope for recovery after brain damage? And
what enables humans to learn language so easily?
Biological Explanations
of Behavior
Commonsense explanations of behavior often refer
to intentional goals such as, He did this because he
was trying to . . . or She did that because she wanted
A
5
3
2
Conscious Experience c
4
6
50
B
2 5
3
Explaining birdsong in terms of hormones,
1
brain activity, and evolutionary selection
probably does not trouble you. But how
B C
would you feel about a physical explana-
tion of your own actions and experiences?
Suppose you say, I became frightened be-
cause I saw a man with a gun, and a neu- Figure 1.5 Ren Descartess conception of brain and mind
roscientist says, You became frightened Descartes understood how light from an object reached the retinas at
because of increased electrochemical ac- the back of the eyes. From there, he assumed the information was all
tivity in the central amygdala of your channeled back to the pineal gland, a small unpaired organ in the brain.
brain. Is one explanation right and the (Source: From Descartes Treaties on Man)
problem. The questions remain: Why is consciousness leads to consciousness or causes consciousness. According
to the identity position, brain activity does not cause or lead to
a property of brain activity? Is it important or just an consciousness any more than consciousness leads to brain activ-
accident, like the noises a machine makes? What kind ity. Each is the same as the other.
Research Fields Research positions ordinarily require a PhD. Researchers are employed by universities,
hospitals, pharmaceutical firms, and research institutes.
Neuroscientist Studies the anatomy, biochemistry, or physiology of the nervous system. (This broad term
includes any of the next five, as well as other specialties not listed.)
Behavioral neuroscientist Investigates how functioning of the brain and other organs influences behavior.
(almost synonyms: psychobiologist,
biopsychologist, or physiological
psychologist).
Cognitive neuroscientist Uses brain research, such as scans of brain anatomy or activity, to analyze and explore
peoples knowledge, thinking, and problem solving.
Neuropsychologist Conducts behavioral tests to determine the abilities and disabilities of people with various
kinds of brain damage and changes in their condition over time. Most neuropsychologists
have a mixture of psychological and medical training; they work in hospitals and clinics.
Psychophysiologist Measures heart rate, breathing rate, brain waves, and other body processes and how they
vary from one person to another or one situation to another.
Neurochemist Investigates the chemical reactions in the brain.
Comparative psychologist Compares the behaviors of different species and tries to relate them to their habitats and
(almost synonyms: ethologist, ways of life.
animal behaviorist)
Evolutionary psychologist Relates behaviors, especially social behaviors, including those of humans, to the functions
(almost synonym: sociobiologist) they have served and, therefore, the presumed selective pressures that caused them to evolve.
Practitioner Fields of Psychology In most cases, their work is not directly related to neuroscience. However, practitioners
often need to understand it enough to communicate with a clients physician.
Clinical psychologist Requires PhD or PsyD. Employed by hospital, clinic, private practice, or college. Helps
people with emotional problems.
Counseling psychologist Requires PhD or PsyD. Employed by hospital, clinic, private practice, or college. Helps
people make educational, vocational, and other decisions.
School psychologist Requires masters degree or PhD. Most are employed by a school system. Identifies educa-
tional needs of schoolchildren, devises a plan to meet the needs, and then helps teachers
implement it.
Medical Fields Practicing medicine requires an MD plus about 4 years of additional study and practice
in a specialization. Physicians are employed by hospitals, clinics, medical schools and in
private practice. Some conduct research in addition to seeing patients.
Neurologist Treats people with brain damage or diseases of the brain.
Neurosurgeon Performs brain surgery.
Psychiatrist Helps people with emotional distress or troublesome behaviors, sometimes using drugs
or other medical procedures.
Allied Medical Field These fields ordinarily require a masters degree or more. Practitioners are employed
by hospitals, clinics, private practice, and medical schools.
Physical therapist Provides exercise and other treatments to help people with muscle or nerve problems, pain,
or anything else that impairs movement.
Occupational therapist Helps people improve their ability to perform functions of daily life, for example, after a
stroke.
Social worker Helps people deal with personal and family problems. The activities of a clinical social
worker overlap those of a clinical psychologist.
as physical therapy. These various fields of practice Anyone who pursues a career in research needs to
range from neurologists (who deal exclusively with stay up to date on new developments by attending con-
brain disorders) to social workers and clinical psy- ventions, consulting with colleagues, and reading the
chologists (who need to distinguish between adjust- primary research journals, such as Journal of Neuro-
ment problems and possible signs of brain disorder). science, Neurology, Behavioral Neuroscience, Brain
Figure 1.7
How DNA controls DNA
Self-replicating
development of molecule
the organism
The sequence of bases
along a strand of DNA Each base determines one base of the RNA.
determines the order
of bases along a strand
of RNA; RNA in turn
RNA
controls the sequence of Copy of one strand
amino acids in a protein of the DNA
molecule.
135
Millions of years ago
65
54
38
Bats
25 Horses and
Monkeys
rhinoceroses
7
Rabbits
Carnivores Humans
Whales
2.5
and dolphins
0
Platypus Elephants Cattle and Insectivores Apes Rodents Marsupials
sheep
(b)
that feature? You may have heard people say some- Jean Lamarcks theory of evolution through the inher-
thing like, Because we hardly ever use our little toes, itance of acquired characteristics, known as Lamarck-
they will get smaller and smaller in each succeeding ian evolution. According to this idea, if giraffes stretch
generation. This idea is a carryover of the biologist their necks as far out as possible, their offspring will
F. J. Hierschel/Okapia/Photo Researchers
toe sizewithout causing some other problemand
then we would have to wait for people with this mu-
tation to outreproduce people with other genes.)
Have humans stopped evolving? Because mod-
ern medicine can keep almost anyone alive, and be-
cause welfare programs in prosperous countries pro-
vide the necessities of life for almost everyone, some
people assert that humans are no longer subject to the
principle of survival of the fittest. Therefore, the ar- Sometimes a sexual display, such as a peacocks spread
gument goes, human evolution has slowed or stopped. of its tail feathers, leads to great reproductive success
The flaw in this argument is that the key to evolu- and therefore to the spread of the associated genes. In
tion is not survival but reproduction. For you to spread a slightly changed environment, this gene could become
your genes, of course you have to survive long enough maladaptive. For example, if an aggressive predator with
to reproduce, but what counts is how many healthy good color vision enters the range of the peacock, the
children (and nieces and nephews) you have. Thus, birds slow movement and colorful feathers could seal its
keeping everyone alive doesnt stop human evolution. doom.
If some people have more children than others do, their
genes will spread in the population.
Does evolution mean improvement? It de- ulation, which is operationally defined as the number
pends on what you mean by improvement. By defini- of copies of ones genes that endure in later genera-
tion, evolution improves the average fitness of the pop- tions. For example, if you have more children than av-
erage, you are by definition evolutionarily fit, regardless
of whether you are successful in any other sense. You
also increase your fitness by supporting your brother,
sister, nieces and nephews, or anyone else with the
same genes you have. Any gene that spreads is by def-
inition fit. However, genes that increase fitness at one
time and place might be disadvantageous after a change
in the environment. For example, the colorful tail feath-
ers of the male peacock enable it to attract females but
might become disadvantageous in the presence of a
new predator that responds to bright colors. In other
words, the genes of the current generation evolved be-
cause they were fit for previous generations; they may
Alain Le Garsmeur/CORBIS
Animals are used in many kinds of research studies, some dealing with behavior and others with
the functions of the nervous system.
Answers to
STOP & CHECK
STOP & CHECK
1. Describe reasons biological psychologists conduct Questions
much of their research on nonhuman animals.
1. Sometimes the mechanisms of behavior are easier
2. How does the minimalist position differ from the to study in a nonhuman species. We are curious
abolitionist position? about animals for their own sake. We study animals
Check your answers on this page. to understand human evolution. Certain proce-
dures are illegal or unethical with humans. (p. 25)
2. A minimalist wishes to limit animal research to
studies with little discomfort and much potential
value. An abolitionist wishes to eliminate all an-
imal research, regardless of how the animals are
treated or how much value the research might pro-
duce. (p. 25)
Terms
altruistic behavior (p. 19) evolutionary psychology (p. 19) monozygotic twins (p. 14)
artificial selection (p. 16) fitness (p. 18) multiplier effect (p. 15)
autosomal gene (p. 13) functional explanation (p. 4) mutation (p. 14)
binocular rivalry (p. 8) gene (p. 12) ontogenetic explanation (p. 4)
biological psychology (p. 2) hard problem (p. 6) phenylketonuria (PKU) (p. 15)
chromosome (p. 12) heritability (p. 14) physiological explanation (p. 3)
crossing over (p. 13) heterozygous (p. 12) problem of other minds (p. 6)
deoxyribonucleic acid (DNA) homozygous (p. 12) recessive (p. 13)
(p. 12) identity position (p. 6) reciprocal altruism (p. 20)
dizygotic twins (p. 14) kin selection (p. 20) recombination (p. 14)
dominant (p. 13) Lamarckian evolution (p. 17) ribonucleic acid (RNA) (p. 12)
dualism (p. 5) materialism (p. 5) sex-limited gene (p. 14)
easy problems (p. 6) mentalism (p. 6) sex-linked gene (p. 13)
enzyme (p. 12) mindbody or mindbrain solipsism (p. 6)
evolution (p. 16) problem (p. 5) X chromosome (p. 13)
evolutionary explanation (p. 4) monism (p. 5) Y chromosome (p. 13)
26 Chapter Ending
ter, click the appropriate chapter, and then find the
appropriate links to additional sites. You can also check
for suggested articles available on InfoTrac College
Edition. The sites for this chapter are:
http://www.thomsonedu.com
National Society for Phenylketonuria Home Page Go to this site for the link to ThomsonNOW, your one-stop study
http://www.nspku.org shop. Take a Pre-Test for this chapter, and ThomsonNOW will
generate a Personalized Study Plan based on your test results.
Statement on Use of Animals in Research
The Study Plan will identify the topics you need to review and
http://www.esf.org/ftp/pdf/SciencePolicy/ESPB9.pdf direct you to online resources to help you master these topics.
U.S. government statement on animal care and use You can then take a Post-Test to help you determine the con-
cepts you have mastered and what you still need to work on.
http://oacu.od.nih.gov/index.htm
Here are three sites that you may find helpful at many
points throughout the text:
Dana Foundation for brain information
http://www.dana.org
Exploring Biological
Psychology CD
Binocular Rivalry (Try It Yourself)
Genetics and Evolution (Try It Yourself)
Evolutionary Studies (video)
Offspring of Parents Homozygous and Heterozygous
for Brown Eyes (animation)
RNA, DNA, and Protein (animation)
Selection and Random Drift (Try It Yourself)
Critical Thinking (essay questions)
Chapter Quiz (multiple-choice questions)
Chapter Ending 27
2
Nerve Cells and
Nerve Impulses
Terms
Suggestions for Further Reading
Websites to Explore
A nervous system, composed of many individual
cells, is in some regards like a society of people
who work together and communicate with one an-
Exploring Biological Psychology CD other or even like elements that form a chemical com-
ThompsonNOW pound. In each case, the combination has properties
that are unlike those of its individual components. We
begin our study of the nervous system by examining
single cells; later, we examine how cells act together.
Advice: Parts of this chapter and the next assume
that you understand basic chemical concepts such as
positively charged ions. If you need to refresh your
memory, read Appendix A.
29
Module 2.1
The Cells of the
Nervous System
Anatomy of
Neurons and Glia
Spinal cord:
The nervous system consists of two kinds of cells: neu- 1 billion neurons
rons and glia. Neurons receive information and trans-
mit it to other cells. Glia provide a number of functions
that are difficult to summarize, and we shall defer that
discussion until later in the chapter. According to one
estimate, the adult human brain contains approximately
100 billion neurons (R. W. Williams & Herrup, 1988)
(Figure 2.1). An accurate count would be more difficult
than it is worth, and the actual number varies from per-
son to person.
The idea that the brain is composed of individual
cells is now so well established that we take it for
granted. However, the idea was in doubt as recently
as the early 1900s. Until then, the best microscopic
views revealed little detail about the organization of
the brain. Observers noted long, thin fibers between
Figure 2.1 Estimated numbers of neurons
one neurons cell body and another, but they could not
in humans
see whether each fiber merged into the next cell or
Because of the small size of many neurons and the variation
stopped before it (Albright, Jessell, Kandel, & Posner,
in cell density from one spot to another, obtaining an accurate
2001). Then, in the late 1800s, Santiago Ramn y Cajal
count is difficult. (Source: R. W. Williams & Herrup, 1988)
used newly developed staining techniques to show
that a small gap separates the tips of one neurons fibers
from the surface of the next neuron. The brain, like
the rest of the body, consists of individual cells. whom we shall discuss in Chapter 3; the other was the
Spanish investigator Santiago Ramn y Cajal (1852
1934). (See photo and quote on the pages inside the
E X T E N S I O N S A N D A P P L I C AT I O N S back cover.) Cajals early career did not progress alto-
Santiago Ramn y Cajal, gether smoothly. At one point, he was imprisoned in a
solitary cell, limited to one meal a day, and taken out
a Pioneer of Neuroscience
daily for public floggingsat the age of 10for the
Two scientists are widely recognized as the main found- crime of not paying attention during his Latin class (Ca-
ers of neuroscience. One was Charles Sherrington, jal, 1937). (And you thought your teachers were strict!)
(nuclear (ribosomes)
envelope)
(nucleolus)
Endoplasmic reticulum
(isolation, modification, transport
Nucleus of proteins and other substances)
(membrane-enclosed region
containing DNA; hereditary control)
Plasma membrane
(control of material Mitochondrion
exchanges, mediation of cell- (aerobic energy
environment interactions) metabolism)
Figure 2.2 An electron micrograph of parts of a neuron from the cerebellum of a mouse
The nucleus, membrane, and other structures are characteristic of most animal cells. The plasma
membrane is the border of the neuron. Magnification approximately x 20,000. (Source: Micrograph
courtesy of Dennis M. D. Landis)
Sensory
Axon endings
Nucleus Skin
Soma
surface
Efferent
(from A)
Variations Among Neurons
Neurons vary enormously in size, shape, and func-
tion. The shape of a given neuron determines its con-
nections with other neurons and thereby determines
Figure 2.8 Cell structures and axons its contribution to the nervous system. The wider the
It all depends on the point of view. An axon from A to B is branching, the more connections with other neurons.
an efferent axon from A and an afferent axon to B, just as a The function of a neuron is closely related to its
train from Washington to New York is exiting Washington shape (Figure 2.9). For example, the dendrites of the
and approaching New York. Purkinje cell of the cerebellum (Figure 2.9a) branch
extremely widely within a single plane; this cell is ca-
tem. Within the nervous system, a given neuron is an pable of integrating an enormous amount of incoming
efferent from the standpoint of one structure and an information. The neurons in Figures 2.9c and 2.9e also
afferent from the standpoint of another. (You can re- have widely branching dendrites that receive and in-
member that efferent starts with e as in exit; afferent tegrate information from many sources. By contrast,
starts with a as in admission.) For example, an axon certain cells in the retina (Figure 2.9d) have only short
that is efferent from the thalamus may be afferent to branches on their dendrites and therefore pool input
the cerebral cortex (Figure 2.8). If a cells dendrites from only a few sources.
Apical
dendrite
Dendrites
Basilar
dendrites
Axon
(a)
Axon
(c)
10 m
(b) (d)
Schwann
cell
Astrocyte
Capillary
(small blood vessel)
Radial glia
Axon
Migrating neuron
Microglia
Microglia
Fat-
solu
STOP & CHECK
ble
mol
ecu
le
3. What is one major advantage of having a blood-brain
Gluc barrier?
ose
tran 4. What is a disadvantage of the blood-brain barrier?
spor
t
5. Which chemicals cross the blood-brain barrier on
Ami their own?
no-a
cid 6. Which chemicals cross the blood-brain barrier by
tran
spor
t active transport?
Check your answers on page 38.
Charged
molecules
+
CO2
Cell wall tight
The Nourishment of
junction Vertebrate Neurons
CO2
Most cells use a variety of carbohydrates and fats for
Endothelial cell nutrition, but vertebrate neurons depend almost en-
O2 tirely on glucose, a simple sugar. (Cancer cells and the
Large
testis cells that make sperm also rely overwhelmingly
O2
molecule on glucose.) The metabolic pathway that uses glucose
requires oxygen; consequently, the neurons consume
an enormous amount of oxygen compared with cells
of other organs (Wong-Riley, 1989).
Blood vessel Brain tissue Why do neurons depend so heavily on glucose?
Although neurons have the enzymes necessary to me-
tabolize fats and several sugars, glucose is practically
Figure 2.12 The blood-brain barrier
the only nutrient that crosses the blood-brain bar-
Most large molecules and electrically charged molecules
rier in adults. The exceptions to this rule are ketones
cannot cross from the blood to the brain. A few small,
(a kind of fat), but ketones are seldom available in
uncharged molecules such as O2 and CO2 cross easily; so
large amounts (Duelli & Kuschinsky, 2001), and large
can certain fat-soluble molecules. Active transport systems
amounts of ketones cause medical complications.
pump glucose and amino acids across the membrane.
Although neurons require glucose, a glucose short-
age is rarely a problem. The liver can make glucose
from many kinds of carbohydrates and amino acids,
If the blood-brain barrier is such a good defense, as well as from glycerol, a breakdown product from
you might ask, why dont we have similar walls fats. An inability to use glucose can be a problem, how-
around our other organs? The answer is that the bar- ever. Many chronic alcoholics have a diet deficient in
rier that keeps out harmful chemicals also keeps out vitamin B1, thiamine, a chemical that is necessary for
many useful ones, including sources of nutrition. For the use of glucose. Prolonged thiamine deficiency can
organs that can afford to risk a viral infection, a tight lead to death of neurons and a condition called Kor-
barrier would be more costly than it is worth. Getting sakoffs syndrome, marked by severe memory impair-
nutrition into the brain requires an active transport, a ments (Chapter 13).
protein-mediated process that expends energy to pump
chemicals from the blood into the brain. Chemicals
that are actively transported into the brain include glu- Module 2.1
cose (the brains main fuel), amino acids (the building
In Closing: Neurons
blocks of proteins), and certain vitamins and hormones
(Brightman, 1997). The brain also has an active trans- What does the study of individual neurons tell us about
port system for moving certain chemicals from the brain behavior? Perhaps the main lesson is that our experi-
to the blood (King, Su, Chang, Zuckerman, & Paster- ence and behavior do not follow from the properties
nak, 2001). of any one neuron. Just as a chemist must know about
Answers to
Summary
STOP & CHECK
1. In the late 1800s, Santiago Ramn y Cajal used
newly discovered staining techniques to establish
Questions
that the nervous system is composed of separate 1. Dendrites, soma (cell body), axon, and presynaptic
cells, now known as neurons. (p. 30) terminal (p. 36)
2. Neurons receive information and convey it to other 2. Astrocytes (p. 36)
cells. The nervous system also contains glia.
3. The blood-brain barrier keeps out most viruses,
(pp. 30, 35)
bacteria, and other harmful substances. (p. 37)
3. Neurons have four major parts: a cell body, den-
4. The blood-brain barrier also keeps out most nutri-
drites, an axon, and presynaptic terminals. Their
ents. (p. 37)
shapes vary greatly depending on their functions
and their connections with other cells. (p. 32) 5. Small, uncharged molecules such as oxygen and
carbon dioxide cross the blood-brain barrier pas-
4. Glia do not convey information over great dis-
sively. So do chemicals that dissolve in the fats of
tances, but they aid the functioning of neurons in
the membrane. (p. 37)
many ways. (p. 35)
6. Glucose, amino acids, and some vitamins and hor-
mones cross by active transport. (p. 37)
Intracellular Amplifier
microelectrode
Reference
Soma microelectrode
Fritz Goro
Axon Computer
(a) (b)
as Figure 2.13 shows. The diameter of the electrode As we shall see in Chapter 3, certain kinds of stimula-
must be as small as possible so that it can enter the cell tion can open the sodium channels. When the mem-
without causing damage. By far the most common elec- brane is at rest, potassium channels are nearly but not
trode is a fine glass tube filled with a concentrated salt entirely closed, so potassium flows slowly.
solution and tapering to a tip diameter of 0.0005 mm Sodium ions are more than ten times more con-
or less. This electrode, inserted into the neuron, is centrated outside the membrane than inside because
connected to recording equipment. A reference elec-
trode placed somewhere outside the cell completes
the circuit. Connecting the electrodes to a voltmeter,
we find that the neurons interior has a negative poten-
tial relative to its exterior. The actual potential varies Membrane
K+
from one neuron to another; a typical level is 70 milli- of neuron
volts (mV), but it can be either higher or lower than that.
60
STOP & CHECK
mV
65
65 action potential?
20 1 ms
40 Time
K+
K+
+
Propagation of the K+
+
Action Potential Na+
Na+
+ + + +
+ +
Local current flow
+ +
Myelin + + + +
sheath Na+
Axon (a)
Node of Na+
Ranvier + + + +
+ + +
K Local current flow
+ +
+ + + +
Cutaway view of axon wrapped in myelin
Na+
Figure 2.18 An axon surrounded by a myelin
(b)
sheath and interrupted by nodes of Ranvier
The inset shows a cross-section through both the axon and Figure 2.19 Saltatory conduction in a
the myelin sheath. Magnification approximately x 30,000. The myelinated axon
anatomy is distorted here to show several nodes; in fact, An action potential at the node triggers flow of current
the distance between nodes is generally about 100 times to the next node, where the membrane regenerates the
as large as the nodes themselves. action potential.
Terms
absolute refractory period (p. 44) glucose (p. 37) oligodendrocyte (p. 35)
action potential (p. 42) graded potential (p. 47) polarization (p. 39)
active transport (p. 37) hyperpolarization (p. 42) presynaptic terminal (p. 33)
afferent axon (p. 33) interneuron (p. 34) propagation of the action potential
all-or-none law (p. 44) intrinsic neuron (p. 34) (p. 45)
astrocyte (p. 35) local anesthetic (p. 44) radial glia (p. 35)
axon (p. 33) local neuron (p. 47) refractory period (p. 44)
axon hillock (p. 45) membrane (p. 31) relative refractory period (p. 44)
blood-brain barrier (p. 36) microglia (p. 35) resting potential (p. 39)
cell body, or soma (p. 33) mitochondrion (pl.: mitochondria) ribosome (p. 32)
concentration gradient (p. 41) (p. 31) saltatory conduction (p. 46)
dendrite (p. 32) motor neuron (p. 32) Schwann cell (p. 35)
dendritic spine (p. 32) myelin (p. 46) selective permeability (p. 40)
depolarization (p. 42) myelin sheath (p. 33) sensory neuron (p. 32)
efferent axon (p. 33) myelinated axon (p. 46) sodium-potassium pump (p. 41)
electrical gradient (p. 39) neuron (p. 30) thiamine (vitamin B1) (p. 37)
endoplasmic reticulum (p. 32) node of Ranvier (p. 33) threshold of excitation (p. 42)
glia (p. 35) nucleus (p. 31) voltage-activated channel (p. 43)
I
Module 3.3
Drugs and Synapses f you had to communicate with someone without
Drug Mechanisms using sound, what would you do? Chances are, your
Common Drugs and Their Synaptic Effects first choice would be a visual code, such as written
In Closing: Drugs and Behavior words or sign language. Your second choice would
Summary probably be some sort of touch code or a system of elec-
Answers to Stop & Check Questions trical impulses. You might not even think of passing
Thought Question chemicals back and forth. Chemical communication
is, however, the primary method of communication for
Terms your neurons. Considering how well the human ner-
Suggestions for Further Reading vous system works, chemical communication is evi-
Websites to Explore dently more versatile than we might have guessed.
Exploring Biological Psychology CD Neurons communicate by transmitting chemicals at
ThomsonNOW specialized junctions called synapses, which are cen-
tral to all information processing in the brain.
51
Module 3.1
The Concept of the Synapse
52 Chapter 3 Synapses
comes relaxed. Let us consider each of these points and mission too weak to reach the threshold for an action
their implications. potential in the postsynaptic neuron, the cell that re-
ceives the message. (The neuron that delivers the syn-
Speed of a Reflex and Delayed aptic transmission is the presynaptic neuron.) Sher-
rington proposed that this subthreshold excitation
Transmission at the Synapse begins to decay shortly after it starts but can combine
When Sherrington pinched a dogs foot, the dog flexed with a second excitation that quickly follows it. A rapid
that leg after a very short but measurable delay. During succession of pinches produces a series of weak activa-
that delay, an impulse had to travel up an axon from the tions at the synapse, each adding its effect to what was
skin receptor to the spinal cord, and then an impulse left of the previous ones. If enough excitations occur
had to travel from the spinal cord back down the leg rapidly enough, they combine to exceed the threshold
to a muscle. Sherrington measured the total distance of the postsynaptic neuron.
that the impulse traveled from skin receptor to spinal Decades after Sherrington, John Eccles (1964) in-
cord to muscle and calculated the speed at which the serted microelectrodes into neurons to measure changes
impulse must have traveled to produce the response in the electrical potential across the membrane. He at-
within the measured delay. He found that the speed tached stimulating electrodes to axons of presynaptic
of conduction through the reflex arc varied but was neurons while recording from the postsynaptic neuron.
never more than about 15 meters per second (m/s). In For example, after he had briefly stimulated an axon,
contrast, previous research had measured action po- Eccles recorded a slight depolarization of the mem-
tential velocities along sensory or motor nerves at about brane of the postsynaptic cell (point 1 in Figure 3.3).
40 m/s. Even if the measurements were not exactly ac- Note that this partial depolarization is a graded
curate, Sherrington concluded that some process was potential. Unlike action potentials, which are always
slowing conduction through the reflex, and he inferred depolarizations, graded potentials may be either de-
that the delay must occur where one neuron commu- polarizations (excitatory) or hyperpolarizations (in-
nicates with another (Figure 3.2). This idea is critical, hibitory). A graded depolarization is known as an ex-
as it established the existence of synapses. Sherrington, citatory postsynaptic potential (EPSP). Like the action
in fact, introduced the term synapse. potentials discussed in Chapter 2, an EPSP occurs when
sodium ions enter the cell. However, in most cases,
transmission at a single synapse does not open enough
Temporal Summation sodium gates to reach the threshold. Unlike an action
Sherringtons work with reflex arcs suggested that re- potential, an EPSP decays over time and space; that
peated stimuli within a brief time have a cumulative is, its magnitude fades rapidly.
effect. He referred to this phenomenon as temporal When Eccles stimulated an axon twice in close
summation. A light pinch of the dogs foot did not succession, he recorded two consecutive EPSPs in the
evoke a reflex, but when Sherrington rapidly repeated postsynaptic cell. If the delay between EPSPs was short
the pinch several times, the leg flexed. Sherrington sur- enough, temporal summation occurred; that is, the
mised that a single pinch produced a synaptic trans- second EPSP added to what was left of the first one
(point 2 in Figure 3.3). The summation of two EPSPs
might or might not exceed the threshold of the post-
synaptic cell depending on the size of the EPSPs, the
A B time between them, and the threshold of the postsyn-
aptic cell. At point 3 in Figure 3.3, three consecutive
The speed of conduction along an axon is about 40 m/s. EPSPs combine to exceed the threshold and produce
an action potential.
C D E Spatial Summation
The speed of conduction through a reflex arc is slower and Sherringtons work with reflex arcs also suggested that
more variable, sometimes 15 m/s or less. Presumably, the synapses have the property of spatial summation: Sev-
delay occurs at the synapse.
eral synaptic inputs originating from separate locations
combine their effects on a neuron. Sherrington again
began with a pinch too weak to elicit a reflex. This time,
Figure 3.2 Sherringtons evidence for synaptic delay instead of pinching one point twice, he pinched two
An impulse traveling through a synapse in the spinal cord points at the same time. Although neither pinch alone
is slower than one traveling a similar distance along an elicited a response, the two together did. Sherrington
uninterrupted axon. concluded that pinching two points on the foot acti-
Threshold
50
55
60
65
70
vated two sensory neurons, whose axons converged both increase the depolarization of the postsynaptic
onto one neuron in the spinal cord. Excitation from cell and therefore increase the probability of an action
either axon excited that neuron but did not reach the potential (Figure 3.4).
threshold. Two excitations exceeded the threshold and You might guess that the synapses closer to the
elicited an action potential (point 4 in Figure 3.3). cell body of the postsynaptic cell might have a big-
Again, Eccles confirmed Sherringtons inference, ger effect than synapses on more distant parts of the
demonstrating that several axons were capable of pro- dendrites because the distant inputs might decay in
ducing EPSPs that summate their effects on a postsyn- strength as they travel toward the cell body. Surpris-
aptic cell. Note that temporal and spatial summation ingly, however, the synapses on remoter parts of the
dendrites produce larger EPSPs,
so their contribution to the cells
response approximately equals
Temporal summation
(several impulses from that of closer synapses (Magee &
one neuron over time) Cook, 2000).
Inhibitory Synapses
When Sherrington vigorously
pinched a dogs foot, the flexor
Action potential muscles of that leg contracted and
travels along axon so did the extensor muscles of
the other three legs (Figure 3.5).
At the same time, the dog relaxed
the extensor muscles of the stim-
ulated leg and the flexor muscles
of the other legs. Sherrington ex-
plained these results by assuming
certain connections in the spinal
Spatial summation (impulses from cord: A pinch on the foot sends a
several neurons at the same time) message along a sensory neuron
to an interneuron (an interme-
diate neuron) in the spinal cord,
Figure 3.4 Temporal and spatial summation which in turn excites the motor
54 Chapter 3 Synapses
Eccles and later researchers physiologically dem-
onstrated the inhibitory synapses that Sherrington
had inferred. At these synapses, input from the axon
hyperpolarizes the postsynaptic cell. That is, it in-
creases the negative charge within the cell, moving it
further from the threshold, and thus decreasing the
probability of an action potential (point 5 in Figure
3.3). This temporary hyperpolarization of a mem-
branecalled an inhibitory postsynaptic potential, or
IPSPresembles an EPSP in many ways. An IPSP oc-
curs when synaptic input selectively opens the gates
for potassium ions to leave the cell (carrying a posi-
tive charge with them) or for chloride ions to enter the
Flexor muscles contract Extensor muscles contract cell (carrying a negative charge). Inhibition is more
than just the absence of excitation; it is an active
brake that suppresses excitation.
Figure 3.5 Antagonistic muscles Today, we take the concept of inhibition for
Flexor muscles draw an extremity toward the trunk of the granted, but at Sherringtons time, the idea was con-
body, whereas extensor muscles move an extremity away troversial, as no one could imagine a mechanism to
from the body. accomplish it. Establishing the idea of inhibition was
critical not just for neuroscience but for psychology
as well. For example, Sigmund Freud, who developed
neurons connected to the flexor muscles of that leg his theories in the same era as Sherrington, proposed
(Figure 3.6). Sherrington surmised that the inter- that when a sexual energy was blocked from its de-
neuron also sends a message to block activity of mo- sired outlet, it would spill over into some other activ-
tor neurons connected to the extensor muscles in the ity. Today, we would simply talk about inhibiting an
same leg, as well as the flexor muscles of the three unwelcome activity; we see no need for the associated
other legs. energy to flow somewhere else.
Brain neuron
Excitatory
synapse Excitatory synapse
Skin
Intrinsic neuron
Inhibitory
synapse
Excitatory
Sensory synapse
neuron
Summary
Relationship Among EPSP, 1. The synapse is the point of communication between
IPSP, and Action Potential two neurons. Charles S. Sherringtons observations
of reflexes enabled him to infer the properties of
A given neuron may have anywhere from a few syn- synapses. (p. 52)
apses on its surface to thousands, some of them exci- 2. Because transmission through a reflex arc is slower
tatory and others inhibitory. Any number and combi- than transmission through an equivalent length of
nation of synapses may be active at any time, yielding axon, Sherrington concluded that some process at
both temporal and spatial summation. The probability the synapses delays transmission. (p. 53)
of an action potential depends on the ratio of EPSPs 3. Graded potentials (EPSPs and IPSPs) summate their
to IPSPs at a given moment. effects. The summation of graded potentials from
Most neurons have a spontaneous firing rate, a stimuli at different times is temporal summation.
periodic production of action potentials even without The summation of graded potentials from different
synaptic input. In such neurons, the EPSPs increase the locations is spatial summation. (p. 53)
frequency of action potentials above the spontaneous
4. A single stimulation at a synapse produces a brief
rate, whereas IPSPs decrease it below that rate. For ex-
graded potential in the postsynaptic cell. An exci-
ample, if the neurons spontaneous firing rate is 10 ac-
tatory graded potential (depolarizing) is an EPSP.
tion potentials per second, a stream of EPSPs might in-
An inhibitory graded potential (hyperpolarizing) is
crease the rate to 15 or more, whereas a preponderance
an IPSP. (pp. 53, 55)
of IPSPs might decrease it to 5 or fewer.
5. An EPSP occurs when gates open to allow sodium
to enter the neurons membrane; an IPSP occurs
when gates open to allow potassium to leave or
Module 3.1 chloride to enter. (pp. 53, 55)
In Closing: The Neuron as Decision Maker 6. The EPSPs on a neuron compete with the IPSPs;
When we learn the basics of any scientific field, we the balance between the two increases or de-
sometimes take them for granted, as if people always creases the neurons frequency of action potentials.
knew them. For example, we are taught that Earth and (p. 56)
the other planets rotate around the sun, and we dont
always pause to marvel at how Copernicus, hundreds
of years ago, drew that conclusion from some obser- Answers to
vations recorded before the invention of telescopes.
Sherringtons accomplishment is also amazing. Just STOP & CHECK
imagine measuring reflexive leg movements and in- Questions
ferring the existence of synapses and their major prop-
erties long before the invention of oscilloscopes or 1. Sherrington found that the velocity of conduction
electron microscopes. through a reflex arc was significantly slower than
56 Chapter 3 Synapses
you explain this finding? Remember that all action
the velocity of an action potential along an axon. potentialswhether produced by strong or weak
Therefore, some delay must occur at the junction stimulitravel at the same speed along a given axon.
between one neuron and the next. (p. 56) 2. A pinch on an animals right hind foot excites a
2. Temporal summation is the combined effect of sensory neuron that excites an interneuron that ex-
quickly repeated stimulation at a single synapse. cites the motor neurons to the flexor muscles of that
Spatial summation is the combined effect of sev- leg. The interneuron also inhibits the motor neu-
eral nearly simultaneous stimulations at several rons connected to the extensor muscles of the leg.
synapses onto one neuron. (p. 56) In addition, this interneuron sends impulses that
3. Sherrington found that a reflex that stimulates a reach the motor neuron connected to the extensor
flexor muscle sends a simultaneous message that muscles of the left hind leg. Would you expect the
inhibits nerves to the extensor muscles of the same interneuron to excite or inhibit that motor neuron?
limb. (p. 56) (Hint: The connections are adaptive. When an ani-
mal lifts one leg, it must put additional weight on
4. During an EPSP, sodium gates open. During an IPSP,
the other legs to maintain balance.)
potassium or chloride gates open. (p. 56)
3. Suppose neuron X has a synapse onto neuron Y,
which has a synapse onto Z. Presume that no other
Thought Questions neurons or synapses are present. An experimenter
finds that stimulating neuron X causes an action po-
1. When Sherrington measured the reaction time of tential in neuron Z after a short delay. However, she
a reflex (i.e., the delay between stimulus and re- determines that the synapse of X onto Y is inhibi-
sponse), he found that the response occurred faster tory. Explain how the stimulation of X might pro-
after a strong stimulus than after a weak one. Can duce excitation of Z.
58 Chapter 3 Synapses
was the exception. Finally, in the 1950s, researchers cleft, the space between the presynaptic and post-
established that chemical transmission is the predom- synaptic neurons.
inant type of communication throughout the nervous 4. The released molecules diffuse across the cleft, at-
system. That discovery revolutionized our understand- tach to receptors, and alter the activity of the post-
ing and led to research developing new drugs for psy- synaptic neuron.
chiatric uses (Carlsson, 2001). 5. The neurotransmitter molecules separate from their
receptors. Depending on the neurotransmitter, it
may be converted into inactive chemicals.
6. The neurotransmitter molecules may be taken back
The Sequence of Chemical into the presynaptic neuron for recycling or may
Events at a Synapse diffuse away. In some cases, empty vesicles are re-
turned to the cell body.
Understanding the chemical events at a synapse is fun- 7. Although the research is not yet conclusive, it is
damental to biological psychology. Here are the major likely that some postsynaptic cells send negative
events at a synapse: feedback messages to slow further release of neuro-
transmitter by the presynaptic cell.
1. The neuron synthesizes chemicals that serve as
neurotransmitters. It synthesizes the smaller neuro- Figure 3.8 summarizes these steps. Lets now consider
transmitters in the axon terminals and the larger each step in more detail.
ones (peptides) in the cell body.
2. The neuron transports the peptide neurotransmit-
ters to the axon terminals. (It doesnt have to trans-
Types of Neurotransmitters
port smaller neurotransmitters because they are At a synapse, one neuron releases chemicals that af-
formed in the terminals.) fect a second neuron. Those chemicals are known as
3. Action potentials travel down the axon. At the pre- neurotransmitters. Research has gradually identified
synaptic terminal, an action potential enables cal- more and more chemicals believed or suspected to be
cium to enter the cell. Calcium releases neurotrans- neurotransmitters; the current count is a hundred or
mitters from the terminals and into the synaptic more (Borodinsky et al., 2004). We shall consider many
Transporter
protein
3 Action
potential
8 causes
calcium to
enter, releasing
neurotransmitter
Synaptic
6 cleft 4 Neurotransmitter
binds to receptor
5 Separation from
receptors
7
Postsynaptic
8 Negative feedback sites
neuron
respond to retrograde
transmitter or to presynaptic
cells own transmitter.
60 Chapter 3 Synapses
Acetyl coenzyme A Phenylalanine Tryptophan Figure 3.9 Pathways in
(from metabolism) (from diet) (from diet) the synthesis of acetylcholine,
dopamine, norepinephrine,
+
epinephrine, and serotonin
Choline Arrows represent chemical reactions.
Tyrosine 5-hydroxytryptophan
(from metabolism or diet)
62 Chapter 3 Synapses
Images not available due to copyright restrictions
the second messenger communicates to areas within ing, the brain needs rapid, quickly changing infor-
the cell. The second messenger may open or close ion mation, the kind that ionotropic synapses bring.
channels in the membrane or alter the production of In contrast, hunger, thirst, fear, and anger constitute
proteins or activate a portion of a chromosome. Note the long-term changes in the probabilities of many behav-
contrast: An ionotropic synapse has effects localized iors. Metabotropic synapses are better suited for that
to one point on the membrane, whereas a metabotropic kind of function. Metabotropic synapses also mediate
synapse, by way of its second messenger, influences ac- at least some of the input for taste (Huang et al., 2005)
tivity in a larger area of the cell and over a longer time. and pain (Levine, Fields, & Basbaum, 1993), which are
Ionotropic and metabotropic synapses contribute slower and more enduring experiences than vision or
to different aspects of behavior. For vision and hear- hearing.
Membrane
2.
2 Receptor bends,
releasing G-protein
G-protein
3.
3 G-protein activates a
second messenger
such as cyclic AMP, which
Figure 3.12 Sequence of events at a metabolic alters a metabolic pathway,
synapse, using a second messenger within the turns on a gene in the nucleus,
postsynaptic neuron or opens or closes an ion channel
Hypothalamus Various releasing hormones Promote or inhibit release of various hormones by pituitary
Anterior pituitary Thyroid-stimulating hormone (TSH) Stimulates thyroid gland
Luteinizing hormone (LH) Increases production of progesterone (female), testosterone (male);
stimulates ovulation
Follicle-stimulating hormone (FSH) Increases production of estrogen and maturation of ovum (female)
and sperm production (male)
ACTH Increases secretion of steroid hormones by adrenal gland
Prolactin Increases milk production
Growth hormone (GH), also known Increases body growth, including the growth spurt during puberty
as somatotropin
Posterior pituitary Oxytocin Controls uterine contractions, milk release, certain aspects of
parental behavior, and sexual pleasure
Vasopressin (also known as Constricts blood vessels and raises blood pressure, decreases
antidiuretic hormone) urine volume
Pineal Melatonin Increases sleepiness, influences sleepwake cycle, also has role in
onset of puberty
Thyroid Thyroxine Increase metabolic rate, growth, and maturation
Triiodothyronine
Parathyroid Parathyroid hormone Increases blood calcium and decreases potassium
Adrenal cortex Aldosterone Reduces secretion of salts by the kidneys
Cortisol, corticosterone Stimulate liver to elevate blood sugar, increase metabolism of
proteins and fats
Adrenal medulla Epinephrine, norepinephrine Similar to effects of sympathetic nervous system
Pancreas Insulin Increases entry of glucose to cells and increases storage as fats
Glucagon Increases conversion of stored fats to blood glucose
Ovary Estrogens Promote female sexual characteristics
Progesterone Maintains pregnancy
Testis Androgens Promote sperm production, growth of pubic hair, and male sexual
characteristics
Liver Somatomedins Stimulate growth
Kidney Renin Converts a blood protein into angiotensin, which regulates blood
pressure and contributes to hypovolemic thirst
Thymus Thymosin (and others) Support immune responses
Fat cells Leptin Decreases appetite, increases activity, necessary for onset of puberty
64 Chapter 3 Synapses
Hormones Protein and peptide hormones attach to membrane re-
A hormone is a chemical that is secreted, in most ceptors where they activate a second messenger within
cases by a gland but also by other kinds of cells, and the cellexactly the same process as at a metabotropic
conveyed by the blood to other organs, whose activity synapse. In fact, many chemicalsincluding epineph-
it influences. A neurotransmitter is like a signal on a rine, norepinephrine, insulin, and oxytocinserve as
telephone line: It conveys a message directly and ex- both neurotransmitters and hormones.
clusively from the sender to the receiver. Hormones Just as circulating hormones modify brain activity,
function more like a radio station: They convey a mes- hormones secreted by the brain control the secretion
sage to any receiver that happens to be tuned in to the of many other hormones. The pituitary gland, attached
right station. Figure 3.13 presents the major endocrine to the hypothalamus (Figure 3.14), consists of two dis-
(hormone-producing) glands. Table 3.2 lists some im- tinct glands, the anterior pituitary and the posterior
portant hormones and their principal effects. pituitary, which release different sets of hormones (see
Hormones are particularly useful for coordinating Table 3.2). The posterior pituitary, composed of neural
long-lasting changes in multiple parts of the body. For tissue, can be considered an extension of the hypo-
example, birds that are preparing to migrate secrete thalamus. Neurons in the hypothalamus synthesize
hormones that change their eating and digestion to the hormones oxytocin and vasopressin (also known
store extra energy for a long journey. Among the vari- as antidiuretic hormone), which migrate down axons
ous types of hormones are protein hormones and pep- to the posterior pituitary, as shown in Figure 3.15.
tide hormones, composed of chains of amino acids. Later, the posterior pituitary releases these hormones
(Proteins are longer chains and peptides are shorter.) into the blood.
The anterior pituitary, composed of glandular
tissue, synthesizes six hormones, although the hypo-
Hypothalamus
Pineal gland
Pituitary gland
Parathyroid glands
Thyroid glands
Thymus
Liver
Optic chiasm Third ventricle
Adrenal gland
Hypothalamus
Kidney
Pancreas
Pituitary
stalk
Figure 3.13 Location of some major endocrine Figure 3.14 Location of the hypothalamus
glands and pituitary gland in the human brain
(Source: Starr & Taggart, 1989) (Source: Starr & Taggart, 1989)
Anterior pituitary
TSH
Thyroid gland
Thyroxine and
triiodothyronine
Excitatory effect
(Arterial flow) Inhibitory effect
Anterior pituitary Posterior pituitary Figure 3.16 Negative feedback in the control
of thyroid hormones
The hypothalamus secretes a releasing hormone that
stimulates the anterior pituitary to release TSH, which
stimulates the thyroid gland to release its hormones. Those
Vasopressin hormones in turn act on the hypothalamus to decrease its
and oxytocin secretion of the releasing hormone.
GH, ACTH,
TSH, FSH,LH,
and prolactin (Arterial flow)
tive feedback system. For example, when the level of
thyroid hormone is low, the hypothalamus releases
TSH-releasing hormone, which stimulates the ante-
Figure 3.15 Pituitary hormones
rior pituitary to release TSH, which in turn causes the
The hypothalamus produces vasopressin and oxytocin,
thyroid gland to secrete more thyroid hormones (Fig-
which travel to the posterior pituitary (really an extension
ure 3.16). For more information about hormones in
of the hypothalamus). The posterior pituitary releases those
general, visit this site: http://www.endo-society.org/
hormones in response to neural signals. The hypothalamus
also produces releasing hormones and inhibiting hormones,
which travel to the anterior pituitary, where they control
the release of six hormones synthesized there. STOP & CHECK
66 Chapter 3 Synapses
aptic neuron, which takes it up and reconnects it with ceived it, sending it again and again. To prevent clut-
acetate already in the cell to form acetylcholine again. tering your inbox, you might add a system that replied
Although this recycling process is highly efficient, it to any message with an automatic reply, Yes, I got your
takes time, and the presynaptic neuron does not re- message; dont send it again.
absorb every molecule it releases. A sufficiently rapid A couple of mechanisms in the nervous system may
series of action potentials at any synapse can deplete serve that function. First, many presynaptic terminals
the neurotransmitter faster than the presynaptic cell re- have receptors sensitive to the same transmitter they
plenishes it, thus slowing or halting transmission (Liu release. Theoretically, these receptors may act as auto-
& Tsien, 1995). receptorsreceptors that detect the amount of trans-
In the absence of acetylcholinesterase, acetylcho- mitter released and inhibit further synthesis and re-
line remains and continues stimulating its receptor. lease after it reaches a certain level. That is, they could
Drugs that block acetylcholinesterase can be helpful for provide negative feedback. However, the evidence on
people with diseases that impair acetylcholine trans- this point is mixed. For example, we should expect
mission, such as myasthenia gravis. autoreceptors to be more strongly stimulated by repet-
Serotonin and the catecholamines (dopamine, nor- itive release of the transmitter than by single release,
epinephrine, and epinephrine) do not break down but the data so far have been inconsistent on this point
into inactive fragments at the postsynaptic membrane (Kalsner, 2001; Starke, 2001).
but simply detach from the receptor. The presynaptic Second, some postsynaptic neurons respond to
neuron takes up most of these neurotransmitter mol- stimulation by releasing special chemicals that travel
ecules intact and reuses them. This process, called re- back to the presynaptic terminal, where they inhibit
uptake, occurs through special membrane proteins further release of transmitter. Nitric oxide is appar-
called transporters. Many of the familiar antidepres- ently one such transmitter. Two others are anandamide
sant drugs, such as fluoxetine (Prozac), block reuptake and 2-AG (sn-2 arachidonylglycerol), both of which
and thereby prolong the effects of the neurotransmitter bind to the same receptors as marijuana extracts. We
on its receptor. (Chapter 15 discusses antidepressants shall discuss them further in the next module, when
in more detail.) we consider drugs more thoroughly. Here, the point is
Some of the serotonin and catecholamine mol- that neurons apparently have ways to inhibit them-
ecules, either before or after reuptake, are converted into selves from excessive release of transmitter.
inactive chemicals that cannot stimulate the receptor.
The enzymes that convert catecholamine transmitters
into inactive chemicals are COMT (catechol-o-methyl-
Synapses and Personality
transferase) and MAO (monoamine oxidase), which The nervous system uses, according to current esti-
affects serotonin as well as catecholamines. We return mates, about 100 neurotransmitters in various areas.
to MAO in the discussion of antidepressant drugs. Many of the neurotransmitters have multiple kinds of
The peptide neurotransmitters (or neuromodula- receptors, each with different properties. Why such
tors) are neither inactivated nor reabsorbed. They sim- an abundance of mechanisms? Presumably, each kind
ply diffuse away. of synapse has a specific function in behavior or in the
development of the nervous system. If each kind of
synapse has its own specific role in behavior, then any-
one who has more or less than the normal amount of
STOP & CHECK some neurotransmitter or receptor (for genetic or other
reasons) should have some altered behavioral tenden-
9. What happens to acetylcholine molecules after they cies. That is, theoretically, variations in synapses should
stimulate a postsynaptic receptor? have something to do with variations in personality.
In the 1990s, researchers studied variations in
10. What happens to serotonin and catecholamine
dopamine receptors and found that people with one
molecules after they stimulate a postsynaptic
form of the D2 receptor (one of five types of dopamine
receptor?
receptors) were more likely than others to develop se-
Check your answers on page 68. vere alcoholism. Later research suggested that this gene
is not specific to alcoholism but instead increases the
probability of a variety of pleasure-seeking behaviors,
Negative Feedback including alcohol consumption, other recreational drug
use, overeating, and habitual gambling (K. Blum, Cull,
from the Postsynaptic Cell Braverman, & Comings, 1996).
Suppose someone had a habit of sending you an e-mail Similarly, researchers studying variations in the D4
message and then, worried that you might not have re- receptor found that people with an alternative form of
68 Chapter 3 Synapses
10. Most serotonin and catecholamine molecules are to know whether B is just an extension of axon A
reabsorbed by the presynaptic terminal. Some or whether A formed an excitatory synapse on some
of their molecules are broken down into inactive neuron in the ganglion, whose axon is axon B. How
chemicals, which then float away in the blood. could an experimenter determine the answer? You
(p. 67) should be able to think of more than one good
method. Presume that the anatomy within the gan-
glion is so complex that you cannot simply trace
the course of an axon through it.
Thought Questions
2. Transmission of visual and auditory information
1. Suppose that axon A enters a ganglion (a cluster of relies largely on ionotropic synapses. Why is iono-
neurons) and axon B leaves on the other side. An tropic better than metabotropic for these purposes?
experimenter who stimulates A can shortly there- For what purposes might metabotropic synapses
after record an impulse traveling down B. We want be better?
cause leakage
from vesicles
m
DA
e
DA C
Amphetamine DA PA e)
increases release DA DO ctiv
a
(in
DA Cocaine blocks
release reuptake reuptake. So do
Typical methylphenidate
ine receptor
antipsychotic Dopam and tricyclic
drug, such antidepressants,
but less strongly.
as haloperidol,
blocks receptor Postsynaptic neuron
70 Chapter 3 Synapses
hormones are the same as those of other animal species
(Cravchik & Goldman, 2000). So if a plant evolves some
chemical to attract wasps to its flowers, to deter cater- STOP & CHECK
pillars from eating its leaves, or whatever, that chemi-
cal is likely to affect humans also.
1. Is a drug with high affinity and low efficacy an agonist
Another part of the answer is that many of our
or an antagonist?
neurotransmitters are also found in plants. For exam-
ple, plants not only have glutamate, but they also have Check your answers on page 77.
ionotropic glutamate receptors chemically similar to
our own (Lam et al., 1998). Exactly what these recep-
tors do in plants is uncertain; perhaps they serve for
some type of communication within a plant, even
though the plant has no nervous system. Evidently, a Common Drugs and
small number of chemicals are so well suited to con- Their Synaptic Effects
veying information that evolution has had no reason
to replace them. We categorize drugs based on their predominant ac-
tion. For example, amphetamine and cocaine are stim-
ulants; opiates are narcotics; LSD is a hallucinogen.
Drug Mechanisms Despite their differences, however, nearly all abused
drugs directly or indirectly stimulate the release of
Drugs can either facilitate or inhibit transmission at dopamine, especially in the nucleus accumbens, a
synapses. A drug that blocks the effects of a neuro- small subcortical area rich in dopamine receptors
transmitter is called an antagonist; a drug that mimics (Figure 3.18). Dopamine is in most cases an inhibitory
or increases the effects is called an agonist. (The term transmitter; however, in the nucleus accumbens, sus-
agonist is derived from a Greek word meaning con- tained bursts of dopamine inhibit cells that release the
testant. We derive our term agony from the same root. inhibitory transmitter GABA. Thus, by inhibiting re-
An antagonist is an antiagonist, or member of the op- lease of an inhibitor, dopamine has the net effect of ex-
posing team.) A drug that is a mixed agonistantagonist citing the nucleus accumbens (Hjelmstad, 2004).
is an agonist for some behavioral effects of the neuro-
transmitter and an antagonist for others, or an agonist
at some doses and an antagonist at others.
Stimulant Drugs
Drugs influence synaptic activity in many ways. Stimulant drugs increase excitement, alertness, and
As in Figure 3.17, which illustrates a dopamine syn- motor activity, while elevating mood. They decrease
apse, a drug can increase or decrease the synthesis of fatigue. Whereas nearly all drugs elevate dopamine
the neurotransmitter, cause it to leak from its vesicles, activity in the nucleus accumbens in some way, stim-
increase its release, decrease its reuptake, block its ulant drugs do so directly, especially at dopamine re-
breakdown into inactive chemicals, or directly stimu- ceptor types D2, D3, and D4 (R. A. Harris, Brodie, &
late or block the postsynaptic receptors. Dunwiddie, 1992; Wise & Bozarth, 1987).
Investigators say that a drug has an affinity for a Because dopamine is in many cases an inhibitory
particular type of receptor if it binds to that receptor, transmitter, drugs that increase activity at dopamine
fitting somewhat like a lock and key. Drugs vary in their synapses decrease the activity in much of the brain.
affinities from strong to weak. The efficacy of a drug Figure 3.19 shows the results of a PET scan, which
is its tendency to activate the receptor. So, for example, measures relative amounts of activity in various brain
a drug that binds tightly to a receptor but fails to stim- areas (London et al., 1990). (PET scans are a method of
ulate it has a high affinity but a low efficacy. visualizing brain anatomy, as described in Chapter 4.)
You may have noticed that the effectiveness and How, you might wonder, could drugs that decrease
side effects of tranquilizers, antidepressants, and other brain activity lead to behavioral arousal? One hypoth-
drugs vary from one person to another. Why? One rea- esis is that high dopamine activity mostly decreases
son is that most drugs affect several kinds of recep- background noise in the brain and therefore increases
tors. People vary in their abundance of each kind of the signal-to-noise ratio (Mattay et al., 1996; Willson,
receptor. For example, one person might have a rela- Wilman, Bell, Asghar, & Silverstone, 2004).
tively large number of dopamine type D4 receptors and Amphetamine stimulates dopamine synapses by
relatively few D1 or D2 receptors, whereas someone increasing the release of dopamine from the presyn-
else has more D1, fewer D4, and so forth. Therefore, a aptic terminal. The presynaptic terminal ordinarily re-
drug with an affinity for dopamine receptors will af- absorbs released dopamine through a protein called the
fect different kinds of dopamine receptors in different dopamine transporter. Amphetamine reverses the trans-
people (Cravchik & Goldman, 2000). porter, causing the cell to release dopamine instead of
72 Chapter 3 Synapses
common with cocaine. Larger amounts of methyl- CTL MA
phenidate produce effects resembling cocaines. >0.5 >0.5
Many people wonder whether prolonged use of 0.3 0.3
methylphenidate in childhood makes people more
likely to abuse drugs later. According to a review of the 0.1 0.1
Average Average
few studies conducted on this issue, children who take gray matter gray matter
methylphenidate are less likely than others to abuse
(a) (b)
drugs during adolescence or early adulthood (Wilens,
Faraone, Biederman, & Gunawardene, 2003). Of course, Figure 3.20 Brain thinning in MDMA users
the researchers could not randomly assign children to The figure on the left is the average brain scan for 21 adults
methylphenidate and control conditions. However, who had never been substance abusers; the figure on the
experiments with rats point to the same conclusion. right is the average of 22 adults who had used MDMA
For example, experimenters gave young rats moderate frequently for years. Red represents the thickest area of
doses of methylphenidate and then, months later, gave brain cells, followed by yellow, green, blue, and purple. Note
them cocaine and tested their preference for the room the apparent thinning in certain areas for the MDMA users.
where they received cocaine versus an adjoining room (Source: From Structural abnormalities in the brains of human
where they had not received it. Compared to other rats, subjects who use methamphetamine, by P. M. Thompson et al.,
those with early exposure to methylphenidate showed Journal of Neuroscience, 24. Copyright 2004 by the Society for
a lower preference for the stimuli associated with co- Neuroscience. Reprinted with permission.
caine and in some cases an avoidance of them (Ander-
sen, Arvanitogiannis, Pliakas, LeBlanc, & Carlezon,
2002; Carlezon, Mague, & Andersen, 2003; Mague, An-
dersen, & Carlezon, 2005). Although these studies con-
tradict the worry that early methylphenidate treatment
STOP & CHECK
might lead to later drug abuse, prolonged use of methyl-
phenidate leads to other long-term disadvantages, in- 2. How does amphetamine influence dopamine synapses?
cluding increased fear responses and a possibly in-
3. How does cocaine influence dopamine synapses?
creased risk of depression (Bolaos, Barrot, Berton,
Wallace-Black, & Nestler, 2003; Carlezon et al., 2003). 4. Why is methylphenidate generally less disruptive to
The drug methylenedioxymethamphetamine behavior than cocaine is despite the drugs similar
(MDMA, or ecstasy) is a stimulant at low doses, in- mechanisms?
creasing the release of dopamine. At higher doses (com- 5. Does cocaine increase or decrease overall brain activity?
parable to the doses people use recreationally), it also Check your answers on page 77.
increases serotonin release, producing hallucinogenic
effects as well. In monkey studies, MDMA not only
stimulates axons that release serotonin but also de-
stroys them (McCann, Lowe, & Ricaurte, 1997).
Nicotine
Research on humans is difficult because research- Nicotine, a compound present in tobacco, has long
ers cannot randomly assign people to MDMA and con- been known to stimulate one type of acetylcholine re-
trol groups. Many studies have found that repeated ceptor, conveniently known as the nicotinic receptor,
MDMA users, in comparison to nonusers, have more which is found both in the central nervous system and
anxiety, depression, sleep problems, memory deficits, at the nerve-muscle junction of skeletal muscles. Nico-
attention problems, and impulsiveness, even a year or tinic receptors are abundant on neurons that release
two after quitting (Montoya, Sorrentino, Lukas, & Price, dopamine in the nucleus accumbens, so nicotine in-
2002). Like laboratory animals, humans exposed to creases dopamine release there (Levin & Rose, 1995;
MDMA have long-term deficits in serotonin synthesis Pontieri, Tanda, Orzi, & DiChiara, 1996). In fact, nico-
and release (Parrott, 2001). One study found that re- tine increases dopamine release in mostly the same
peated MDMA users have thinner cell layers in certain cells of the nucleus accumbens that cocaine does (Pich
brain areas, as shown in Figure 3.20 (P. M. Thompson et al., 1997).
et al., 2004). We do not know, of course, how many of One consequence of repeated exposure to nicotine,
the users had behavioral problems or thin cell layers as demonstrated in rat studies, is that after the end of
in the brain prior to using MDMA. (Maybe people who nicotine use, the nucleus accumbens cells responsible
already have anxiety, memory problems, and so forth for reinforcement become less responsive than usual
are more likely than others to use MDMA.) Also, some (Epping-Jordan, Watkins, Koob, & Markou, 1998). That
MDMA users abuse other drugs as well. Still, the evi- is, many events, not just nicotine itself, become less
dence suggests serious risks associated with this drug. reinforcing than they used to be.
74 Chapter 3 Synapses
anandamide (from the Sanskrit word ananda, mean- The report that time passes more slowly under
ing bliss) (Calignano, LaRana, Giuffrida, & Piomelli, marijuanas influences is harder to explain, but what-
1998; DiMarzo et al., 1994) and the more abundant ever the cause, we can demonstrate it in rats as well:
sn-2 arachidonylglycerol, abbreviated 2-AG (Stella, Consider a rat that has learned to press a lever for food
Schweitzer, & Piomelli, 1997). The cannabinoid recep- on a fixed-interval schedule, where just the first press
tors are peculiar in being located on the presynaptic of any 30-second period produces food. With practice,
neuron, not the postsynaptic one. When glutamate de- a rat learns to wait after each press before it starts press-
polarizes the postsynaptic neuron, the postsynaptic ing again. Under the influence of marijuana, rats press
neuron releases anandamide or 2-AG, which travels sooner after each reinforcer. For example, instead of
back to the presynaptic neuron, temporarily decreas- waiting 20 seconds, a rat might wait only 10 or 15. Ev-
ing transmitter release (Kreitzer & Regehr, 2001; R. I. idently, the 10 or 15 seconds felt like 20 seconds; time
Wilson & Nicoll, 2002). Cannabinoids also diffuse to was passing more slowly (Han & Robinson, 2001).
inhibit the release of GABA from other nearby neurons
(Galante & Diana, 2004). That is, cannabinoids put the
brakes on the release of both glutamate, which is exci-
Hallucinogenic Drugs
tatory, and GABA, which is inhibitory (Kreitzer & Drugs that distort perception are called hallucinogenic
Regehr, 2001; Ohno-Shosaku, Maejima, & Kano, 2001; drugs. Many hallucinogenic drugs, such as lysergic
R. I. Wilson & Nicoll, 2001). In some cases, the result acid diethylamide (LSD), chemically resemble sero-
is decreased release of transmitter in a fairly wide area tonin (Figure 3.21) and stimulate serotonin type 2A
(Kreitzer, Carter, & Regehr, 2002). (5-HT2A) receptors at inappropriate times or for longer
The functions of cannabinoids are partly known, than usual durations. You could compare this effect to
partly unknown. After a rapid burst of glutamate re- a key that almost fits a lock, so that when you get it in,
lease, cannabinoids feed back to retard further gluta- its hard to get it out. (The analogy is not perfect, but it
mate release (the negative feedback discussed on page conveys the general idea.)
67). Excessive glutamate stimulation can be harmful, O === C N(C2H5)2
even fatal, to a postsynaptic cell, so halting it is clearly
helpful (Marsicano et al., 2003). The value of blocking NCH3
GABA is less obvious but may be related to the forma-
tion of memories (Chevaleyre & Castillo, 2004; Gerde-
man, Ronesi, & Lovinger, 2002).
HO CH2CH2NH2
Marijuana stimulates cannabinoid receptors at
times when they would not ordinarily receive stimula- N N
tion. In effect, they give glutamate-releasing neurons the H H
message, I already received your message before the Serotonin LSD
message is sent, shutting down the normal flow of in-
formation. Why are these effects pleasant or habit form- Figure 3.21 Resemblance of the neurotransmitter
ing? Actually, of course, not all of the effects of mari- serotonin to LSD, a hallucinogenic drug
juana are pleasant. But remember that virtually all
abused drugs increase the release of dopamine in the Note that we understand the chemistry better than
nucleus accumbens. Cannabinoids do so indirectly. One the psychology. LSD exerts its effects at 5-HT2A recep-
place in which they inhibit GABA release is the ventral tors, but why do effects at those receptors produce hal-
tegmental area of the midbrain, a major source of axons lucinations? Table 3.3 summarizes the effects of some
that release dopamine in the nucleus accumbens. When commonly abused drugs.
cannabinoids inhibit GABA there, the result is less in-
hibition (therefore increased activity) of the neurons
that release dopamine in the nucleus accumbens (Cheer,
Wassu, Heien, Phillips, & Wightman, 2004). STOP & CHECK
Researchers have tried to explain some of mari-
juanas other behavioral effects. Cannabinoids relieve
8. What are the effects of cannabinoids on neurons?
nausea by inhibiting serotonin type 3 synapses (5-HT3),
which are known to be important for nausea (Fan, 9. If incoming serotonin axons were destroyed, LSD
1995). Cannabinoid receptors are abundant in areas of would still have its normal effects on the postsynaptic
the hypothalamus that influence feeding, and mice cell. However, if incoming dopamine and norepineph-
lacking these receptors show decreased appetite under rine axons were destroyed, amphetamine and cocaine
some circumstances (DiMarzo et al., 2001). Presum- would have no effects. Explain the difference.
ably, excess cannabinoid activity would produce extra Check your answers on page 77.
appetite.
Amphetamine Excitement, alertness, elevated mood, Increases release of dopamine and several other transmitters
decreased fatigue
Cocaine Excitement, alertness, elevated mood, Blocks reuptake of dopamine and several other transmitters
decreased fatigue
Methylphenidate Increased concentration Blocks reuptake of dopamine and others, but gradually
(Ritalin)
MDMA (ecstasy) Low dose: stimulant Releases dopamine
Higher dose: sensory distortions Releases serotonin, damages axons containing serotonin
Nicotine Mostly stimulant effects Stimulates nicotinic-type acetylcholine receptor, which
(among other effects) increases dopamine release in nucleus
accumbens
Opiates Relaxation, withdrawal, decreased pain Stimulates endorphin (e.g., heroin, morphine) receptors
Cannabinoids Altered sensory experiences, decreased Excites negative-feedback receptors on presynaptic cells;
(marijuana) pain and nausea, increased appetite those receptors ordinarily respond to anandamide and 2AG
Hallucinogens Distorted sensations Stimulates serotonin type 2A receptors (5-HT2A)
(e.g., LSD)
76 Chapter 3 Synapses
7. Opiates stimulate endorphin synapses, which in-
hibit GABA synapses on certain cells that release
Answers to
dopamine. By inhibiting an inhibitor, opiates in-
STOP & CHECK crease the release of dopamine. (p. 74)
Questions 8. Cannabinoids released by the postsynaptic neuron
attach to receptors on presynaptic neurons, where
1. Such a drug is therefore an antagonist because, by they inhibit further release of glutamate as well as
occupying the receptor, it prevents the normal ef- GABA. (p. 75)
fects of the transmitter. (p. 71) 9. Amphetamine acts by releasing norepinephrine
2. Amphetamine causes the dopamine transporter to and dopamine from the presynaptic neurons. If
release dopamine instead of reabsorbing it. (p. 73) those neurons are damaged, amphetamine is in-
3. Cocaine interferes with reuptake of released dopa- effective. In contrast, LSD directly stimulates the re-
mine. (p. 73) ceptor on the postsynaptic membrane. (p. 75)
4. The effects of a methylphenidate pill develop and
decline in the brain much more slowly than do those
of cocaine. (p. 73) Thought Question
5. Cocaine decreases total activity in the brain because 1. People who take methylphenidate (Ritalin) for con-
it stimulates activity of dopamine, which is an in- trol of attention-deficit disorder often report that,
hibitory transmitter in many cases. (p. 73) although the drug increases their arousal for a while,
6. Nicotine excites acetylcholine receptors on neurons they feel a decrease in alertness and arousal a few
that release dopamine and thereby increases dopa- hours later. Explain.
mine release. (p. 74)
Terms
acetylcholine (p. 60) exocytosis (p. 61) posterior pituitary (p. 65)
acetylcholinesterase (p. 66) G-protein (p. 62) postsynaptic neuron (p. 53)
affinity (p. 71) hallucinogenic drugs (p. 75) presynaptic neuron (p. 53)
2-AG (p. 75) hormone (p. 65) protein hormone (p. 65)
agonist (p. 71) inhibitory postsynaptic potential purines (p. 60)
amino acids (p. 60) (IPSP) (p. 55) reflex (p. 52)
amphetamine (p. 71) ionotropic effect (p. 62) reflex arc (p. 52)
anandamide (p. 75) MAO (p. 67) releasing hormone (p. 66)
antagonist (p. 71) metabotropic effect (p. 62) reuptake (p. 67)
anterior pituitary (p. 65) methylphenidate (p. 72) second messenger (p. 63)
autoreceptors (p. 67) monoamines (p. 60) spatial summation (p. 53)
cannabinoids (p. 74) neuromodulator (p. 64) spontaneous firing rate (p. 56)
catecholamine (p. 60) neurotransmitter (p. 59) stimulant drugs (p. 71)
cocaine (p. 72) nicotine (p. 73) synapse (p. 52)
COMT (p. 67) nitric oxide (p. 60) temporal summation (p. 53)
9-tetrahydrocannabinol (9-THC) nucleus accumbens (p. 71) transporter (p. 67)
(p. 74) opiate drugs (p. 74) vasopressin (p. 65)
efficacy (p. 71) oxytocin (p. 65) vesicle (p. 61)
endocrine glands (p. 65) peptide (p. 60)
excitatory postsynaptic potential peptide hormone (p. 65)
(EPSP) (p. 53) pituitary gland (p. 65)
78 Chapter Ending
AChE Inhibitors
Exploring Biological
Psychology CD
Postsynaptic Potentials (animation)
Release of Neurotransmitter (animation)
Cholinergic Synapse (animation) This animation shows how a drug can block reuptake of a
Release of ACh (animation) neurotransmitter. You can find this animation in Chapter 3 on
AChE Inactivates ACh (animation) the CD.
Chapter Ending 79
4
Anatomy of the
Nervous System
T
The Parietal Lobe
The Temporal Lobe rying to learn neuroanatomy (the anatomy of the
The Frontal Lobe nervous system) from a book is like trying to learn
How Do the Parts Work Together? geography from a road map. A map can tell you that
In Closing: Functions of the Cerebral Cortex Mystic, Georgia, is about 40 km north of Enigma, Geor-
Summary gia. Similarly, a book can tell you that the habenula is
Answers to Stop & Check Questions about 4.6 mm from the interpeduncular nucleus in a
Thought Question rats brain (slightly farther in a human brain). But these
little gems of information will seem both mysterious
Module 4.3 and enigmatic unless you are concerned with that part
Research Methods of Georgia or that area of the brain.
Correlating Brain Anatomy with Behavior This chapter does not provide a detailed road map
Recording Brain Activity of the nervous system. It is more like a world globe,
Effects of Brain Damage describing the large, basic structures (analogous to the
Effects of Brain Stimulation continents) and some distinctive features of each.
Brain and Intelligence The first module introduces key neuroanatomical
In Closing: Research Methods and Their Limits terms and outlines overall structures of the nervous
Summary system. In the second module, we concentrate on the
Answers to Stop & Check Questions structures and functions of the cerebral cortex, the
Thought Question largest part of the mammalian central nervous system.
The third module deals with the main methods that
Terms researchers use to discover the behavioral functions
Suggestions for Further Reading of different brain areas.
Websites to Explore Be prepared: This chapter contains a huge number
Exploring Biological Psychology CD of new terms. You should not expect to memorize all
ThomsonNOW of them at once, and it will pay to review this chapter
repeatedly.
Opposite: New methods allow researchers to examine
living brains. Source: Peter Beck/CORBIS
81
Module 4.1
Structure of the Vertebrate
Nervous System
Thalamus
Hypothalamus
Pituitary gland
Pons
Medulla
Cerebellum
Peripheral Nervous System
Somatic (blue): Controls voluntary muscles and conveys sensory
information to the central nervous system
Autonomic (red): Controls involuntary muscles
Sympathetic: Expends energy
Parasympathetic: Conserves energy
Dorsal Toward the back, away from the Proximal Located close (approximate) to the
ventral (stomach) side. The top of point of origin or attachment
the brain is considered dorsal
Distal Located more distant from the point
because it has that position in four-
of origin or attachment
legged animals.
Ipsilateral On the same side of the body (e.g.,
Ventral Toward the stomach, away from the
two parts on the left or two on the
dorsal (back) side
right)
Anterior Toward the front end
Contralateral On the opposite side of the body
Posterior Toward the rear end (one on the left and one on the right)
Superior Above another part Coronal plane A plane that shows brain structures
(or frontal plane) as seen from the front
Inferior Below another part
Sagittal plane A plane that shows brain structures
Lateral Toward the side, away from the
as seen from the side
midline
Horizontal plane A plane that shows brain structures
Medial Toward the midline, away from the
(or transverse plane) as seen from above
side
or
r
Horizontal
teri
vous system is a complex three-dimensional structure, rio
te
plane
Pos
we need more terms to describe it. As Figure 4.2 and
An
The Autonomic
Nervous System
The autonomic nervous system consists of neurons
that receive information from and send commands to
the heart, intestines, and other organs. It is comprised
of two parts: the sympathetic and parasympathetic
nervous systems (Figure 4.6). The sympathetic nervous
E X T E N S I O N S A N D A P P L I C AT I O N S
Goose Bumps
Each segment of the spinal cord sends sensory in-
formation to the brain and receives motor commands Erection of the hairs, known as goose bumps or
from the brain. All that information passes through goose flesh, occurs when we are cold. What does it
tracts of axons in the spinal cord. If the spinal cord is have to do with the fight-or-flight functions associated
cut at a given segment, the brain loses sensation from with the sympathetic nervous system? Part of the an-
that segment and all segments below it; the brain also swer is that we also get goose bumps when we are
loses motor control over all parts of the body served frightened. You have heard the expression, I was so
by that segment and the lower ones. frightened my hairs stood on end. You may also have
seen a frightened cat erect its fur. Human body hairs
are so short that erecting them accomplishes nothing,
but a cat with erect fur looks bigger and potentially
frightening. A frightened porcupine erects its quills,
which are just modified hairs (Richter & Langworthy,
1933). The behavior that makes the quills so useful,
their erection in response to fear, evidently evolved be-
fore the quills themselves did.
Manfred Kage/Peter Arnold, Inc.
Cervical
nerves
(8 pairs)
Lungs
Stomach
Celiac
ganglion Pancreas
Liver Thoracic
nerves
(12 pairs)
Adrenal
Muscles that erect hairs gland
Sweat Kidney
gland
Small intestine
Lumbar
Large intestine nerves
(5 pairs)
(Most ganglia
near spinal cord) Bladder
Sacral
Pelvic nerves
nerve (5 pairs)
Uterus Coccygeal
nerve
Sympathetic outflow Parasympathetic outflow (1 pair)
Genitals
Figure 4.6 The sympathetic nervous system (red lines) and parasympathetic
nervous system (blue lines)
Note that the adrenal glands and hair erector muscles receive sympathetic input only.
(Source: Adapted from Biology: The Unity and Diversity, 5th Edition, by C. Starr and R. Taggart, p. 340.
Copyright 1989 Wadsworth.)
thetic systems act in opposition to one another, both in a chain near the spinal cord. Rather, long pregangli-
are constantly active to varying degrees, and many stim- onic axons extend from the spinal cord to parasympa-
uli arouse parts of both systems. thetic ganglia close to each internal organ; shorter post-
The parasympathetic nervous system is also known ganglionic fibers then extend from the parasympathetic
as the craniosacral system because it consists of the ganglia into the organs themselves. Because the para-
cranial nerves and nerves from the sacral spinal cord sympathetic ganglia are not linked to one another, they
(see Figure 4.6). Unlike the ganglia in the sympathetic act somewhat more independently than the sympa-
system, the parasympathetic ganglia are not arranged thetic ganglia do. Parasympathetic activity decreases
I. Olfactory Smell
II. Optic Vision
III. Oculomotor Control of eye movements, pupil constriction
IV. Trochlear Control of eye movements
V. Trigeminal Skin sensations from most of the face; control of jaw muscles for chewing and swallowing
VI. Abducens Control of eye movements
VII. Facial Taste from the anterior two-thirds of the tongue; control of facial expressions, crying, salivation, and
dilation of the heads blood vessels
VIII. Statoacoustic Hearing, equilibrium
IX. Glossopharyngeal Taste and other sensations from throat and posterior third of the tongue; control of swallowing,
salivation, throat movements during speech
X. Vagus Sensations from neck and thorax; control of throat, esophagus, and larynx; parasympathetic nerves
to stomach, intestines, and other organs
XI. Accessory Control of neck and shoulder movements
XII. Hypoglossal Control of muscles of the tongue
Cranial nerves III, IV, and VI are coded in red to highlight their similarity: control of eye movements.
Cranial nerves VII, IX, and XII are coded in green to highlight their similarity: taste and control of
tongue and throat movements. Cranial nerve VII has other important functions as well. Nerve X (not
highlighted) also contributes to throat movements, although it is primarily known for other functions.
Cranial nerve V
Pons
ter 8), and many older textbooks describe the cerebel- means a covering, such as a rug on the floor. The teg-
lum as important for balance and coordination. True, mentum covers several other midbrain structures, al-
people with cerebellar damage are clumsy and lose though it is covered by the tectum.) The tegmentum in-
their balance, but the functions of the cerebellum ex- cludes the nuclei for the third and fourth cranial nerves,
tend far beyond balance and coordination. People with parts of the reticular formation, and extensions of the
damage to the cerebellum have trouble shifting their pathways between the forebrain and the spinal cord or
attention back and forth between auditory and visual hindbrain. Another midbrain structure is the substan-
stimuli (Courchesne et al., 1994). They have much dif- tia nigra, which gives rise to the dopamine-containing
ficulty with timing, including sensory timing. For ex- pathway that deteriorates in Parkinsons disease (see
ample, they are poor at judging whether one rhythm is Chapter 8).
faster than another.
The Forebrain
The Midbrain
The forebrain is the most anterior and most promi-
As the name implies, the midbrain is in the middle of nent part of the mammalian brain. It consists of two
the brain, although in adult mammals it is dwarfed cerebral hemispheres, one on the left side and one on
and surrounded by the forebrain. In birds, reptiles, am- the right (Figure 4.11). Each hemisphere is organized
phibians, and fish, the midbrain is a larger, more promi- to receive sensory information, mostly from the contra-
nent structure. The roof of the midbrain is called the lateral (opposite) side of the body, and to control mus-
tectum. (Tectum is the Latin word for roof; the same cles, mostly on the contralateral side, by way of axons
root shows up in the geological term plate tectonics.) to the spinal cord and the cranial nerve nuclei.
The two swellings on each side of the tectum are the The outer portion is the cerebral cortex. (Cerebrum
superior colliculus and the inferior colliculus (see Fig- is a Latin word meaning brain; cortex is a Latin word
ures 4.8 and 4.10); both are part of important routes for meaning bark or shell.) Under the cerebral cortex
sensory information. are other structures, including the thalamus, which is
Under the tectum lies the tegmentum, the inter- the main source of input to the cerebral cortex. A set
mediate level of the midbrain. (In Latin, tegmentum of structures known as the basal ganglia plays a major
Frontal lobe
Thalamus
Corpus callosum
Midbrain
Pituitary gland
Pons
Cerebellum
Medulla
Spinal cord
Central canal of
spinal cord
Anterior
Frontal lobe of
cerebral cortex
Frontal lobe Corpus callosum
Lateral ventricles
Occipital lobe (posterior parts)
Posterior
Figure 4.11 Dorsal view of the brain surface and a horizontal section through the brain
role in certain aspects of movement. A number of other important for motivations and emotions, such as eat-
interlinked structures, known as the limbic system, ing, drinking, sexual activity, anxiety, and aggression.
form a border (or limbus, the Latin word for border) The structures of the limbic system are the olfactory
around the brainstem. These structures are particularly bulb, hypothalamus, hippocampus, amygdala, and cin-
Thalamus
Hypothalamus
Mamillary body
Hippocampus
Amygdala
Olfactory bulb
Frontal
lobe of Longitudinal
cerebral fissure
cortex
Olfactory
bulbs
Cerebral
cortex Corpus Temporal
callosum lobe of
Lateral cerebral Optic
Dorsal ventricles cortex nerves
Ventral Basal
ganglia
Medulla
Temporal
Left lobes Cerebellum Spinal cord
Right
Anterior commissure
gulate gyrus of the cerebral cortex. Figure 4.12 shows cortex. In later chapters, we return to each of these
the positions of these structures in three-dimensional areas as they become relevant.
perspective. Figures 4.10 and 4.13 show sagittal (from
the side) and coronal (from the front) sections through
the human brain. Figure 4.13 also includes a view of the
Thalamus
ventral surface of the brain. The thalamus and hypothalamus together form the di-
In describing the forebrain, we begin with the sub- encephalon, a section distinct from the rest of the fore-
cortical areas; the next module focuses on the cerebral brain, which is known as the telencephalon. The thal-
Frontal cortex
Optic tract
Dorsomedial nucleus
amus is a structure in the center of the forebrain. The contains a number of distinct nuclei, which we exam-
term is derived from a Greek word meaning ante- ine in Chapters 10 and 11. Partly through nerves and
room, inner chamber, or bridal bed. It resembles partly through hypothalamic hormones, the hypothal-
two avocados joined side by side, one in the left hemi- amus conveys messages to the pituitary gland, altering
sphere and one in the right. Most sensory information its release of hormones. Damage to any hypothalamic
goes first to the thalamus, which then processes it and nucleus leads to abnormalities in motivated behav-
sends the output to the cerebral cortex. The one clear iors, such as feeding, drinking, temperature regulation,
exception to this rule is olfactory information, which sexual behavior, fighting, or activity level. Because of
progresses from the olfactory receptors to the olfactory these important behavioral effects, the rather small hy-
bulbs and from the bulbs directly to the cerebral cor- pothalamus attracts a great deal of research attention.
tex without passing through the thalamus.
Many nuclei of the thalamus receive their primary Pituitary Gland
input from one of the sensory systems, such as vision,
and then transmit the information to a single area of the The pituitary gland is an endocrine (hormone-
cerebral cortex, as in Figure 4.14. The cerebral cortex producing) gland attached to the base of the hypothal-
then sends information back to the thalamus, prolong- amus by a stalk that contains neurons, blood vessels,
ing and magnifying certain kinds of input at the expense and connective tissue (see Figure 4.10). In response to
of others, apparently serving to focus attention on par- messages from the hypothalamus, the pituitary syn-
ticular stimuli (Komura et al., 2001). thesizes and releases hormones into the bloodstream,
which carries them to other organs.
Hypothalamus
The hypothalamus is a small area near the base of the
Basal Ganglia
brain just ventral to the thalamus (see Figures 4.10 The basal ganglia, a group of subcortical structures
and 4.12). It has widespread connections with the rest lateral to the thalamus, include three major structures:
of the forebrain and the midbrain. The hypothalamus the caudate nucleus, the putamen, and the globus pal-
Caudate
nucleus
Thalamus
lidus (Figure 4.15). Some au-
thorities include several other
structures as well. The basal Globus Putamen
ganglia have been conserved pallidus (lateral)
through evolution, and the ba- (medial)
sic organization is about the Amygdala
same in mammals as in amphibians (Marin,
Smeets, & Gonzlez, 1998).
The basal ganglia have multiple subdivisions,
each of which exchanges information with a different
part of the cerebral cortex. The connections are most
abundant with the frontal areas of the cortex, which
are responsible for planning sequences of be-
havior and for certain aspects of memory
and emotional expression (Graybiel,
Aosaki, Flaherty, & Kimura, 1994). In
conditions such as Parkinsons dis-
ease and Huntingtons disease, in
which the basal ganglia deterio-
rate, the most prominent symp-
tom is impaired movement, but
people also show depression, defi-
cits of memory and reasoning, and
attentional disorders.
Basal Forebrain
Several structures lie on the dorsal surface of
the forebrain, including the nucleus basalis, which
receives input from the hypothalamus and basal
ganglia and sends axons that release acetylcho- Nucleus basalis
line to widespread areas in the cerebral cortex (Fig-
ure 4.16). We might regard the nucleus basalis as
an intermediary between the emotional arousal of the
hypothalamus and the information processing of the
cerebral cortex. The nucleus basalis is a key part of
the brains system for arousal, wakefulness, and atten-
tion, as we consider in Chapter 9. Patients with Parkin-
sons disease and Alzheimers disease have impair- Figure 4.16 The basal forebrain
ments of attention and intellect because of inactivity The nucleus basalis and other structures in this area send
or deterioration of their nucleus basalis. axons throughout the cortex, increasing its arousal and
wakefulness through release of the neurotransmitter
Hippocampus acetylcholine. (Source: Adapted from Cholinergic Systems
in Mammalian Brain and Spinal Cord, by N. J. Woolf, Progress
The hippocampus (from a Latin word meaning sea in Neurobiology, 37, pp. 475524, 1991)
horse, a shape suggested by the hippocampus) is a
large structure between the thalamus and the cerebral
cortex, mostly toward the posterior of the forebrain, cussion is that the hippocampus is critical for storing
as shown in Figure 4.12. We consider the hippocam- certain kinds of memories but not all. A debate contin-
pus in more detail in Chapter 12; the gist of that dis- ues about how best to describe the class of memories
Lateral ventricles
Third
ventricle
Lateral
ventricles
Thalamus
Cerebral Fourth
aqueduct ventricle
Central canal Anterior Posterior
of spinal cord
(a) (b)
(visual, auditory, and somatosensory) but absent from illustrates the idea of columns, although in nature they
the motor cortex. are not so straight. The cells within a given column
The cells of the cortex are also organized into col- have similar properties to one another. For example,
umns of cells perpendicular to the laminae. Figure 4.22 if one cell in a column responds to touch on the palm
of the left hand, then the other cells in that column do
too. If one cell responds to a horizontal pattern of light
at a particular location in the retina, then the other cells
in the column respond to the same pattern in nearly the
same location.
We now turn to some of the specific parts of the
cortex. Researchers distinguish 50 or more areas of the
cerebral cortex based on differences in the thickness of
May no
Surface of cortex
Prefrontal
cortex Visual
Auditory
Olfactory
bulb
Olfaction
Occipital
lobe Audition
(vision) Vision
Temporal lobe
(hearing, advanced
visual processing)
Somesthesis
Movement
(a) (b)
Precentral gyrus
(primary motor
cortex)
Leg
Hip
Trun
Knee
Hip
Nec
Trun lder
Hea
Sho
Arm w
Elb arm
Arm ow
k
k
Fo
Elb
d
Postcentral gyrus
k
u
o
re
Ha
W
(primary Fi
Th nge
nd
ris
Ha
somatosensory Fi
Ey um rs t
nd
cortex) Th nge
No e b N um rs
se Broeck b
Fac Toes Eye w
e
Fac
Lips e
Genitals
Teeth Lips
Gums
Jaw Jaw
ue
Tong Tongu
e
r nx
y l
Pha ina ing
om llow
Swa
Dr. Dana Copeland
d
ab
ra-
Int
Figure 4.25
Species differences
in prefrontal cortex
Note that the prefrontal
cortex (blue area)
constitutes a larger
proportion of the human
brain than of these other
species. (Source: After The Squirrel monkey Cat Rhesus monkey
Prefrontal Cortex by J. M.
Fuster, 1989, Raven Press.
Reprinted by permission.)
Wrist
Head STOP & CHECK
Finger Eyes
UpperFacial
LowerMuscles
Tongue
Platy FRONTAL
mus sma
cle L ASSOCIA- 4. What is meant by the binding problem and what
arynx TION
CENTRE are two hypotheses to explain it?
VISUAL
AREA
Check your answer on page 104.
OCCIPITO-TEMPORAL
ASSOCIATION CENTRE
AUDITORY AREA
SOMSTHETIC AREA
PARTIETAL ASSOCIA-
Module 4.2
TION CENTER
In Closing: Functions of the Cerebral Cortex
The human cerebral cortex is so large that we easily
slip into thinking of it as the brain, with all of the rest
VISUAL
of the brain almost trivial. In fact, only mammals have
AREA
a true cerebral cortex, and many mammals have only a
FRONTAL
ASSOCIATION small one. So subcortical areas by themselves can pro-
CENTRE
duce very complex behaviors, and a cerebral cortex
by itself cannot do anything at all (because it would
OCCIPITO-TEMPORAL
ASSOCIATION CENTRE not be connected to any sense organs or muscles).
OLFACTORY AREA
What, then, is the function of the cerebral cortex?
The primary function seems to be one of elaborating
Figure 4.27 An old, somewhat misleading view sensory material. Even fish, which have no cerebral
of the cortex cortex, can see, hear, and so forth, but they do not rec-
Note the designation association centre in this illustration ognize and remember all the complex aspects of sen-
of the cortex from an old introductory psychology textbook sory stimuli that mammals do. In a television adver-
(Hunter, 1923). Todays researchers are more likely to regard tisement, one company says that it doesnt make any
those areas as additional sensory areas. products, but it makes lots of products better. The same
could be said for the cerebral cortex.
ce
ur
so
y
X-ra
ctor
ete
yd
Dan McCoy/Rainbow
-ra
(a) (b)
second to another.
Figure 4.32 shows an MEG record of brain re-
sponses to a brief tone heard in the right ear. The dia-
gram represents a human head as viewed from above,
with the nose at the top (Hari, 1994). Researchers using
MEG can identify the times at which various brain areas
respond and thereby trace a wave of brain activity from
its point of origin to all the other areas that process it Figure 4.33 A PET scanner
(Salmelin, Hari, Lounasmaa, & Sams, 1994). A person engages in a cognitive task while attached to this
Another method, positron-emission tomography apparatus that records which areas of the brain become
(PET), provides a high-resolution image of activity in more active and by how much.
Nasal
bone
Computerized axial tomography (CAT) Maps brain areas, but requires exposure to x-rays
Magnetic resonance imaging (MRI) Maps brain areas in detail, using magnetic fields
Record from electrodes in brain Invasive; used with laboratory animals, seldom humans
Electroencephalograph (EEG) Records from scalp; measures changes by ms, with but low resolution of location of
the signal
Evoked potentials Similar to EEG but in response to stimuli
Magnetoencephalograph (MEG) Similar to EEG but measures magnetic fields
Positron emission tomography (PET) Measures changes over both time and location but requires exposing brain to radiation
Functional magnetic resonance Measures changes over about 1 second, identifies location within 12 mm, no use of
imaging (fMRI) radiation
Study victims of stroke etc. Used with humans; each person has different damage
Lesion Controlled damage in laboratory animals
Ablation Removal of a brain area
Gene-knockout Effects wherever that gene is active (e.g., a receptor)
Transcranial magnetic stimulation Intense application temporarily inactivates a brain area
Stimulating electrodes Invasive; used with laboratory animals, seldom with humans
Transcranial magnetic stimulation Brief, mild application activates underlying brain area
1500 1400
1400 1300
1300 1200
1200 1100
1100 1000
1000 900
1000 1100 1200 1300 1400 1500 1600 1700 600 800 1000 1200 1400 1600
Cerebral volume of twin 1 Cerebral volume of twin 1
same in men and women. Because the surface is lined of behavior? In the rest of this text, we concentrate on
with cells (gray matter), the sexes are nearly equal in these types of questions.
gray matter, whereas men have more white matter
(Allen, Damasio, Grabowski, Bruss, & Zhang, 2003). An
MRI study found a .34 correlation between teenagers
IQ scores and the gray matter volumes of their brains
STOP & CHECK
(Frangou, Chitins, & Williams, 2004).
So a tentative conclusion is that, other things being 7. Why do recent studies show a stronger relationship
equal, more gray matter is associated with better per- between brain size and IQ than older studies did?
formance on intellectual tests. However, lets go back
8. What evidence indicated that the genes that control
to species comparisons: The difference between human
human brain size also influence IQ?
brains and those of chimpanzees and gorillas is more
a matter of increased white matter in humans than in- Check your answers on page 116.
creased gray matter (Schoenemann, Sheehan, & Glotzer,
2005). So the species difference seems to imply that
white matter (i.e., connectivity within the brain) is im-
portant for intelligence. At this point, we have far bet-
ter data than before about the brain and intelligence, Module 4.3
but the overall picture is still confusing. In Closing: Research Methods
On the other hand, how important is this ques-
tion, really? It has been a matter of curiosity for many
and Their Limits
people for a long time, but it has no great theoretical im- Descriptions of the history of science sometimes high-
portance or practical applications. Relating total brain light a single study that conclusively established one
to total intelligence is like relating the geographical theory or another. Such events are rare. Far more often,
area of a country to the size of its population: Sure, the researchers gradually accumulate evidence that points
correlation is positive, but it overlooks a host of more in a particular direction, until eventually that view
interesting variables. Progress in both psychology and becomes dominant. Even in those rare cases when a
neuroscience depends on making finer-grained distinc- single study appears to have been decisive, research-
tions. How do the anatomy, chemistry, and other fea- ers often identify it as decisive only in retrospect, after
tures of each part of the brain relate to specific aspects several additional studies have confirmed the finding.
Terms
ablation (p. 109) gamma waves (p. 102) phrenology (p. 105)
anterior (p. 83) ganglion (pl.: ganglia) (p. 84) pituitary gland (p. 92)
anterior commissure (p. 96) gene-knockout approach (p. 110) pons (p. 88)
autonomic nervous system (p. 83) gray matter (p. 84) positron-emission tomography
basal ganglia (p. 92) gyrus (pl.: gyri) (p. 84) (PET) (p. 108)
Bell-Magendie law (p. 84) hindbrain (p. 87) postcentral gyrus (p. 98)
binding problem (p. 102) hippocampus (p. 93) posterior (p. 83)
brainstem (p. 87) horizontal plane (p. 83) precentral gyrus (p. 100)
central canal (p. 94) hypothalamus (p. 92) prefrontal cortex (p. 100)
central nervous system (CNS) inferior (p. 83) prefrontal lobotomy (p. 100)
(p. 82) inferior colliculus (p. 89) primates (p. 96)
central sulcus (p. 98) ipsilateral (p. 83) proximal (p. 83)
cerebellum (p. 88) Klver-Bucy syndrome (p. 100) raphe system (p. 88)
cerebral cortex (p. 96) lamina (pl.: laminae) (pp. 84, 96) reticular formation (p. 88)
cerebrospinal fluid (CSF) (p. 94) lateral (p. 83) sagittal plane (p. 83)
column (pp. 84, 97) lesion (p. 109) sham lesion (p. 109)
computerized axial tomography limbic system (p. 90) somatic nervous system (p. 83)
(CT or CAT scan) (p. 105) spinal cord (p. 84)
magnetic resonance imaging (MRI)
contralateral (p. 83) (p. 106) stereotaxic instrument (p. 109)
coronal plane (p. 83) magnetoencephalograph (MEG) substantia nigra (p. 89)
corpus callosum (p. 96) (p. 108) sulcus (pl.: sulci) (p. 84)
cranial nerve (p. 87) medial (p. 83) superior (p. 83)
delayed-response task (p. 101) medulla (p. 87) superior colliculus (p. 89)
distal (p. 83) meninges (p. 94) sympathetic nervous system
dorsal (p. 83) midbrain (p. 89) (p. 85)
dorsal root ganglion (p. 84) nerve (p. 84) tectum (p. 89)
electroencephalograph (EEG) neuroanatomy (p. 81) tegmentum (p. 89)
(p. 107) nucleus (p. 84) temporal lobe (p. 98)
evoked potentials or evoked nucleus basalis (p. 93) thalamus (p. 91)
responses (p. 107)
occipital lobe (p. 98) tract (p. 84)
fissure (p. 84)
parasympathetic nervous system transcranial magnetic stimulation
forebrain (p. 89) (p. 85) (p. 110)
frontal lobe (p. 100) parietal lobe (p. 98) ventral (p. 83)
functional magnetic resonance peripheral nervous system (PNS) ventricle (p. 94)
imaging (fMRI) (p. 109) (p. 82) white matter (p. 84)
The CD includes this virtual reality brain that you can rotate to
various positions and dissect with a click.
Terms
Suggestions for Further Reading
Websites to Explore
Exploring Biological Psychology CD
S ome assembly required. Have you ever bought
a package with those ominous words? Some-
times all you have to do is attach a few parts. But some-
ThomsonNOW times you face page after page of incomprehensible in-
structions. I remember putting together my daughters
bicycle and wondering how something that looked so
simple could be so complicated.
The human nervous system requires an enormous
amount of assembly, and the instructions are different
from those for a bicycle. Instead of, Put this piece here
and that piece there, the instructions are, Put these
axons here and those dendrites there, and then wait
to see what happens. Keep the connections that work
the best and discard the others. Continue periodically
making new connections and keeping only the most
successful ones.
Therefore, we say that the brains anatomy is plas-
tic; it is constantly changing, within limits. The brain
changes rapidly in early development and continues
changing throughout life.
Opposite: An enormous amount of brain development has
already occurred by the time a person is 1 year old.
Source: Dr. Dana Copeland
121
Module 5.1
Development of the Brain
Neural
groove
velopment of neurons: proliferation, migration, differ- long the cell proliferation lasts in days and the num-
entiation, myelination, and synaptogenesis. ber of new neurons produced per day. For example, the
Proliferation is the production of new cells. Early main differences between human brains and chim-
in development, the cells lining the ventricles of the panzee brains are due to the fact that neurons continue
brain divide. Some cells remain where they are (as stem proliferating longer in humans (Rakic, 1998; Vrba,
cells), continuing to divide and redivide. Others be- 1998). Evidently, a small genetic change can produce a
come primitive neurons and glia that begin migrating major difference in outcome.
to other locations. The developmental process is about After cells have differentiated as neurons or glia,
the same in all vertebrates, varying in two factors: how they migrate (move) toward their eventual destinations
in the brain. Different kinds of
cells originate in different loca-
Midbrain Midbrain Hindbrain tions at different times, and each
must migrate substantial dis-
Forebrain tances, following specific chem-
Hindbrain ical paths, to reach its final des-
tination (Marn & Rubenstein,
Forebrain Cranial
nerves 2001). Some move radially from
the inside of the brain to the out-
Spinal cord side; others move tangentially
along the surface of the brain; and
3 weeks 7 weeks some move tangentially and then
radially (Nadarajah & Parnave-
Midbrain Forebrain las, 2002). Chemicals in families
known as immunoglobulins and
Forebrain
chemokines help to guide neu-
ron migration. A deficit in these
chemicals can lead to impaired
Hindbrain
migration, decreased brain size,
Cerebellum decreased axon growth, and men-
Midbrain Medulla tal retardation (Berger-Sweeney
(hidden) & Hohmann, 1997; Crossin &
Krushel, 2000; Tran & Miller,
2003). On the other extreme, ex-
cesses of immunoglobulins have
11 weeks At birth
been linked to some cases of
Figure 5.3 Human brain at four stages of development schizophrenia (Crossin & Krushel,
Chemical processes develop the brain to an amazing degree even before the start of 2000; Poltorak et al., 1997). The
any experience with the world. Detailed changes in development continue to occur brain has many kinds of immuno-
throughout life. globulins and chemokines, pre-
leg moved in synchrony with its neighbor because each No. The current estimate is that humans have
axon had found exactly the correct muscle. only about 30,000 genes totalfar too few to mark
Since the time of Weisss work, most of the research each of the brains many billions of neurons. Never-
on axon growth has dealt with how sensory axons find theless, nearly all axons grow to almost exactly their
their way to the correct targets in the brain. (The issues correct targets (Kozloski, Hamzei-Sichani, & Yuste,
are the same as those for axons finding their way to 2001). Without an extensive system of individual
muscles.) In one study, Roger Sperry, a former student markers, neurons still manage to find their way.
of Weiss, cut the optic nerves of some newts. (See photo But how?
and quote on the pages inside the back cover.) The dam- A growing axon follows a path of cell-surface
aged optic nerve grew back and connected with the molecules, attracted by some chemicals and repelled
tectum, which is the main visual area of fish, amphib- by others, in a process that steers the axon in the cor-
ians, reptiles, and birds (Figure 5.5). Sperry found that rect direction (Yu & Bargmann, 2001). Some axons
when the new synapses formed, the newt regained nor- follow a trail based on one attractive chemical until
mal vision. they reach an intermediate location where they be-
Then Sperry (1943) repeated the experiment, but come insensitive to that chemical and start following
this time, after he cut the optic nerve, he rotated the eye a different attractant (Shirasaki, Katsumata, & Mura-
by 180 degrees. When the axons grew back to the tec- kami, 1998; H. Wang & Tessier-Lavigne, 1999). Even-
tum, which targets would they contact? Sperry found tually, axons sort themselves over the surface of their
that the axons from what had originally been the dor- target area by following a gradient of chemicals. For
sal portion of the retina (which was now ventral) grew example, one chemical in the amphibian tectum is
back to the area responsible for vision in the dorsal a protein known as TOPDV (TOP for topography; DV
retina. Axons from what had once been the ventral for dorsoventral). This protein is 30 times more con-
retina (now dorsal) also grew back to their original tar- centrated in the axons of the dorsal retina than of the
gets. The newt now saw the world upside down and ventral retina and 10 times more concentrated in the
backward, responding to stimuli in the sky as if they ventral tectum than in the dorsal tectum. As axons
were on the ground and to stimuli on the left as if they from the retina grow toward the tectum, the retinal
were on the right (Figure 5.6). Each axon regenerated axons with the greatest concentration of TOPDV con-
to the area of the tectum where it had originally been, nect to the tectal cells with the highest concentration
presumably by following a chemical trail. of that chemical; the axons with the lowest concen-
tration connect to the tectal cells with the lowest con-
Chemical Gradients centration. A similar gradient of another protein
The next question was: How specific is the axons aim? aligns the axons along the anteriorposterior axis
Must an axon from the retina find the tectal cell with (J. R. Sanes, 1993) (Figure 5.7). (By analogy, you could
exactly the right chemical marker on its surface, like a think of men lining up from tallest to shortest, pair-
key finding the right lock? Does the body have to syn- ing up with women who lined up from tallest to short-
thesize a separate chemical marker for each of the bil- est, so the tallest man dated the tallest woman and so
lions of axons in the nervous system? forth.)
Ventral Ventral
Retina Tectum
Old ventral Dorsal
&
Number of motor neurons
STOP CHECK
175,000
Richard Coss
declines in old age but much less in peo- prise. What was more surprising was that
ple who remain physically active (Col- PET and fMRI scans indicated substantial
combe et al., 2003). activity in the occipital cortex of the blind
(a) (b)
People with extensive academic people while they performed these tasks
education also tend to have longer and Figure 5.11 Effect (Burton et al., 2002; Sadato et al., 1996,
more widely branched dendrites than of a stimulating 1998). Evidently, touch information had
people with less formal education (B. Ja- environment on invaded this cortical area, which is or-
cobs, Schall, & Scheibel, 1993). It is neuronal branching dinarily devoted to vision alone.
tempting to assume that the increased (a) A jewel fish reared in To double-check this conclusion, re-
learning led to brain changes, but we isolation develops neurons searchers asked blind and sighted people
cannot draw cause-and-effect conclu- with fewer branches. (b) A to perform the same kind of task during
sions from correlational studies. Per- fish reared with others has temporary inactivation of the occipital cor-
haps people who already had wider den- more neuronal branches. tex. Remember from Chapter 4 (p. 110)
drites succeeded more in school and that intense magnetic stimulation on the
therefore stayed longer. Or of course, scalp can temporarily inactivate neurons
both explanations could be correct. beneath the magnet. Applying this procedure to the
occipital cortex of blind people interferes with their
ability to identify Braille symbols or to notice the dif-
Effects of Special Experiences ference between one tactile stimulus and another. The
The detailed structure and function of the brain can be same procedure does not impair any aspect of touch
modified by experiences. For example, neurons become for sighted people. In short, blind people use the oc-
more responsive and more finely tuned to stimuli that cipital cortex to help identify what they feel; sighted
have been important or meaningful in the past (e.g., people do not (L. G. Cohen et al., 1997). One conclu-
Fritz, Shamma, Elhilali, & Klein, 2003; L. I. Zhang, Bao, sion is that touch information invades the occipital
& Merzenich, 2001). How much plasticity might occur cortex of blind people. A broader conclusion is that
after experiences that are far different from the average? brain areas modify their function based on the kinds
of stimuli they receive.
Brain Adaptations in On the average, blind people also outperform
People Blind Since Infancy sighted people on many verbal skills. (If you cant see,
One way to ask this question is to consider what hap- you pay more attention to what you hear, including
pens to the brain if one sensory system is nearly or en- words.) One example is the task, When you hear the
tirely absent. Recall the experiment on ferrets, in which name of an object (e.g., apple), say as quickly as pos-
axons of the visual system, unable to contact their nor- sible the name of an appropriate action for that object
mal targets, attached instead to the brain areas usually (e.g., eat ). Again, performing this task activates parts
devoted to hearing and managed to convert them into of the occipital cortex in blind people but not in
more-or-less satisfactory visual areas (p. 124). Might sighted people. Furthermore, the amount of activity in
anything similar happen in the brains of people born the occipital cortex (for blind people) correlates with
deaf or blind? their performance on the task (Amedi, Raz, Pianka,
You will hear many people say that blind people Malack, & Zohary, 2003). Also, inactivating the occip-
become better than usual at touch and hearing, or that ital cortex by intense transcranial magnetic stimula-
deaf people become especially acute at touch and vi- tion interferes with verbal performance by blind peo-
132 Chapter 5 Development and Plasticity of the Brain
ple but not by sighted people (Amedi, Floel, Knecht, The most strongly affected areas related to hand con-
Zohary, & Cohen, 2004). So the occipital cortex of blind trol and vision (presumably because of its importance
people serves certain verbal functions as well as touch. for reading music). A related study on stringed instru-
As the occipital cortex increases its response to ment players found that a larger than normal section
touch, sound, and verbal stimuli, does it decrease its of the postcentral gyrus was devoted to representing
response to visual stimuli? Applying brief transcra- the fingers of the left hand, which they use to control
nial magnetic stimulation (enough to stimulate, not the strings (Elbert, Pantev, Wienbruch, Rockstroh, &
enough to inactivate) over the occipital cortex causes Taub, 1995). The area devoted to the left fingers was
sighted people to report seeing flashes of light. When largest in those who began their music practice early
the same procedure is applied to people who com- (and therefore also continued for more years).
pletely lost their sight because of eye injuries more These results imply that practicing a skill reorga-
than 10 years ago, most report seeing nothing or see- nizes the brain, within limits, to maximize performance
ing flashes only rarely after stimulation in certain lo- of that skill. Part of the mechanism is that sustained at-
cations (Gothe et al., 2002). Note that for this experi- tention to anything releases dopamine, and dopamine
ment it is essential to test people who once had normal acts on the cortex to expand the response to stimuli
vision and then lost it because we can ask them whether active during the dopamine release (Bao, Chan, & Mer-
they see anything in response to the magnetic stimula- zenich, 2001).
tion. Someone blind since birth presumably would not
understand the question. When Brain Reorganization
Goes Too Far
Effects of Prolonged Practice Ordinarily, the expanded cortical representation of
Cognitive psychologists have found that extensive personally important information is beneficial. How-
practice of a skill, such as playing chess or working ever, in extreme cases, the reorganization creates
crossword puzzles, makes someone ever more adept problems. As mentioned, when people play string in-
at that skill but not necessarily at anything else (Erics- struments many hours a day for years, the representa-
son & Charness, 1994). Presumably, developing ex- tion of the left hand increases in the somatosensory
pertise changes the brain in ways that improve the cortex. Similar processes occur in people who play
abilities required for a particular skill. In a few cases, piano and other instruments.
researchers have identified some of
the relevant brain changes.
Precentral and postcentral gyri
One study used magnetoenceph- (Body sensations and motor control, including fingers)
alography (MEG; see p. 108) to re-
cord responses of the auditory cortex
to pure tones. The responses in pro-
fessional musicians were about twice
as strong as those for nonmusicians.
Then an examination of their brains,
using MRI, found that one area of the
temporal cortex in the right hemi-
sphere was about 30% larger in the
professional musicians (Schneider
et al., 2002). Of course, we do not
know whether their brains were
equal before they started practicing;
perhaps those with certain predis-
positions are more likely than oth-
ers to develop a passion for music.
Another study used a type of
MRI scan to compare the entire
Inferior visual cortex Left inferior frontal gyrus
brains of professional keyboard
(Vision, such as reading music)
players, amateur keyboard players,
and nonmusicians. Several areas Figure 5.12 Brain correlates of extensive music practice
showed that gray matter was thicker Areas marked in red showed thicker gray matter among professional keyboard
in the professionals than in the am- players than in amateurs and thicker gray matter among amateurs than in
ateurs, and thicker in the amateurs nonmusicians. Areas marked in yellow showed even stronger differences in that
than the nonmusicians, including same direction. (Source: From Brain structures differ between musicians and non-
the structures highlighted in Fig- musicians, by C. Gaser & G. Schlaug, Journal of Neuroscience, 23. Copyright 2003 by the
ure 5.12 (Gaser & Schlaug, 2003). Society for Neuroscience. Reprinted with permission.)
Leg
Trun k
Hip
cortex)
Nec d
Hea
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Elb rm
Fo
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Postcentral gyrus
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(primary Fi
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somatosensory Thu ger
cortex) E ye mb s
No
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Fac e Toes
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Lips
Teeth Genitals
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Jaw 34
2
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a r y nx l
na
Dr. Dana Copeland
Ph mi
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Int
cortex)
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Arm w
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Postcentral gyrus
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(primary Fi
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Int
cortex)
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Arm w
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Fo nd
Postcentral gyrus
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(primary Fi
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Teeth Genitals
5
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Dr. Dana Copeland
P h
d om
ab
ra-
Int
a stroke, and a bullet wound victim. For a good col- damaged neurons (Kim, Park, Hong, & Koh, 1999). To
lection of information about stroke, visit this website: get significant benefit, a patient should receive tPA
http://www.stroke.org/ within 3 hours after a stroke, although slight benefits
In ischemia, neurons are deprived of blood and are possible up to 6 hours after the stroke. Unfortu-
therefore lose much of their oxygen and glucose sup- nately, by the time a patients family gets the patient to
plies. In hemorrhage, they are flooded with blood and the hospital, and then waits through procedures in the
therefore with excess oxygen, calcium, and other prod- emergency room, the delay is usually too long (Stahl,
ucts. Both ischemia and hemorrhage lead to many of Furie, Gleason, & Gazelle, 2003).
the same problems, including edema (the accumula- Even when it is too late for tPA to save the cells in
tion of fluid), which increases pressure on the brain the immediate vicinity of the ischemia or hemorrhage,
and increases the probability of additional strokes (Un- hope remains for cells in the penumbra (Latin for al-
terberg, Stover, Kress, & Kiening, 2004). Both ischemia most shadow), the region that surrounds the imme-
and hemorrhage also impair the sodium-potassium diate damage (Hsu, Sik, Gallyas, Horvth, & Buzski,
pump, leading to an accumulation of sodium inside 1994; Jonas, 1995) (Figure 5.14). One idea is to pre-
neurons. The combination of edema and excess sodium vent overstimulation by blocking glutamate synapses
provokes excess release of the transmitter glutamate or preventing positive ions from entering neurons.
(Rossi, Oshima, & Attwell, 2000), which overstimulates However, most such methods have produced disap-
neurons: Sodium and other ions enter the neurons in pointing results (Lee, Zipfel, & Choi, 1999). A some-
excessive amounts, faster than the sodium-potassium what promising new drug opens potassium channels
pump can remove them. The excess positive ions block (Gribkoff et al., 2001). As calcium or other positive ions
metabolism in the mitochondria and kill the neurons enter a neuron, potassium exits through these open
(Stout, Raphael, Kanterewicz, Klann, & Reynolds, channels, reducing overstimulation.
1998). As neurons die, glia cells proliferate, removing One possible reason blocking overstimulation has
waste products and dead neurons. not worked better is that stroke-damaged neurons also
If someone had a stroke and you called a hospital, can die from understimulation (Colbourne, Suther-
what advice would you probably get? As recently as the land, & Auer, 1999; Conti, Raghupathi, Trojanowski, &
1980s, the staff would have been in no great hurry to McIntosh, 1998). It is possible that a stroke reactivates
see the patient because they had little to offer anyway. the mechanisms of apoptosis. According to research
Today, it is possible to reduce the effects of an ischemic with laboratory animals, injections of neurotrophins
stroke if physicians act quickly. A drug called tissue and other drugs that block apoptosis can improve re-
plasminogen activator (tPA) breaks up blood clots covery from a stroke (Barinaga, 1996; Choi-Lundberg
(Barinaga, 1996) and has a mixture of other effects on et al., 1997; Levivier, Przedborski, Bencsics, & Kang,
erated, leaving vacant synaptic sites at several levels also feels a sensation in the phantom hand. It is possi-
of the CNS. Axons representing the face sprouted into ble to map out which part of the face stimulates sen-
those sites in the spinal cord, brainstem, and thalamus sation in which part of the phantom hand, as shown in
(Florence & Kaas, 1995; E. G. Jones & Pons, 1998). (Or Figure 5.18 (Aglioti, Smania, Atzei, & Berlucchi, 1997).
perhaps axons from the face had already innervated The relationship between phantom sensations and
those sites but were overwhelmed by the normal input. brain reorganization enables us to understand some
After removal of the normal input, the weaker synapses otherwise puzzling observations. Note in Figure 4.24
became stronger.) Also, lateral connections sprouted
from the face-sensitive cortical areas into the previously
hand-sensitive areas of the cortex, according to results
from histochemistry (Florence, Taub, & Kaas, 1998) (see
Methods 5.1).
Brain scan studies confirm that the same processes
occur with humans. Now consider what happens when
cells in a reorganized cortex become activated. Previ-
ously, those neurons responded to an arm, and now
they receive information from the face. Does the re-
sponse feel like stimulation on the face or on the arm?
The answer: It feels like the arm (Davis et al.,
1998). Physicians have long noted that many people
with amputations experience a phantom limb, a con-
tinuing sensation of an amputated body part. That ex-
perience can range from occasional tingling to intense
pain. It is possible to have a phantom hand, foot, in-
testines, breast, penis, or anything else that has been
amputated. Sometimes the phantom sensation fades Image not available due to copyright restrictions
within days or weeks, but sometimes it lasts a lifetime
(Ramachandran & Hirstein, 1998).
Until the 1990s, no one knew what caused phan-
tom pains, and most believed that the sensations were
coming from the stump of the amputated limb. Some
physicians even performed additional amputations,
removing more and more of the limb in a futile attempt
to eliminate the phantom sensations. But modern meth-
ods have demonstrated that phantom limbs develop
only if the relevant portion of the somatosensory cor-
tex reorganizes and becomes responsive to alternative
inputs (Flor et al., 1995). For example, axons represent-
ing the face may come to activate the cortical area pre-
viously devoted to an amputated hand. Whenever the
face is touched, the person still feels it on the face but
Gray matter
Correct
Damage to visual cortex
Ventral root (motor)
Chapter Ending
Key Terms and Activities
Terms
apoptosis (p. 128) fetal alcohol syndrome (p. 130) neurotrophin (p. 129)
closed head injury (p. 137) focal hand dystonia (p. 135) penumbra (p. 138)
collateral sprout (p. 141) ganglioside (p. 141) phantom limb (p. 143)
deafferent (p. 144) hemorrhage (p. 137) proliferation (p. 123)
denervation supersensitivity ischemia (p. 137) stem cells (p. 125)
(p. 141) migration (p. 123) stroke (or cerebrovascular
diaschisis (p. 139) myelination (p. 124) accident) (p. 137)
differentiation (p. 124) nerve growth factor (NGF) synaptogenesis (p. 125)
disuse supersensitivity (p. 141) (p. 128) tissue plasminogen activator (tPA)
edema (p. 138) neural Darwinism (p. 128) (p. 138)
Websites http://www.thomsonedu.com
to Explore Go to this site for the link to ThomsonNOW, your one-stop study
shop, Take a Pre-Test for this chapter, and ThomsonNOW will
You can go to the Biological Psychology Study generate a Personalized Study Plan based on your test results.
Center and click this link. While there, you The Study Plan will identify the topics you need to review and
can also check for suggested articles available direct you to online resources to help you master these topics.
on InfoTrac College Edition. The Biological Psychol- You can then take a Post-Test to help you determine the con-
ogy Internet address is: cepts you have mastered and what you still need work on.
http://psychology.wadsworth.com/book/kalatbiopsych9e/
151
Module 6.1
Visual Coding and the
Retinal Receptors
Cornea
Lens
Ciliary muscle
(controls the lens) Optic
Retina nerve
Optic nerve
Horizontal cell
Amacrine cell
Ganglion cells
Bipolar cells
Receptors
Visual Receptors:
Rods and Cones
The vertebrate retina contains two types of receptors:
rods and cones (Figure 6.6). The rods, which are most
abundant in the periphery of the human retina, re-
spond to faint light but are not useful in bright day-
light because bright light bleaches them. Cones, which
are most abundant in and around the fovea, are less
active in dim light, more useful in bright light, and es-
Chase Swift
sential for color vision. Because of the distribution of
rods and cones, you have good color vision in the fovea
but not in the periphery. The differences between fo-
Figure 6.5 A behavioral consequence of how veal and peripheral vision are summarized in Table 6.1.
receptors are arranged on the retina Although rods outnumber cones about 20 to 1 in the
One owlet has turned its head almost upside down to see human retina, cones have a much more direct route to
above itself. Birds of prey have a great density of receptors the brain. Remember the midget ganglion cells: In the
on the upper half of the retina, enabling them to see below fovea (all cones), each receptor has its own line to the
them in great detail during flight. But they see objects brain. In the periphery (mostly rods), each receptor
above themselves poorly, unless they turn their heads. Take shares a line with tens or hundreds of others. A typi-
another look at the prairie falcon at the start of this chapter. cal count shows about 10 cone-driven responses in the
It is not a one-eyed bird; it is a bird that has tilted its head. brain for every rod-driven response (Masland, 2001).
Do you now understand why? The ratio of rods to cones varies among species. A
20-to-1 ratio may sound high, but in fact, the ratio is
much higher in rodents and other
species that are active at night.
An extreme case is that of South
American oilbirds, which live in
caves and emerge only at night to
search for fruits. They have about
15,000 rods per cone. Further-
more, as an adaptation to detect
faint lights at night, their rods are
packed three deep throughout the
retina (Martin, Rojas, Ramrez, &
McNeil, 2004).
Both rods and cones contain
photopigments, chemicals that re-
lease energy when struck by light.
E. R. Lewis, F. S. Werblin, & Y. Y. Zeevi
Receptors Cones in the fovea itself; cones and Proportion of rods increases toward the periphery;
rods mix in the surrounding area the extreme periphery has only rods
Convergence of Just a few receptors send their input Increasing numbers of receptors send input to
receptors to each postsynaptic cell each postsynaptic cell
Brightness sensitivity Useful for distinguishing among bright Responds well to faint lights; less useful for making
lights; responds poorly to faint lights distinctions in bright light
Sensitivity to detail Good detail vision because few receptors Poor detail vision because so many receptors send
funnel their input to a postsynaptic cell their input to the same postsynaptic cell
Color vision Good (many cones) Poor (few cones)
100
color vision.
Figure 6.8 shows wavelength-sensitivity
functions for the three cone types: short- 75
wavelength, medium-wavelength, and long-
wavelength. Note that each cone responds 50
to a broad band of wavelengths but to some
more than others.
25
According to the trichromatic theory,
we discriminate among wavelengths by
the ratio of activity across the three types 0
400 450 500 550 600 650
of cones. For example, light at 550 nm ex-
Wavelength (nanometers)
cites the medium-wavelength and long-
wavelength receptors about equally and Figure 6.8 Response of rods and three kinds of cones to
the short-wavelength receptor almost not various wavelengths of light
at all. This ratio of responses among the Note that each kind responds somewhat to a wide range of wavelengths
three cones determines a perception of yel- but best to wavelengths in a particular range. (Source: From J. K. Bowmaker
low. More intense light increases the activ- and H. J. A. Dartnell, Visual pigments of rods and cones in a human retina,
ity of all three cones without much change Journal of Physiology, 298, 501511. Copyright 1980. Reprinted by permission
in their ratio of responses. As a result, the of the author.)
Furthermore, the three kinds of cones are distributed at it under a bright light, without moving your eyes,
randomly within the retina (Roorda, Metha, Lennie, & for a full minute. (The brighter the light and the longer
Williams, 2001; Roorda & Williams, 1999). Figure 6.9 you stare, the stronger the effect.) Then look at a plain
shows the distribution of short-, medium-, and long-
wavelength cones in two peoples retinas, with colors
artificially added to distinguish the three cone types.
Note how few short-wavelength cones are present.
Note also the patches of all medium- or all long-wave-
length cones, which vary from one person to another.
Some people have more than 10 times as many of one
kind as the other over the whole retina. Surprisingly,
these variations do not produce any easily measured
difference in peoples ability to discriminate one color
from another (Miyahara, Pokorny, Smith, Baron, &
Baron, 1998). Evidently, the brain can compensate for
variations in input over a wide range.
That compensation breaks down, however, in the
periphery of the retina, where cones are scare and their
connections are somewhat haphazard (Diller et al.,
2004; Martin, Lee, White, Solomon, & Rtiger, 2001).
Try this: Get someone to mark a colored dot on the tip
of your finger, or attach a colored sticker, without tell-
ing you the color. Slowly move it from behind your
head into your field of vision and then gradually to-
ward your fovea. At what point do you see the color?
You will see your finger long before you can iden-
tify the color. As a rule, the smaller the dot, the far-
ther you will have to move it into your visual field
that is, the part of the world that you seebefore you
can identify the color. The On- ONLINE
line Try It Yourself exercise
Color Blindness in the Visual try it try it
Periphery provides a further yourself yourself
illustration.
The trichromatic theory correctly predicted the dis- Figure 6.10 Stimulus for demonstrating negative
covery of three kinds of cones, but it is incomplete as color afterimages
a theory of color vision. For example, try the follow- Stare at any point on the face under bright light for about
ing demonstration: Pick any point in the top portion a minute and then look at a white field. You should see a
of Figure 6.10such as the tip of the noseand stare negative afterimage.
For the afterimage of the surrounding box, you in fact, if you used a bright enough light, you saw a
saw red, as expected from the theory. But what about green afterimage.
the circle inside? Theoretically, you should have seen Here is another demonstration: First, look at Fig-
a gray or black afterimage (the opposite of white), but ure 6.12. Note that although it shows four red quarter-
(b)
(c)
6. Suppose a bipolar cell received excitatory input from ing on the screen than when the set was off and the
medium-wavelength cones and inhibitory input from screen appeared gray. What accounts for the black
all three kinds of cones. When it is highly excited, perception?
what color would one see? When it is inhibited, what 8. Figure 6.8 shows 500 nm light as blue and 550 nm
color perception would result? light as yellow. Why should we nevertheless not call
7. When a television set is off, its screen appears gray. them blue light and yellow light?
When you watch a program, parts of the screen ap- Check your answers on page 164.
pear black, even though more light is actually show-
Color Vision Deficiency sponse to a slightly longer wavelength than other peo-
ples receptors (Stockman & Sharpe, 1998). The gene
Encyclopedias describe many examples of discover- controlling this receptor is on the X chromosome, so
ies in astronomy, biology, chemistry, and physics, but because men have only one X chromosomemen have
what are psychologists discoveries? You might give only one of these LW receptor types or the other. How-
that question some thought. One of my nominations ever, women have two X chromosomes. In each cell,
is color blindness, better described as color vision de- one X chromosome is activated and the other is inhib-
ficiency, an impairment in perceiving color differences ited, apparently at random.2 Therefore, nearly half of
compared to other people. (Complete color blindness, women have both kinds of long-wavelength genes and
the inability to perceive anything but shades of black two kinds of red receptors (Neitz, Kraft, & Neitz, 1998).
and white, is rare.) Before color vision deficiency was Several studies have found that such women draw
discovered in the 1600s, people assumed that vision slightly finer color distinctions than other people do.
copies the objects we see (Fletcher & Voke, 1985). That That is, sometimes two objects that seem the same color
is, if an object is round, we see the roundness; if it is to other people look different to women with two kinds
yellow, we see the yellowness. Investigators discovered of LW receptors (Jameson, Highnote, & Wasserman,
that it is possible to have otherwise satisfactory vision 2001). This effect is small, however, and emerges only
without seeing color. That is, color depends on what with careful testing.
our brains do with incoming light; it is not a property For more information about the retina and vision
of the light by itself. and vision in general, this site provides an excellent
Several types of color vision deficiency occur. For treatment: http://www.webvision.med.utah.edu
genetic reasons, some people lack one or two of the
types of cones. Some have all three kinds, but one kind
has abnormal properties (Nathans et al., 1989). In the
most common form of color vision deficiency, people
have trouble distinguishing red from green because STOP & CHECK
their long- and medium-wavelength cones have the
same photopigment instead of different ones. The gene
9. As mentioned on page 158, most people can use
causing this deficiency is on the X chromosome. About
varying amounts of three colors to match any other
8% of men are red-green color blind compared with
color that they see. Who would be an exception to
less than 1% of women (Bowmaker, 1998).
this rule, and how many would they need?
Check your answer on page 164.
E X T E N S I O N S A N D A P P L I C AT I O N S
People with Four Cone Types
Might anyone have more than three kinds of cones? 2There is a good reason one X chromosome per cell is inhibited.
Apparently, some women do. People vary in the gene The genes on any chromosome produce proteins. If both X chro-
controlling the long-wavelength (LW) cone receptor mosomes were active in women, then either women would be get-
ting an overdose of the X-related proteins or men would be getting
(sensitive mainly in the red end of the spectrum). Con- an underdose. Because only one X chromosome is active per cell
sequently, some peoples LW receptors have a peak re- in women, both men and women get the same dose (Arnold, 2004).
(b) Photo of a
cross-section through the retina.
This section from the periphery of
the retina has relatively few ganglion
cells; a slice closer to the fovea
Image not available due to copyright restrictions would have a greater density.
Ganglion
cells retina.
Axons from Lateral inhibition is the retinas way
(b) ganglion cells of sharpening contrasts to emphasize
the border between one object and the
next. We begin with the rods and cones.
They have spontaneous levels of activ-
ity, and light striking them actually de-
creases their output. They have inhibitory synapses
Processing in the Retina onto the bipolar cells, and therefore, light decreases
their inhibitory output. Because you and I have trou-
The human retina contains roughly 120 million rods ble thinking in double negatives, for simplicitys sake,
and 6 million cones. We cannot intelligently pro- lets think of their output as excitation of the bipolar
cess that many independent messages; we need to cells. In the fovea, each cone attaches to just one bi-
Direction of light
(a)
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15
3. If light strikes only one receptor, what is the net effect
Receptors (excitatory or inhibitory) on the nearest bipolar cell
that is directly connected to that receptor? What is
the effect on bipolar cells off to the sides? What causes
Horizontal
cell that effect?
12 13
4. Examine Figure 6.17. You should see grayish dia-
1 2 3 4 5 6 8 9 10 11 14 15 Bipolar
7 monds at the crossroads among the black squares.
cells
Explain why.
Check your answers on page 183.
Low activity Low activity
High activity
Greatest activity
Pathways to the Lateral
Least activity Geniculate and Beyond
Look out your window. Perhaps you see someone walk-
ing by. Although your perception of that person seems
to be an integrated whole, different parts of your brain
are analyzing different aspects. One set of neurons iden-
tifies the persons shape, another set concentrates on
the colors, and another sees the speed and direction of
movement (Livingstone, 1988; Livingstone & Hubel,
1988; Zeki & Shipp, 1988). Your visual pathway begins
its division of labor long before it reaches the cerebral
cortex. Even at the level of the ganglion cells in the
retina, different cells react differently to the same input.
Each cell in the visual system of the brain has what
we call a receptive field, which is the part of the visual
field that either excites it or inhibits it. For a receptor,
the receptive field is simply the point in space from
which light strikes the receptor. Other visual cells de-
rive their receptive fields from the pattern of excita-
tory and inhibitory connections to them. For example,
a ganglion cell is connected to a group of bipolar cells,
which in turn are connected to receptors, so the recep-
Figure 6.17 An illustration of lateral tive field of the ganglion cell is the combined recep-
inhibition tive fields of those receptors, as shown in Figure 6.18.
Do you see dark diamonds at the crossroads? The receptive fields of the ganglion cells converge to
ulate have receptive fields that resemble those of the ever, many people with area V1 damage do not show
ganglion cellseither an excitatory or an inhibitory blindsight or show it for stimuli only in certain loca-
central portion and a surrounding ring of the opposite tions (Schrli, Harman, & Hogben, 1999; Wessinger,
effect. Again, some have large (magnocellular) recep- Fendrich, & Gazzaniga, 1997). An alternative explana-
tive fields, and others have small (parvocellular) fields. tion is that tiny islands of healthy tissue remain within
However, after the information reaches the cerebral cor- an otherwise damaged visual cortex, not large enough
tex, the receptive fields begin to become more special- to provide conscious perception but nevertheless
ized and complicated. enough for blindsight (Fendrich, Wessinger, & Gaz-
zaniga, 1992). Some patients experience blindsight
after extensive (but not total) damage to the optic nerve,
so the surviving island theory does appear valid for
Pattern Recognition some cases (Wst, Kasten, & Sabel, 2002).
in the Cerebral Cortex Perhaps both hypotheses are correct. In some pa-
tients, a small amount of recordable activity in area V1
Most visual information from the lateral geniculate nu- accompanies blindsight, supporting the islands ex-
cleus of the thalamus goes to the primary visual cortex planation. In other patients, no activity in V1 is appar-
in the occipital cortex, also known as area V1 or the ent, and blindsight evidently depends on other brain
striate cortex because of its striped appearance. It is areas (Morland, L, Carroll, Hofmann, & Pambakian,
the area of the cortex responsible for the first stage of 2004). In one study, experimenters temporarily sup-
visual processing. Area V1 is apparently essential for pressed a wide area of the visual cortex by transcranial
most, if not all, conscious vision. If you close your eyes magnetic stimulation (see page 110). Although people
and imagine a visual scene, activity increases in area were not aware of a spot flashed on the screen during
V1 (Kosslyn & Thompson, 2003). People who become the period of suppression, the spot influenced their
blind because of eye damage continue having visual eye movements (Ro, Shelton, Lee, & Chang, 2004). That
dreams if their visual cortex is intact. However, peo- result also indicates that activity outside V1 can pro-
ple with extensive damage to the visual cortex report duce visually guided behavior.
no conscious vision, no visual imagination, and no vi- Still, all these blindsight responses occur without
sual images in their dreams (Hurovitz, Dunn, Domhoff, consciousness. The conclusion remains that conscious
& Fiss, 1999). visual perception requires strong activity in area V1.
Nevertheless, some people with extensive area V1
damage show a surprising phenomenon called blind-
sight, an ability to respond in some ways to visual in-
Pathways in the Visual Cortex
formation that they report not seeing. If a light flashes The primary visual cortex sends information to the sec-
within an area where they report no vision, they can ondary visual cortex (area V2), which processes the
nevertheless point to it or turn their eyes toward it, information further and transmits it to additional areas,
while insisting that they saw nothing and are only as shown in Figure 6.19. The connections in the visual
guessing (Bridgeman & Staggs, 1982; Weiskrantz, War- cortex are reciprocal; for example, V1 sends informa-
rington, Sanders, & Marshall, 1974). tion to V2 and V2 returns information to V1. Neuro-
The explanation remains controversial. After dam- scientists have distinguished 30 to 40 visual areas in
age to area V1, other branches of the optic nerve de- the brain of a macaque monkey (Van Essen & Deyoe,
liver visual information to the superior colliculus (in 1995) and believe that the human brain has even more.
the midbrain) and several other areas, including parts Within the cerebral cortex, a ventral pathway, with
of the cerebral cortex (see Figure 6.16a). Perhaps those mostly parvocellular input, is sensitive to details of
areas control the blindsight responses (Cowey & Stoerig, shape. Another ventral pathway, with mostly magno-
1995; Moore, Rodman, Repp, & Gross, 1995). How- cellular input, is sensitive to movement. Still another,
METHODS 6.1
Microelectrode Recordings
David Hubel and Torsten Wiesel pioneered the use of tip or a narrow glass tube containing a salt solution and
microelectrode recordings to study the properties of in- a metal wire. They direct the electrode either next to or
dividual neurons in the cerebral cortex. In this method, into a single cell and then record its activity while they
investigators begin by anesthetizing an animal and drill- present various stimuli, such as patterns of light. Re-
ing a small hole in the skull. Then they insert a thin elec- searchers use the results to determine what kinds of
trodeeither a fine metal wire insulated except at the stimuli do and do not excite the cell.
simple cells, and they found that as soon as the simple End-stopped, or hypercomplex, cells resemble
cells stopped responding, the complex cells stopped complex cells with one additional feature: An end-
too (Martinez & Alonso, 2001). stopped cell has a strong inhibitory area at one end of
its bar-shaped receptive field. The cell responds to a
bar-shaped pattern of light anywhere in its broad re-
Time when stimulus is present
ceptive field provided that the bar does not extend
beyond a certain point (Figure 6.23). Table 6.3 sum-
marizes the properties of simple, complex, and end-
stopped cells.
High response
High response
Low response
Figure 6.22 The receptive field of a complex Figure 6.23 The receptive field of an
cell in the visual cortex end-stopped cell
Like a simple cell, its response depends on a bar of lights The cell responds to a bar in a particular orientation (in this
angle of orientation. However, a complex cell responds the case, horizontal) anywhere in its receptive field provided
same for a bar in any position within the receptive field. that the bar does not extend into a strongly inhibitory area.
174 Chapter 6 Vision
Table 6.3 Summary of Cells in the Primary Visual Cortex
Characteristic Simple Cells Complex Cells End-Stopped Cells
Disorders of
Object Recognition
Damage to the shape pathway of the cortex should lead
to specialized deficits in the ability to recognize objects.
Neurologists have reported such cases for decades, al-
though they frequently met with skepticism. Now that
we understand how such specialized defects might
arise, we find them easier to accept.
An inability to recognize objects despite otherwise
satisfactory vision is called visual agnosia (meaning
visual lack of knowledge). It usually results from
Mirror image Figure-ground reversal damage somewhere in the temporal cortex. A person
with brain damage might be able to point to visual ob-
Figure 6.25 Three transformations of an jects and slowly describe them but fail to recognize
original drawing what they are or mean. For example, one patient, when
In the inferior temporal cortex, cells that respond strongly to shown a key, said, I dont know what that is; perhaps
the original respond about the same to the contrast reversal a file or a tool of some sort. When shown a stethoscope,
and mirror image but not to the figureground reversal. he said that it was a long cord with a round thing at
Note that the figureground reversal resembles the original the end. When he could not identify a pipe, the exam-
very strongly in terms of the pattern of light and darkness; iner told him what it was. He then replied, Yes, I can
however, it is not perceived as the same object. (Source: Based see it now, and pointed out the stem and bowl of the
on Baylis & Driver, 2001) pipe. Then the examiner asked, Suppose I told you
that the last object was not really a pipe? The patient
replied, I would take your word for it. Perhaps its not
our capacity for shape constancythe ability to rec- really a pipe (Rubens & Benson, 1971).
ognize an objects shape even as it changes location or Many other types of agnosia occur. One closed head
direction. However, although shape constancy helps us injury patient could recognize faces of all kinds, in-
recognize an object from different angles, it interferes cluding cartoons and face pictures made from objects
in reading, where we need to treat mirror-image letters (Figure 6.26). However, he could not recognize any of
Fusiform
gyrus
Dr. Dana Copeland
EXTENSIONS
A N D A P P L I C AT I O N S
Suppressed Vision During
Eye Movements
The temporal cortex has cells that dis-
tinguish between moving objects and vi-
sual changes due to head movements. An
additional mechanism prevents confu-
Figure 6.30 Stimuli that excite the ventral part of area MST sion or blurring during eye movements.
Cells here respond when an object moves relative to its background. They Before the explanation, try this demonstra-
therefore react either when the object moves or when the object is steady tion: Look at yourself in a mirror and fo-
and the background moves. cus on your left eye. Then shift your focus
Summary
1. The optic nerves of the two eyes join at the optic
chiasm, where half of the axons from each eye
When you are instructed to attend to the lines in
cross to the opposite side of the brain. Most of the
one direction or the other, activity increases in the V1
axons then travel to the lateral geniculate nucleus
neurons that respond to lines in that orientation. As
of the thalamus, which communicates with the vi-
your attention shifts from one orientation to the other,
sual cortex. (p. 166)
so does activity in the two sets of neurons. In fact, some-
one who is monitoring your visual cortex with fMRI 2. Lateral inhibition is a mechanism by which stim-
could tell you which set of lines you were attending ulation in any area of the retina suppresses the re-
to (Kamitani & Tong, 2005). sponses in neighboring areas, thereby enhancing
Also, if you are told to pay attention to color or mo- the contrast at lightdark borders. (p. 167)
tion, activity increases in the areas of your visual cor- 3. Lateral inhibition in the vertebrate retina occurs
tex responsible for color or motion perception (Chawla, because receptors stimulate bipolar cells and also
Rees, & Friston, 1999). In fact, activity increases in those stimulate the much wider horizontal cells, which
areas even before the stimulus (Driver & Frith, 2000). inhibit both the stimulated bipolar cells and those
Somehow the instructions prime those areas, so that to the sides. (p. 168)
they can magnify their responses to any appropriate 4. Each neuron in the visual system has a receptive
stimulus. They in turn feed back to area V1, enhancing field, an area of the visual field to which it is con-
that areas response to the stimulus. Again, it appears nected. Light in the receptive field excites or in-
that the feedback increase in V1 responses is necessary hibits the neuron depending on the lights location,
for attention or conscious awareness of a stimulus wavelength, movement, and so forth. (p. 169)
(Pascual-Leone & Walsh, 2001).
5. The mammalian vertebrate visual system has a
partial division of labor. In general, the parvocel-
lular system is specialized for perception of color
Module 6.2 and fine details; the magnocellular system is spe-
In Closing: From Single Cells to Vision cialized for perception of depth, movement, and
In this module, you have read about single cells that overall patterns. (p. 170)
respond to shape, movement, and other aspects of vi- 6. Area V1 is apparently essential to conscious vi-
sion. Does any single cell identify what you see? sual perception. Without it, people report no vi-
Several decades ago, the early computers used to sion, even in dreams. However, some kinds of re-
crash frequently. Some of the pioneers of computer sci- sponse to light (blindsight) can occur after damage
ence were puzzled. A single neuron in the brain, they to V1, despite the lack of conscious perception.
realized, was surely no more reliable than a single com- (p. 171)
realistic one and a distorted one (Cassia, Turati, & Sim- nch for perset
iits of photogr
why later t
ion, 2004). Evidently, a newborns concept of face is
Face Circles Newsprint White Yellow Red
not well developed, except that it requires most of its
content (e.g., eyes) to be on top. Figure 6.31 Amount of time infants spend looking
Of course, a person gradually becomes more and at various patterns
more adept at recognizing faces. Some day you may go Even in the first 2 days after birth, infants look more at faces
to a high school reunion and quickly recognize people than at most other stimuli. (Source: Based on Fantz, 1963)
Here is a fascinating demonstration that combines (Gdecke & Bonhoeffer, 1996; Horton & Hocking, 1996),
visual attention with motor control: Hold one hand to and many of the normal properties develop even for
the left of a childs head and the other hand to the right. animals with retinal damage (Rakic & Lidow, 1995;
Tell the child that when you wiggle a finger of one Shatz, 1996) or those reared in complete darkness (Lein
hand, he or she should look at the other hand. Before & Shatz, 2001; White, Coppola, & Fitzpatrick, 2001).
about age 5 or 6 years, most children find it almost im- However, without normal visual experience, those
possible to ignore the wiggling finger and to look at the properties deteriorate. The visual system can mature to
other one. Ability to perform this task smoothly devel- a certain point without experience, but it needs visual
ops gradually all the way to age 18, requiring areas of the experience to maintain and fine-tune its connections.
prefrontal cortex that mature slowly. Even
some adultsespecially those with neuro- try it
logical or psychiatric disordershave trou- yourself
Early Lack of Stimulation of One Eye
ble on this task (Munoz & Everling, 2004). What would happen if a young animal could see with
To examine visual development in more detail, one eye but not the other? For cats and primates
investigators turn to studies of animals. The research which have both eyes pointed in the same direction
in this area has greatly expanded our understanding most neurons in the visual cortex receive binocular
of brain development and has helped alleviate certain input (stimulation from both eyes). As soon as a kit-
human abnormalities. ten opens its eyes at about age 9 days, each neuron re-
sponds to approximately corresponding areas in the
two retinasthat is, areas that focus on about the same
point in space (Figure 6.33).
Early Experience If an experimenter sutures one eyelid shut for a
and Visual Development kittens first 4 to 6 weeks of life, synapses in the visual
cortex gradually become unresponsive to input from
Developing axons of the visual system approach their the deprived eye (Rittenhouse, Shouval, Paradiso, &
targets by following chemical gradients, as discussed Bear, 1999). Consequently, even after the deprived eye
in Chapter 5. In a newborn mammal, the lateral genic- is opened, the kitten does not respond to it (Wiesel,
ulate and visual cortex already resemble an adults 1982; Wiesel & Hubel, 1963).
Early Exposure to a
Limited Array of Patterns
If a kitten spends its entire early sensitive period wear-
ing goggles with horizontal lines painted on them (Fig-
ure 6.34), nearly all its visual cortex cells become re-
sponsive only to horizontal lines (Stryker & Sherk,
1975; Stryker, Sherk, Leventhal, & Hirsch, 1978). Even
after months of later normal experience, the cat does
not respond to vertical lines (D. E. Mitchell, 1980).
What happens if human infants are exposed mainly
to vertical or horizontal lines instead of both equally?
You might wonder how such a bizarre thing could hap-
Sue Ford/Science Source/Photo Researchers
Left Right
retina retina
Left Right
retina retina
Terms
astigmatism (p. 187) law of specific nerve energies receptor potential (p. 152)
binocular input (p. 185) (p. 152) retina (p. 153)
bipolar cell (p. 153) lazy eye (p. 187) retinal disparity (p. 186)
blind spot (p. 154) magnocellular neuron (p. 170) retinex theory (p. 161)
blindsight (p. 171) midget ganglion cell (p. 155) rod (p. 156)
color constancy (p. 161) motion blindness (p. 181) saccade (p. 181)
color vision deficiency (p. 163) MST (p. 179) secondary visual cortex (or
complex cell (p. 173) MT (or area V5) (p. 179) area V2) (p. 171)
cone (p. 156) negative color afterimage (p. 160) sensitive period (or critical period)
opponent-process theory (p. 160) (p. 186)
dorsal stream (p. 172)
optic nerve (p. 154) shape constancy (p. 177)
end-stopped cell (p. 174)
parvocellular neuron (p. 170) simple cell (p. 173)
feature detector (p. 175)
photopigment (p. 156) stereoscopic depth perception
fovea (p. 155)
(p. 179)
ganglion cell (p. 153) primary visual cortex (or area V1)
(p. 171) strabismus (or strabismic
horizontal cell (p. 166) amblyopia) (p. 186)
prosopagnosia (p. 178)
hypercomplex cell (p. 174) trichromatic theory (or Young-
psychophysical observations
inferior temporal cortex (p. 176) Helmholtz theory) (p. 158)
(p. 158)
koniocellular neuron (p. 170) ventral stream (p. 172)
pupil (p. 153)
lateral geniculate nucleus (p. 166) visual agnosia (p. 177)
receptive field (p. 169)
lateral inhibition (p. 169) visual field (p. 159)
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195
Module 7.1
Audition
Dimensions of Sound
Sound waves vary in amplitude and frequency. The Low amplitude
amplitude of a sound wave is its intensity. A very in-
tense compression of air, such as that produced by a
Higher amplitude
bolt of lightning, produces sound waves of great ampli-
tude. Loudness, the perception of intensity, is related
to amplitude but not the same as it. When a sound
doubles its amplitude, its loudness increases but does
0.1 second
not double. Many factors influence loudness; for ex-
ample, a rapidly talking person sounds louder than Figure 7.1 Four sound waves
slow music of the same physical amplitude. So if you The time between the peaks determines the frequency of
complain that television advertisements are louder the sound, which we experience as pitch. Here the top line
than the program, one reason is that the people in the represents five sound waves in 0.1 second, or 50 Hza very
advertisements talk faster. low-frequency sound that we experience as a very low pitch.
The frequency of a sound is the number of com- The other three lines represent 100 Hz. The vertical extent
pressions per second, measured in hertz (Hz, cycles of each line represents its amplitude or intensity, which we
per second). Pitch is a perception closely related to fre- experience as loudness.
Anvil
Hammer
Scala Scala
Hair cells
vestibuli media Tympanic
membrane Auditory
Stirrup nerve
(b)
Oval window
(membrane behind stirrup)
Round window
Tectorial membrane
Scala
(c) tympani
Hair cells
Cochlear
neuron
Pitch Perception
Hudspeth, 1985
Our ability to understand speech or enjoy music de-
pends on our ability to differentiate among sounds of
different frequencies. How do we do it? Figure 7.3 Hair cells from the auditory systems
of three species
Frequency Theory (a, b) Hair cells from a frog sacculus, an organ that detects
ground-borne vibrations. (c) Hair cells from the cochlea
and Place Theory of a cat. (d) Hair cells from the cochlea of a fence lizard.
According to the frequency theory, the basilar mem- Kc = kinocilium, one of the components of a hair bundle.
brane vibrates in synchrony with a sound, causing au-
ditory nerve axons to produce action potentials at the
same frequency. For example, a sound at 50 Hz would The current theory combines modified versions of
cause 50 action potentials per second in the auditory both frequency and place theories. For low-frequency
nerve. The downfall of this theory in its simplest form sounds (up to about 100 Hzmore than an octave be-
is that the refractory period of a neuron is about 1 1,000 low middle C in music, which is 264 Hz), the basilar
second, so the maximum firing rate of a neuron is about membrane does vibrate in synchrony with the sound
1 Hz, far short of the highest frequencies we hear. waves, in accordance with the frequency theory, and
According to the place theory, the basilar mem- auditory nerve axons generate one action potential per
brane resembles the strings of a piano in that each area wave. Weak sounds activate few neurons, whereas
along the membrane is tuned to a specific frequency stronger sounds activate more. Thus, at low frequen-
and vibrates in its presence. (If you loudly sound one cies, the frequency of impulses identifies the pitch,
note with a tuning fork near a piano, you vibrate the and the number of firing cells identifies the loudness.
piano string tuned to that note.) According to this the- Because of the refractory period of the axon, as
ory, each frequency activates the hair cells at only one sounds exceed 100 Hz, it is harder and harder for a
place along the basilar membrane, and the nervous sys- neuron to continue firing in synchrony with the sound
tem distinguishes among frequencies based on which waves. At higher frequencies, it fires on every second,
neurons are activated. The downfall of this theory is third, fourth, or later wave. Its action potentials are
that the various parts of the basilar membrane are bound phase-locked to the peaks of the sound waves (i.e., they
together too tightly for any part to resonate like a piano occur at the same phase in the sound wave), as illus-
string. trated here:
Sound wave
1000 Hz
Neuron 1
Neuron 2 400 Hz
Neuron 3 1500 Hz 200 Hz
Sum of
neurons
7000 Hz
If we consider the auditory nerve as a whole, we
find that with a tone of a few hundred Hz, each wave
excites at least a few auditory neurons. According to
the volley principle of pitch discrimination, the audi- Base of cochlea (Stiff)
tory nerve as a whole can have volleys of impulses up (by oval window)
to about 4000 per second, even though no individual 20000 Hz
axon approaches that frequency by itself (Rose, Brugge,
Anderson, & Hind, 1967). Beyond about 4000 Hz, even Figure 7.4 The basilar membrane of the
staggered volleys of impulses cant keep pace with the human cochlea
sound waves. Neuroscientists assume that these vol- High-frequency sounds excite hair cells near the base.
leys contribute to pitch perception, although no one Low-frequency sounds excite cells near the apex.
understands how the brain uses the information.
Most human hearing takes place below 4000 Hz,
the approximate limit of the volley principle. For com- ine, they dont enjoy music (Hyde & Peretz, 2004). The
parison, the highest key on a piano is 4224 Hz. Frequen- explanation for this phenomenon is not known.
cies much above that level are not important in music
or human speech, although they are important in the
lives of rats, mice, bats, and other small animals. When
we hear these very high frequencies, we use a mecha- STOP & CHECK
nism similar to the place theory. The basilar membrane
varies from stiff at its base, where the stirrup meets the
1. Through what mechanism do we perceive low-
cochlea, to floppy at the other end of the cochlea, the
frequency sounds (up to about 100 Hz)?
apex (von Bksy, 1956) (Figure 7.4). The hair cells
along the basilar membrane have different properties 2. How do we perceive middle-frequency sounds
based on their location, and they act as tuned resona- (100 to 4000 Hz)?
tors that vibrate only for sound waves of a particular 3. How do we perceive high-frequency sounds
frequency. The highest frequency sounds vibrate hair (above 4000 Hz)?
cells near the base, and lower frequency sounds vibrate Check your answers on page 204.
hair cells farther along the membrane (Warren, 1999).
Actually, the mechanisms of hearing at frequencies well
over 4000 Hz are not entirely understood, as these ultra-
high frequencies alter several of the properties of neu-
rons and their membranes (Fridberger et al., 2004). The Auditory Cortex
Some people are tone deaf. The fancy term for
this condition is amusia. Anyone who was completely Information from the auditory system passes through
insensitive to frequency differences could not under- several subcortical structures, with a crossover in the
stand speech because every sound we make depends midbrain that enables each hemisphere of the forebrain
on fast and slight changes in pitch. However, a substan- to get most of its input from the opposite ear (Glenden-
tial number of peopleby one estimate, about 4% ning, Baker, Hutson, & Masterton, 1992). The informa-
are seriously impaired at detecting small changes in tion ultimately reaches the primary auditory cortex
frequency, such as between C and C#. As you can imag- (area A1) in the superior temporal cortex, as shown
Corresponds to Corresponds to
apex of basilar base of basilar
membrane membrane
Primary Secondary
auditory auditory
(a) cortex cortex
Sound source
Sound Localization
You are walking alone when suddenly you hear a loud
noise. You want to know what produced it (friend or
foe), but equally, you want to know where it came from Figure 7.7 Differential loudness and arrival times
(so you can approach or escape). Determining the di- as cues for sound localization
rection and distance of a sound requires comparing Sounds reaching the closer ear arrive sooner as well as
the responses of the two earswhich are in effect just louder because the head produces a sound shadow.
two points in space. And yet this system is accurate (Source: After Lindsay & Norman, 1972)
Somatosensory Receptors
The skin has many kinds of somatosensory recep-
tors, including those listed in Figure 7.11. Table 7.1
lists the probable functions of these and other recep-
tors (Iggo & Andres, 1982; Par, Smith, & Rice, 2002).
(a) However, each receptor probably contributes to sev-
eral kinds of somatosensory experience. Many re-
spond to more than one kind of stimulus, such as
Semicircular
touch and temperature. Others (not in the table) re-
canals spond to deep stimulation, joint movement, or mus-
cle movement.
A touch receptor may be a simple bare neuron
ending (e.g., many pain receptors), an elaborated neu-
ron ending (Ruffini endings and Meissners corpus-
cles), or a bare ending surrounded by nonneural cells
that modify its function (Pacinian corpuscles). Stim-
ulation of a touch receptor opens sodium channels
in the axon, thereby starting an action potential (Price
Saccule et al., 2000).
and utricle One example of a receptor is the Pacinian cor-
Inner ear puscle, which detects sudden displacements or high-
frequency vibrations on the skin (Figure 7.12). Inside
its onionlike outer structure is the neuron membrane.
(b)
When mechanical pressure bends the membrane, its
resistance to sodium flow decreases, and sodium ions
enter, depolarizing the membrane (Loewenstein, 1960).
Only a sudden or vibrating stimulus can bend the mem-
Otoliths brane; the onionlike outer structure provides mechan-
ical support that resists gradual or constant pressure
on the skin.
Hair cell Certain chemicals can also stimulate the receptors
for heat and cold. The heat receptor responds to cap-
saicin, the chemical that makes jalapeos and similar
peppers taste hot. The coolness receptor responds to
menthol and less strongly to mint (McKemy, Neu-
hausser, & Julius, 2002). So advertisements mention-
ing the cool taste of menthol are literally correct.
Vestibular nerve fibers
E X T E N S I O N S A N D A P P L I C AT I O N S
(c) Tickle
Figure 7.10 Structures for vestibular sensation The sensation of tickle is interesting but poorly under-
(a) Location of the vestibular organs. (b) Structures of the stood. Why does it exist at all? Why do you laugh if
vestibular organs. (c) Cross-section through a utricle. Calcium someone rapidly fingers your armpit, neck, or the soles
carbonate particles, called otoliths, press against different of your feet? Chimpanzees respond to similar sensa-
hair cells depending on the direction of tilt and rate of tions with bursts of panting that resemble laughter. And
acceleration of the head. yet tickling is unlike humor. Most people do not enjoy
206 Chapter 7 The Other Sensory Systems
Table 7.1 Somatosensory Receptors and Their Possible Functions
Receptor Location Responds to
Free nerve ending (unmyelinated Near base of hairs and elsewhere Pain, warmth, cold
or thinly myelinated axons) in skin
Hair-follicle receptors Hair-covered skin Movement of hairs
Meissners corpuscles Hairless areas Sudden displacement of skin; low-frequency vibration
(flutter)
Pacinian corpuscles Both hairy and hairless skin Sudden displacement of skin; high-frequency vibration
Merkels disks Both hairy and hairless skin Tangential forces across skin
Ruffini endings Both hairy and hairless skin Stretch of skin
Krause end bulbs Mostly or entirely in hairless areas, Uncertain
perhaps including genitals
Pacinian
corpuscle
C7
C8
Spinal cord Cervical nerves
(8 pairs) C6
I
T1
First thoracic II
vertebra III
C2 C5
T2
C3
C4
T3
T4
T5
C5 T6
Thoracic nerves T7
(12 pairs) T2 T8
T9
T10
C6 T11
T12
T1
L1
C7 S2
C8 S 2
L2 L2
S2
Lumbar nerves
(5 pairs)
L3 L3
S2 S2
L5 S2 L5
Sacral nerves L4 L4
(5 pairs)
S1 S1
Coccygeal nerves S1
L5
(1 pair)
Figure 7.13 The human central nervous Figure 7.14 Dermatomes innervated by
system (CNS) the 31 sensory spinal nerves
Spinal nerves from each segment of the spinal cord exit Areas I, II, and III of the face are not innervated by the
through the correspondingly numbered opening between spinal nerves but instead by three branches of the fifth
vertebrae. (Source: From C. Starr and R. Taggart, Biology: The cranial nerve. Although this figure shows distinct borders,
Unity and Diversity of Life, 5th edition, 1989, 338. Copyright 1989 the dermatomes actually overlap one another by about
Wadsworth Publishing Co. Reprinted by permission.) one-third to one-half of their width.
Itch
Have you ever wondered, What is itch, anyway? Is it
a kind of pain? A kind of touch? Or something else al- Module 7.2
together? For many years, no one knew, and we still In Closing: The Mechanical Senses
have not identified the receptors responsible for itch.
We humans generally pay so much attention to vision
However, we do know that when your skin rubs against
and hearing that we take our mechanical senses for
certain plants, when an insect crawls along your skin,
granted. However, a mere moments reflection should
or when you have mild tissue damage, your skin re-
reveal how critical they are for survival. At every mo-
leases histamines, and histamines produce an itching
ment, your vestibular sense tells you whether you are
sensation.
standing or falling; your sense of pain can tell you that
Researchers have identified a spinal cord pathway
you have injured yourself. If you moved to a television-
of itch sensation (Andrew & Craig, 2001). Histamines
like universe with only vision and hearing, you might
in the skin excite axons of this pathway, and other
get by if you had already learned what all the sights
kinds of skin stimuli do not. Even when a stimulus re-
and sounds mean. But it is hard to imagine how you
leases histamines, however, this pathway is slow to
could have learned their meaning without much pre-
respond, and when it does respond, the axons trans-
vious experience of touch and pain.
mit impulses at the unusually slow velocity of only
about half a meter per second. At that rate, an action
potential from your foot needs 3 or 4 seconds to reach
your head. For a giraffe or elephant, the delay is even Summary
longer. You might try very rubbing some
1. The vestibular system detects the position and ac-
rough leaves against your ankle. Note how try it
celeration of the head and adjusts body posture
soon you feel the touch sensation and how yourself
and eye movements accordingly. (p. 205)
much more slowly you notice the itchiness.
Itch is useful because it directs you to scratch the 2. The somatosensory system depends on a variety
itchy area and presumably remove whatever is irritat- of receptors that are sensitive to different kinds of
ing your skin. Vigorous scratching produces mild pain, stimulation of the skin and internal tissues. The
and pain inhibits itch. Opiates, which decrease pain, brain maintains several parallel somatosensory
increase itch (Andrew & Craig, 2001). This inhibitory representations of the body. (p. 206)
relationship between pain and itch is the strongest evi- 3. Activity in the primary somatosensory cortex cor-
dence that itch is not a type of pain. responds to what someone is experiencing, even
Answers to
Thought Questions
STOP & CHECK
1. Why is the vestibular sense generally useless under
Questions conditions of weightlessness?
1. The vestibular system enables the brain to shift eye 2. How could you determine whether hypnosis de-
movements to compensate for changes in head po- creases pain by increasing the release of endorphins?
1. Of the following, which use a labeled-line code and How Many Kinds
which use an across-fiber pattern code?
A. A fire alarm
of Taste Receptors?
B. A light switch Traditionally, people in Western society have described
C. The shift key plus another key on a keyboard sweet, sour, salty, and bitter as the primary tastes.
Check your answers on page 227. However, some tastes defy categorization in terms of
these four labels (Schiffman & Erickson, 1980; Schiff-
man, McElroy, & Erickson, 1980). How could we de-
termine how many kinds of taste we have?
Taste
E X T E N S I O N S A N D A P P L I C AT I O N S
Taste refers to the stimulation of the taste buds, the Chemicals That Alter the Taste Buds
receptors on the tongue. When we talk about the taste
of food, we generally mean flavor, which is a combi- One way to identify taste receptor types is to find pro-
nation of taste and smell. Whereas other senses re- cedures that alter one receptor but not others. For ex-
main separate throughout the cortex, taste and smell ample, chewing a miracle berry (native to West Africa)
axons converge onto some of the same cells in an area gives little taste itself but temporarily changes sweet
called the endopiriform cortex (Fu, Sugai, Yoshimura, receptors. Miracle berries contain a protein, miraculin,
& Onoda, 2004). That convergence presumably enables that modifies sweet receptors in such a way that they
taste and smell to combine their influences on food can be stimulated by acids (Bartoshuk, Gentile, Mosko-
selection. witz, & Meiselman, 1974). If you ever get a chance to
chew a miracle berry (and I do recommend it), for the
next half hour anything acidic will taste sweet in ad-
Taste Receptors dition to its usual sour taste.
The receptors for taste are not true neurons but modi- A colleague and I once spent an evening experi-
fied skin cells. Like neurons, taste receptors have ex- menting with miracle berries. We drank straight lemon
citable membranes and release neurotransmitters to juice, sauerkraut juice, even vinegar. All tasted ex-
excite neighboring neurons, which in turn transmit tremely sweet, but we awoke the next day with mouths
information to the brain. Like skin cells, however, taste full of ulcers.
receptors are gradually sloughed off and replaced, each Miraculin was, for a time, commercially available
one lasting about 10 to 14 days (Kinnamon, 1987). in the United States as a diet aid. The idea was that di-
Mammalian taste receptors are in taste buds, lo- eters could coat their tongue with a miraculin pill and
cated in papillae, structures on the surface of the tongue then enjoy unsweetened lemonade and so forth, which
(Figure 7.18). A given papilla may contain from zero would taste sweet but provide almost no calories.
to ten or more taste buds (Arvidson & Friberg, 1980), Have you ever drunk orange juice just after brush-
and each taste bud contains about 50 receptor cells. ing your teeth? Did you wonder why something so won-
In adult humans, taste buds are located mainly derful suddenly tasted so bad? Most toothpastes con-
along the outside edge of the tongue, with few or none tain sodium lauryl sulfate, a chemical that intensifies
in the center. You can demonstrate this principle as bitter tastes and weakens sweet ones, apparently by
follows: Soak a small cotton swab in sugar water, salt- coating the sweet receptors and preventing anything
water, or vinegar. Then touch it lightly on the center of from reaching them (DeSimone, Heck, & Bartoshuk,
your tongue, not too far toward the back. If you get the 1980; Schiffman, 1983).
position right, you will experience little or Another taste-modifying substance is an extract
no taste. Then try it again on the edge of try it from the plant Gymnema sylvestre (R. A. Frank, Mize,
your tongue and notice how much stronger yourself Kennedy, de los Santos, & Green, 1992). Some health-
the taste is. food and herbal-remedy stores sell dried leaves of
Now change the procedure a bit. Wash your mouth Gymnema sylvestre, from which you can brew a tea.
out with water and prepare a cotton swab as before. (Gymnema sylvestre pills wont work for this demon-
Touch the soaked portion to one edge of your tongue stration.) Soak your tongue in the tea for about 30 sec-
and then slowly stroke it to the center of your tongue. onds and then try tasting various substances. Salty,
Now it will seem as if you are moving the taste to the sour, and bitter substances taste the same as usual, but
center of your tongue. In fact, you are getting only a sugar becomes utterly tasteless, as if you had sand on
Taste buds
Further behavioral evidence for
separate types of taste receptors
comes from studies of the following
Foliate papilla type: Soak your tongue for 15 sec-
onds in a sour solution, such as un-
sweetened lemon juice. Then try
Fungiform tasting some other sour solution,
papilla
such as dilute vinegar. You will find
(a)
that the second solution tastes less
sour than usual. Depending on the concentrations of
the lemon juice and vinegar, the second solution may
not taste sour at all. This phenomenon, called adapta-
tion, reflects the fatigue of receptors sensitive to sour
tastes. Now try tasting something salty, sweet, or bit-
ter. These substances taste about the same as usual.
In short, you experience little cross-adaptationre-
duced response to one taste after exposure to another
(McBurney & Bartoshuk, 1973). Evidently, the sour re-
ceptors are different from the other taste
receptors. Similarly, you can show that salt try it
SIU/Peter Arnold, Inc.
Mechanisms of Taste Receptors 2. Suppose you find a new, unusual-tasting food. How
could you determine whether we have a special re-
The saltiness receptor is particularly simple. Recall that ceptor for that food or whether we taste it with a
a neuron produces an action potential when sodium combination of the other known taste receptors?
ions cross its membrane. A saltiness receptor, which
3. Although the tongue has receptors for bitter tastes,
detects the presence of sodium, simply permits sodium
researchers have not found neurons in the brain itself
ions on the tongue to cross its membrane. The higher
that respond more strongly to bitter than to other
the concentration of sodium on the tongue, the greater
tastes. Explain, then, how it is possible for the brain
the receptors response. Chemicals such as amiloride,
to detect bitter tastes.
which prevents sodium from crossing the membrane,
reduce the intensity of salty tastes (DeSimone, Heck, 4. If someone injected into your tongue some chemical
Mierson, & DeSimone, 1984; Schiffman, Lockhead, & that blocks the release of second messengers, how
Maes, 1983). would it affect your taste experiences?
Sourness receptors operate on a different princi- Check your answers on page 227.
ple. When an acid binds to the receptor, it closes potas-
sium channels, preventing potassium from leaving the
cell. The result is an increased accumulation of positive
charges within the neuron and therefore a depolariza-
Taste Coding in the Brain
tion of the membrane (Shirley & Persaud, 1990). Although you may assume that the five kinds of recep-
Sweetness, bitterness, and umami receptors have tors imply five labeled lines to the brain, research sug-
much in common chemically (He et al., 2004). After a gests a more complicated system (Hettinger & Frank,
molecule binds to one of these receptors, it activates a 1992). The receptors converge their input onto the
G-protein that releases a second messenger within the next cells in the taste system, each of which responds
cell, as in the metabotropic synapses discussed in Chap- best to a particular taste, but somewhat to other tastes
ter 3 (Lindemann, 1996). also. The brain can determine what the tongue is tast-
Bitter sensations have long been a puzzle. It is easy ing only by comparing the responses of several kinds
to describe the chemicals that produce a sour taste of taste neurons. In other words, taste depends on
(acids), a salty taste (Na+ ions), or umami (glutamate). a pattern of responses across fibers (R. P. Erickson,
Sweets are more difficult, as they include naturally DiLorenzo, & Woodbury, 1994).
occurring sugars plus artificial chemicals such as sac- Information from the receptors in the anterior two-
charin and aspartame. But bitter substances include a thirds of the tongue is carried to the brain along the
long list of chemicals that apparently have nothing in chorda tympani, a branch of the seventh cranial nerve
common, except that they are to varying degrees toxic. (the facial nerve). Taste information from the poste-
What receptor could identify such a large and diverse rior tongue and the throat is carried along branches of
set of chemicals? The answer is that we have not one the ninth and tenth cranial nerves. What do you sup-
bitter receptor but a whole family of 40 or more (Adler pose would happen if someone anesthetized your
et al., 2000; Matsunami, Montmayeur, & Buck, 2000). chorda tympani? You would no longer taste anything
Most taste cells sensitive to bitterness contain just a in the anterior part of your tongue, but you probably
small number of the possible bitter receptors, not all would not notice because you would still taste with the
40 (Caicedo & Roper, 2001). posterior part. However, the probability is about 40%
One consequence of having so many bitter recep- that you would experience taste phantoms, analo-
tors is that we can detect a great variety of dangerous gous to the phantom limb experience discussed in
chemicals. The other is that because each type of bitter Chapter 5 (Yanagisawa, Bartoshuk, Catalanotto, Kar-
receptor is present in small numbers, we cant detect rer, & Kveton, 1998). That is, you might experience
very low concentrations of bitter substances, the way tastes even when nothing was on your tongue. Evi-
we can with sour or salty substances, for example. dently, the inputs from the anterior and posterior parts
Somatosensory cortex
Ventral
posteromedial
thalamus
Insula (primary taste cortex) Corpus
callosum
Orbital
prefrontal
cortex
Hypothalamus
Amygdala
Nucleus of tractus solitarius
From taste buds on tongue
Figure 7.19 Major routes of impulses related to the sense of taste in the human brain
The thalamus and cerebral cortex receive impulses from both the left and the right sides of the
tongue. (Source: Based on Rolls, 1995)
The variations in taste sensitivity relate to the num- If you would like to classify yourself
ber of fungiform papillae near the tip of the tongue. as a taster, nontaster, or supertaster, follow try it
Supertasters have the most; nontasters have the few- the instructions in Table 7.2. yourself
est. That anatomical difference depends mostly on ge-
netics but also on hormones and other influences.
Womens taste sensitivity rises and falls with their
monthly hormone cycles and reaches its maximum
STOP & CHECK
during early pregnancy, when estradiol levels are very
high (Prutkin et al., 2000). That tendency is probably 5. What are several reasons why some people like
adaptive: During pregnancy, a woman needs to be more spicier foods than others do?
careful than usual to avoid harmful foods.
Check your answer on page 227.
Olfactory
bulb
Olfactory
nerve
(a)
Olfactory
bulb
Olfactory
nerve axons
Olfactory
receptor cell
Olfactory
epithelium Supporting cell
Olfactory
cilia (dendrites)
(b)
Terms
across-fiber pattern principle labeled-line principle (p. 215) place theory (p. 198)
(p. 215) loudness (p. 196) placebo (p. 212)
adaptation (p. 217) nerve deafness (inner-ear primary auditory cortex (area A1)
amplitude (p. 196) deafness) (p. 201) (p. 199)
anosmia (p. 222) nucleus of the tractus solitarius semicircular canal (p. 205)
capsaicin (p. 209) (NTS) (p. 219) somatosensory system (p. 206)
cochlea (p. 198) olfaction (p. 220) specific anosmia (p. 222)
conductive deafness (middle-ear olfactory cell (p. 221) substance P (p. 209)
deafness) (p. 201) opioid mechanisms (p. 210) supertasters (p. 219)
cross-adaptation (p. 217) oval window (p. 197) synesthesia (p. 225)
dermatome (p. 208) Pacinian corpuscle (p. 206) taste bud (p. 216)
endorphin (p. 210) papilla (pl.: papillae) (p. 216) tinnitus (p. 202)
frequency (p. 196) periaqueductal gray area (p. 210) tympanic membrane (p. 197)
frequency theory (p. 198) pheromone (p. 224) volley principle (p. 199)
gate theory (p. 211) pinna (p. 197) vomeronasal organ (VNO) (p. 224)
hair cell (p. 198) pitch (p. 196)
B
The Cerebellum
The Basal Ganglia efore we get started, please try this: Get out a
Brain Areas and Motor Learning pencil and a sheet of paper, and put the pencil
In Closing: Movement Control and Cognition in your nonpreferred hand. For example, if you are
Summary right-handed, put it in your left hand. Now, with that
Answers to Stop & Check Questions hand, draw a face in profilethat is, facing
Thought Question one direction or the other, but not straight try it
ahead. Please do this now before reading yourself
Module 8.3 further.
Disorders of Movement If you tried the demonstration, you probably notice
Parkinsons Disease that your drawing is much more childlike than usual.
Huntingtons Disease It is as if some part of your brain stored the way you
In Closing: Heredity and Environment used to draw as a young child. Now, if you are right-
in Movement Disorders handed and therefore drew the face with your left hand,
Summary why did you draw it facing to the right? At least I as-
Answers to Stop & Check Questions sume you did because more than two-thirds of right-
Thought Questions handers drawing with their left hand draw the profile
facing right. Young children, age 5 or so, when drawing
Terms with the right hand, almost always draw people and
Suggestions for Further Reading animals facing left, but when using the left hand, they
Websites to Explore almost always draw them facing right. But why? The
Exploring Biological Psychology CD short answer is we dont know. We have much to learn
ThomsonNOW about the control of movement and how it relates to
perception, motivation, and other functions.
231
Module 8.1
The Control of Movement
Biceps Triceps
contracts relaxes
Biceps Triceps
relaxes contracts
Ed Reschke
Figure 8.2
An axon branching to innervate separate muscle
fibers within a muscle
Movements can be much more precise where each axon Figure 8.3 A pair of antagonistic muscles
innervates only a few fibers, as with eye muscles, than where The biceps of the arm is a flexor; the triceps is an extensor.
it innervates many fibers, as with biceps muscles. (Source: Starr & Taggart, 1989)
3. Duck breast muscles are red (dark meat) whereas Muscle spindle
chicken breast muscles are white. Which species Golgi tendon organ
probably can fly for a longer time before fatiguing?
4. Why is an ultramarathoner like Bertil Jrlaker probably
mediocre or poor at short-distance races?
Check your answers on page 239.
Units of Movement
The stretch reflex is a simple example of movement.
More complex kinds include speaking, walking, thread-
ing a needle, and throwing a basketball through a hoop
while off balance and evading two defenders. In many
ways, these movements are different from one another,
Figure 8.6 The knee-jerk reflex and they depend on different kinds of control by the
Here is one example of a stretch reflex.
nervous system.
Voluntary and
the lower leg upward. The same reflex contributes to Involuntary Movements
walking; raising the upper leg reflexively moves the
lower leg forward in readiness for the next step. Reflexes are consistent automatic responses to stimuli.
The Golgi tendon organ, another proprioceptor, We generally think of reflexes as involuntary because
responds to increases in muscle tension. Located in they are insensitive to reinforcements, punishments,
the tendons at opposite ends of a muscle, it acts as a and motivations. The stretch reflex is one example;
brake against an excessively vigorous contraction. Some another is the constriction of the pupil in response to
muscles are so strong that they could damage them- bright light.
selves if too many fibers contracted at once. Golgi ten-
don organs detect the tension that results during a E X T E N S I O N S A N D A P P L I C AT I O N S
muscle contraction. Their impulses travel to the spinal
Infant Reflexes
cord, where they inhibit the motor neurons through
messages from interneurons. In short, a vigorous mus- Humans have few reflexes, although infants have sev-
cle contraction inhibits further contraction by activat- eral not seen in adults. For example, if you place an
ing the Golgi tendon organs. object firmly in an infants hand, the infant will reflex-
The proprioceptors not only control important re- ively grasp it tightly (the grasp reflex). If you stroke the
flexes but also provide the brain with information. Here sole of the foot, the infant will reflexively extend the
is an illusion that you can demonstrate yourself: Find big toe and fan the others (the Babinski reflex). If you
a small, dense object and a larger, less dense object touch an infants cheek, the head will turn toward the
that weighs the same as the small one. For example, stimulated cheek, and the infant will begin to suck (the
you might try a lemon and a hollowed-out orange, rooting reflex). The rooting reflex is not a pure reflex,
with the peel pasted back together so it appears to be as its intensity depends on the infants arousal and
intact. Drop one of the objects onto someones hand hunger levels.
while he or she is watching. (The watching is essen- Although such reflexes fade away with time, the
tial.) Then remove it and drop the other object onto connections remain intact, not lost but suppressed by
the same hand. Most people report that the small one axons from the maturing brain. If the cerebral cortex
felt heavier. The reason is that with the larger object, is damaged, the infant reflexes are released from inhi-
people set themselves up with the expectation of a bition. In fact, neurologists and other physicians fre-
heavier weight. The actual weight displaces quently test adults for infant reflexes. A physician who
their proprioceptors less than expected and try it strokes the sole of your foot during a physical exam is
therefore yields the perception of a lighter yourself probably looking for evidence of brain damage. This
object. is hardly the most dependable test, but it is easy. If a
Jo Ellen Kalat
Charles Gupton/Stock, Boston, Inc./Picture Quest
The grasp reflex enables an infant to cling to the mother
while she travels.
Trunk
lder
Arm
ow
Shou
Since the pioneering work of Gustav Fritsch and Ed-
Elb
t
ris
uard Hitzig (1870), neuroscientists have known that
nd
Ha
direct electrical stimulation of the primary motor cor-
s
er
b
ng
texthe precentral gyrus of the frontal cortex, just an- um k
Fi
Th Nec
terior to the central sulcus (Figure 8.8)elicits move- w
Bro ye
ments. The motor cortex has no direct connections to E
the muscles; its axons extend to the brainstem and Toes Face
spinal cord, which generate the activity patterns that Lips
control the muscles (Shik & Orlovsky, 1976). The cere-
bral cortex is particularly important for complex ac- Jaw
tions such as talking, writing, or hand gestures. It is less
important for coughing, sneezing, gagging, laughing, Tongue
or crying (Rinn, 1984). Perhaps the lack of cerebral con- Swall
owin
trol explains why it is hard to perform such actions g
voluntarily. Laughing or coughing voluntarily is not
the same as laughing or coughing spontaneously, and
most people cannot cry or sneeze voluntarily.
Figure 8.9 (which repeats part of Figure 4.24,
p. 99) indicates which area of the motor cortex con- Figure 8.9 Coronal section through the primary
trols which area of the body. For example, the brain motor cortex
area shown next to the hand is active during hand Stimulation at any point in the primary motor cortex is most
movements. In each case, the brain area controls a likely to evoke movements in the body area shown. However,
structure on the opposite side of the body. However, actual results are usually messier than this figure implies:
dont read this figure as implying that each spot in the For example, individual cells controlling one finger may be
motor cortex controls a single muscle. For example, intermingled with cells controlling another finger. (Source:
movement of any finger or the wrist is associated with Adapted from Penfield & Rasmussen, 1950)
hand had been. That is, the stimulation produced a firmed that the finger area of the motor cortex is active
certain outcome, not a fixed set of muscle movements. when pianists listen to familiar music, even if they keep
Depending on the position of the arm, the stimulation their fingers motionless (Haueisen & Knsche, 2001).
might activate biceps muscles, triceps, or whatever. Neurons in part of the inferior parietal cortex of
Although the motor cortex can direct contractions of monkeys are active during a movement and while a
a specific muscle, more often it orders an outcome and monkey watches another monkey do the same move-
leaves it to the spinal cord and other areas to find the ment (Cisek & Kalaska, 2004; Fogassi et al., 2005).
combination of contractions to produce that outcome Brain-scan studies have demonstrated similar neurons
(S. H. Scott, 2004). in humans. These neurons, called mirror neurons, pre-
Just as the visual cortex becomes active when we sumably enable the observer to understand and iden-
imagine seeing something, the motor cortex becomes tify with the movements another individual is making.
active when we imagine movements. For example, ex- They respond when a monkey watches another mon-
pert pianists say that when they listen to familiar, well- key or a human watches another human. They do not
practiced music, they imagine the finger movements respond when a monkey watches a human do some-
and often start tapping the appropriate fingers as if they thing or when a human watches a robot (Tai, Scherfler,
were playing the music. Brain recordings have con- Brooks, Sawamoto, & Castiello, 2004). Evidently, there-
Paralysis Lack of voluntary movement in part of the body. Damage to spinal cord, motor neurons, or their axons.
Paraplegia Loss of sensation and voluntary muscle control Cut through the spinal cord above the segments
in both legs. Reflexes remain. Although no attached to the legs.
messages pass between the brain and the
genitals, the genitals still respond reflexively to
touch. Paraplegics have no genital sensations, but
they can still experience orgasm (Money, 1967).
Quadriplegia Loss of sensation and muscle control in all four Cut through the spinal cord above the segments
extremities. controlling the arms.
Hemiplegia Loss of sensation and muscle control in the arm Cut halfway through the spinal cord or (more
and leg on one side. commonly) damage to one hemisphere of the
cerebral cortex.
Tabes dorsalis Impaired sensation in the legs and pelvic region, Late stage of syphilis. Dorsal roots of the spinal cord
impaired leg reflexes and walking, loss of bladder deteriorate.
and bowel control.
Poliomyelitis Paralysis. Virus that damages cell bodies of motor neurons.
Amyotrophic Gradual weakness and paralysis, starting with the Unknown.
lateral sclerosis arms and later spreading to the legs. Both motor
neurons and axons from the brain to the motor
neurons are destroyed.
fore, their response reflects identification of the viewer but all vertebrates have this pattern. In newborn hu-
with the actor: The actor is like me, doing something mans, the immature primary motor cortex has partial
I might do. Many psychologists believe that the exis- control of both ipsilateral and contralateral muscles.
tence of such neurons is important for the complex so- As the contralateral control improves over the first year
cial behaviors typical of humans and other primates. and a half of life, it displaces the ipsilateral control,
which gradually becomes weaker. In some children
Connections from the with cerebral palsy, the contralateral path fails to ma-
ture, and the ipsilateral path remains relatively strong.
Brain to the Spinal Cord In fact, sometimes part of the clumsiness of children
All the messages from the brain must eventually reach with cerebral palsy comes from competition between
the medulla and spinal cord, which control the mus- the ipsilateral and contralateral paths (Eyre, Taylor,
cles. Diseases of the spinal cord can impair the control Villagra, Smith, & Miller, 2001).
of movement in various ways (Table 8.1). The various In contrast to the dorsolateral tract, the ventro-
axons from the brain organize into two paths, the dorso- medial tract includes axons from the primary motor
lateral tract and the ventromedial tract. Nearly all move- cortex, surrounding areas, and also from many other
ments rely on a combination of both the dorsolateral parts of the cortex. In addition, it includes axons that
and ventromedial tracts, but many movements rely on originate from the midbrain tectum, the reticular for-
one tract more than the other. mation, and the vestibular nucleus, a brain area that
The dorsolateral tract of the spinal cord is a set receives input from the vestibular system (see Fig-
of axons from the primary motor cortex, surrounding ure 8.11). Axons of the ventromedial tract go to both
areas, and the red nucleus, a midbrain area with out- sides of the spinal cord, not just the contralateral side.
put mainly to the arm muscles (Figure 8.11). Axons of The ventromedial tract controls mainly the muscles
the dorsolateral tract extend directly from the motor of the neck, shoulders, and trunk and therefore such
cortex to their target neurons in the spinal cord. In movements such as walking, turning, bending, stand-
bulges of the medulla called pyramids, the dorsolateral ing up, and sitting down (Kuypers, 1989). Note that
tract crosses from one side of the brain to the contra- these movements are necessarily bilateral; you can
lateral (opposite) side of the spinal cord. (For that rea- move your fingers on just one side, but any movement
son, the dorsolateral tract is also called the pyramidal of your neck or trunk must include both sides.
tract.) It controls movements in peripheral areas, such Suppose someone has suffered a stroke that dam-
as the hands, fingers, and toes. aged the primary motor cortex of the left hemisphere.
Why does each hemisphere control the contra- The result is a loss of the dorsolateral tract from that
lateral side instead of its own side? We do not know, hemisphere and a loss of movement control on the right
side of the body. Eventually, depending on the exact Areas Near the Primary Motor Cortex
location and amount of damage, the person may regain
some muscle control from spared axons in the dorso- A number of areas near the primary motor cortex also
lateral tract. If not, the possibility remains of using the contribute to movement in diverse ways (see Figure
ventromedial tract to approximate the intended move- 8.8). In the posterior parietal cortex, some neurons
ment. For example, someone who has no direct con- respond primarily to visual or somatosensory stimuli,
trol of the hand muscles might move the shoulders, others respond mostly to current or future movements,
trunk, and hips in a way that repositions the hand in a and still others respond to a complicated mixture of
crude way. Also, because of connections between the the stimulus and the upcoming response (Shadlen &
left and right halves of the spinal cord, normal move- Newsome, 1996). You might think of the posterior
ments of one arm or leg can induce associated move- parietal cortex as keeping track of the position of the
ments on the other side, at least to a limited degree body relative to the world (Snyder, Grieve, Brotchie, &
(Edgley, Jankowska, & Hammar, 2004). Andersen, 1998). Contrast the effects of posterior pari-
etal damage with those of occipital or temporal dam-
age. People with posterior parietal damage can accu-
rately describe what they see, but they have trouble
STOP & CHECK converting their perception into action. Although they
can walk toward something they hear, they cannot walk
toward something they see, nor can they reach out to
1. What evidence indicates that cortical activity repre-
grasp somethingeven after describing its size, shape,
sents the idea of the movement and not just the
and angle. They seem to know what it is but not where
muscle contractions?
it is. In contrast, people with damage to parts of the oc-
2. What kinds of movements does the dorsolateral tract cipital or temporal cortex have trouble describing what
control? The ventromedial tract? they see, but they can reach out and pick up objects,
Check your answers on page 252. and when walking, they step over or go around the
objects in their way (Goodale, 1996; Goodale, Milner,
55 5
Person reports that the
conscious decision
50 occurred here.
10
Brains readiness
potential begins to
rise in preparation The movement
for the movement. itself starts here.
45 15
Cerebellum Nuclei
E
E nial nerve nuclei, and from the cerebral cortex. That
they saw a signal telling where the letter would be on
the screen. For most people, that signal improved their information eventually reaches the cerebellar cortex,
performance even if it appeared just 100 ms before the the surface of the cerebellum (see Figure 8.14).
letter. For people with cerebellar damage, the signal Figure 8.15 shows the types and arrangements of
had to appear nearly a second before the letter to be neurons in the cerebellar cortex. The figure is com-
helpful. Evidently, people with cerebellar damage need plex, but concentrate on these main points:
longer to shift their attention (Townsend et al., 1999). The neurons are arranged in a precise geometrical
So the cerebellum appears to be linked to timing, pattern, with multiple repetitions of the same units.
certain aspects of attention, and probably other abili-
ties as well. Are these separate functions that just hap-
pen to be located in the same place? Or can we reduce
them all to a single theme? (For example, maybe shift-
ing attention requires
timing or aim.) These
unanswered ques- Stellate cell
tions require a care-
ful analysis of be-
havior, not just a
study of the ner-
vous system.
Parallel fibers
Purkinje cell
Basket cell
Substantia
nigra
Summary
1. The primary motor cortex is the main source of
Answers to
brain input to the spinal cord. The spinal cord con- STOP & CHECK
tains central pattern generators that actually control
the muscles. (p. 241)
Questions
2. The primary motor cortex produces patterns rep- 1. Activity in the motor cortex leads to a particular
resenting the intended outcome, not just the mus- outcome, such as movement of the hand to the
cle contractions. Neurons in part of the parietal mouth, regardless of what muscle contractions are
cortex respond to both a self-produced movement necessary given the hands current location. Also,
and an observation of a similar movement by an- neurons in part of the parietal cortex respond sim-
other individual. (p. 242) ilarly to self-produced movement and to observa-
3. The dorsolateral tract, which controls movements tion of a similar movement by another individual.
in the periphery of the body, has axons that cross (p. 244)
from one side of the brain to the opposite side of 2. The dorsolateral tract controls detailed movements
the spinal cord. (p. 243) in the periphery on the contralateral side of the
4. The ventromedial tract controls bilateral move- body. (For example, the dorsolateral tract from the
ments near the midline of the body. (p. 243) left hemisphere controls the right side of the body.)
The ventromedial tract controls the trunk muscles
5. Areas near the primary motor cortexincluding
bilaterally. (p. 244)
the prefrontal, premotor, and supplementary motor
corticesare active in detecting stimuli for move- 3. The posterior parietal cortex is important for per-
ment and preparing for a movement. (p. 245) ceiving the location of objects and the position of
the body relative to the environment, including
6. When people identify the instant when they formed
those objects. The prefrontal cortex responds to
a conscious intention to move, their time precedes
sensory stimuli that call for some movement. The
the actual movement by about 200 ms but follows
premotor cortex is active in preparing a movement
the start of motor cortex activity by about 300 ms.
immediately before it occurs. The supplementary
These results suggest that what we call a conscious
motor cortex is especially active in preparing for
decision is our perception of a process already un-
a rapid sequence of movements. (p. 245)
derway, not really the cause of it. (p. 245)
4. Researchers recorded a readiness potential in the
7. People with damage to part of the parietal cortex
motor cortex, beginning about 500 ms before the
fail to perceive any intention prior to the start of
start of the movement. When people reported
their own movements. (p. 247)
the time that they felt the intention to move, the
8. Some people with damage to the primary motor reported intention occurred about 200 ms before
cortex of the right hemisphere are paralyzed on the movement and therefore 300 ms after the start
the left side but insist that they can still move. Evi- of motor cortex activity that led to the movement.
dently, they fail to receive the feedback that indi- (p. 247)
cates lack of movement. (p. 247)
5. After damage to the parietal cortex, people do not
9. The cerebellum has multiple roles in behavior, in- monitor the processes leading up to a movement.
cluding sensory functions related to perception of When they try to report the time of an intention to
the timing or rhythm of stimuli. Its role in the con- move, they report the same time when the move-
trol of movement is especially important for tim- ment actually began. That is, they are not aware of
ing, aiming, and correcting errors. (p. 247) any intention before the movement itself. (p. 247)
10. The cells of the cerebellum are arranged in a very 6. Anosognosia occurs when someone is paralyzed
regular pattern that enables them to produce out- on the left side but denies having such a problem.
puts of well-controlled duration. (p. 249) It is associated with damage to the motor cortex of
11. The basal ganglia are a group of large subcortical the right hemisphere and parts of the surrounding
structures that are important for selecting and in- areas. (p. 247)
Thalamus Thalamus
Amygdala Amygdala
Putamen Putamen
(a) (b)
Figure 8.17 Connections from the substantia nigra: (a) normal and
(b) in Parkinsons disease
Excitatory paths are shown in green; inhibitory are in red. The substantia nigras axons
inhibit the putamen. Axon loss increases excitatory communication to the globus pallidus.
The result is increased inhibition from the globus pallidus to the thalamus and decreased
excitation from the thalamus to the cerebral cortex. People with Parkinsons disease show
decreased initiation of movement, slow and inaccurate movement, and psychological
depression. (Source: Based on Wichmann, Vitek, & DeLong, 1995)
Chapter Ending
Key Terms and Activities
Terms
aerobic (p. 234) grasp reflex (p. 236) primary motor cortex (p. 241)
anaerobic (p. 234) huntingtin (p. 260) proprioceptor (p. 235)
anosognosia (p. 247) Huntingtons disease (p. 258) Purkinje cell (p. 250)
antagonistic muscles (p. 232) L-dopa (p. 257) readiness potential (p. 246)
Babinski reflex (p. 236) mirror neurons (p. 242) red nucleus (p. 243)
ballistic movement (p. 238) motor program (p. 238) reflex (p. 236)
basal ganglia (caudate nucleus, MPTP, MPP+ (p. 256) rooting reflex (p. 236)
putamen, globus pallidus) muscle spindle (p. 235) skeletal muscle (or striated
(p. 250) muscle) (p. 232)
myasthenia gravis (p. 234)
cardiac muscle (p. 232) slow-twitch fiber (p. 234)
neuromuscular junction (p. 232)
central pattern generator (p. 238) smooth muscle (p. 232)
nuclei of the cerebellum (p. 250)
cerebellar cortex (p. 249) stem cell (p. 258)
parallel fibers (p. 250)
dorsolateral tract (p. 243) stretch reflex (p. 235)
Parkinsons disease (p. 254)
extensor (p. 232) supplementary motor cortex
posterior parietal cortex (p. 244)
fast-twitch fiber (p. 234) (p. 245)
prefrontal cortex (p. 245)
flexor (p. 232) ventromedial tract (p. 243)
premotor cortex (p. 245)
Golgi tendon organ (p. 236) vestibular nucleus (p. 243)
presymptomatic test (p. 259)
Cole, J. (1995). Pride and a daily marathon. Cam- Go to this site for the link to ThomsonNOW, your one-stop study
bridge, MA: MIT Press. Biography of a man who shop, Take a Pre-Test for this chapter, and ThomsonNOW will
lost his sense of touch and proprioception from generate a Personalized Study Plan based on your test results.
the neck down and eventually learned to control The Study Plan will identify the topics you need to review and
his movements strictly by vision. direct you to online resources to help you master these topics.
You can then take a Post-Test to help you determine the con-
Klawans, H. L. (1996). Why Michael couldnt hit. New
cepts you have mastered and what you still need work on.
York: W. H. Freeman. A collection of fascinat-
ing sports examples related to the brain and its
disorders.
Lashley, K. S. (1951). The problem of serial order in
behavior. In L. A. Jeffress (Ed.), Cerebral mecha-
nisms in behavior (pp. 112136). New York: Wiley.
One of the true classic articles in psychology; a
thought-provoking appraisal of what a theory of
movement should explain.
Websites
to Explore
You can go to the Biological Psychology Study
Center and click these links. While there, you
can also check for suggested articles available View an animation of the dorsolateral and ventromedial tracts.
on InfoTrac College Edition. The Biological Psychol-
ogy Internet address is:
http://psychology.wadsworth.com/book/kalatbiopsych9e/
Exploring Biological
Psychology CD
This animation illustrates one example of a reflex.
The Withdrawal Reflex (animation)
The Crossed Extensor Reflex (animation)
Major Motor Areas (animation)
Critical Thinking (essay questions)
Chapter Quiz (multiple-choice questions)
265
Module 9.1
Rhythms of Waking
and Sleeping
5
Time of the middle of sleep on
Females
days without obligations
10 20 30 40 50 60
Age (years)
Duration of the Human follow. Researchers had already known that most peo-
ple can adjust to a 23- or 25-hour day but not to a 22-
Circadian Rhythm or 28-hour day (Folkard, Hume, Minors, Waterhouse,
It might seem simple to determine the duration of the & Watson, 1985; Kleitman, 1963). Later researchers
human circadian rhythm: Put people in an environ- kept healthy adults in rooms with an artificial 28-hour
ment with no cues to time and observe their waking day. None of them could, in fact, synchronize to that
sleeping schedule. However, the results depend on schedule; they all therefore produced their own self-
the amount of light (Campbell, 2000). Under constant generated rhythms of alertness and body temperature.
bright lights, people have trouble sleeping, they com- Those rhythms varied among individuals, with a mean
plain about the experiment, and their rhythms run of 24.2 hours (Czeisler et al., 1999).
faster than 24 hours. In constant darkness, they have Yet another approach is to examine people living
trouble waking up, again they complain about the ex- under unusual conditions. Naval personnel on U.S.
periment, and their rhythms run slower. In several stud- nuclear powered submarines are cut off from sunlight
ies, people were allowed to turn on bright lights when- for months at a time, living under faint artificial light.
ever they chose to be awake and turn them off when In many cases, they have been asked to live on a sched-
they wanted to sleep. Under these conditions, most peo- ule of 6 hours of work alternating with 12 hours of rest.
ple followed a cycle closer to 25 than to 24 hours a day. Even though they sleep (or try to sleep) on this 18-hour
The problem, which experimenters did not realize at schedule, their bodies generate rhythms of alertness
first, was that bright light late in the day lengthens the and body chemistry that average about 24.3 to 24.4
circadian rhythm. hours (Kelly et al., 1999). In short, humans circadian
A different way to run the experiment is to pro- clock generates a rhythm slightly longer than 24 hours
vide light and darkness on a cycle that people cannot when it has nothing to reset it.
Hypothalamus
(c)
Day Night
Concentration
of Tim and Per Normal
Pulse of
bright
light Bright light
late at night
Concentration
phase
of Tim and Per
advances
the rhythm
Behavior Wakefulness Sleep Wake
Module 9.1
In Closing: SleepWake Cycles
Unlike an electric appliance that stays on until some-
one turns it off, the brain periodically turns itself on
Eviatar Nevo
REM
REM
REM
REM
A23 4 32 2 3 432 3 4 2 2 3 2 2 2 A Columns indicate awake (A) and
sleep stages 2, 3, 4, and REM.
Deflections in the line at the
bottom of each chart indicate
11 P.M. 12 P.M. 1 A.M. 2 A.M. 3 A.M. 4 A.M. 5 A.M. shifts in body position. Note that
stage 4 sleep occurs mostly in
the early part of the nights sleep,
REM
REM
REM
A2 3 4 2 3 4 32 3 2 2 3 2 A whereas REM sleep becomes more
prevalent toward the end. (Source:
Based on Dement & Kleitman, 1957a)
REM
REM
REM
A 2 34 2 4 3 4 2 2 3 4 2 2 3 4 2 2 3 2A 2 2 A
REM than in other stages; in this regard, REM is deep William Dement and Nathaniel Kleitman (1957b) found
sleep. REM is also associated with erections in males that people who were awakened during REM sleep re-
and vaginal moistening in females. Heart rate, blood ported dreams 80% to 90% of the time. Later research-
pressure, and breathing rate are more variable in REM ers, however, found that people also sometimes re-
than in stages 2 through 4. In short, REM sleep com- ported dreams when they were awakened from NREM
bines deep sleep, light sleep, and features that are dif- sleep. REM dreams are more likely than NREM dreams
ficult to classify as deep or light. Consequently, it is to include striking visual imagery and complicated
best to avoid the terms deep and light sleep. plots, but not always. Some people with brain dam-
In addition to its steady characteristics, REM sleep age continue to have REM sleep but do not report any
has intermittent characteristics such as facial twitches dreams (Bischof & Bassetti, 2004), and other people
and eye movements, as shown in Figure 9.9(f). The EEG continue to report dreams despite no evidence of REM
record is similar to that for stage 1 sleep, but notice the sleep (Solms, 1997). In short, REM and dreams usu-
difference in eye movements. The stages other than ally overlap, but they are not the same thing.
REM are known as non-REM (NREM) sleep.
Anyone who falls asleep first enters stage 1 and
then slowly progresses through stages 2, 3, and 4 in
order, although loud noises or other intrusions can in-
STOP & CHECK
terrupt this sequence. After about an hour of sleep, the
person begins to cycle back from stage 4 through stages 2. How can an investigator determine whether a sleeper
3, 2, and then REM. The sequence repeats, with each is in REM sleep?
complete cycle lasting about 90 minutes. Early in the
3. During which part of a nights sleep is REM most
night, stages 3 and 4 predominate. Toward morning,
common?
the duration of stage 4 grows shorter and the duration
of REM grows longer. Figure 9.10 shows typical se- Check your answers on page 285.
quences. The tendency to increase REM depends on
time, not on how long you have been asleep. That is, if
you go to sleep later than usual, you still begin to in-
crease your REM at about the same time that you would
have ordinarily (Czeisler, Weitzman, Moore-Ede, Zim- Brain Mechanisms of
merman, & Knauer, 1980). Most depressed people enter Wakefulness and Arousal
REM quickly after falling asleep, even at their normal
time, suggesting that their circadian rhythm is out of Recall from Chapter 1 the distinction between the
synchrony with real time. easy and hard problems of consciousness. The
Initially after the discovery of REM, researchers easy problems include such matters as, Which brain
believed it was almost synonymous with dreaming. areas increase overall alertness, and by what kinds of
GABA Ac
et
ylc
ho
Histamine
li n
e
BA
GA
Basal forebrain
Hista Hypothalamus
Ac
G AB Norepine
ine
y lc ph
et
A
ine
ho r
li n e
Adenos
in
Se
e
r Locus coeruleus
lcholine
ot
A c ety
on
Hista
in
mi
ne
Hi
sta
m in e
Dorsal raphe
Pontomesencephalon
tial for sleep; that is, without the inhibition provided usual rate, and neurons in the brains sensory areas con-
by GABA, sleep would not occur (Gottesmann, 2004). tinue to respond to sounds and other stimuli. Neverthe-
The functions of GABA help explain what we experi- less, we are unconscious. An explanation is that GABA
ence during sleep: During sleep, body temperature and inhibits synaptic activity. A neuron may be active, ei-
metabolic rate decrease slightly, and so does the activ- ther spontaneously or in response to a stimulus, but its
ity of neurons, but by less than we might expect. Spon- axons do not spread the stimulation to other areas be-
taneously active neurons continue to fire at almost their cause of inhibition by GABA. Researchers demonstrated
Getting to Sleep
Sleep requires decreased arousal, largely by means of
adenosine (ah-DENN-o-seen). During metabolic activ-
ity, adenosine monophosphate (AMP) breaks down into
adenosine; thus, when the brain is awake and active,
adenosine accumulates. In most of the brain, adeno-
sine has little effect, but adenosine inhibits the basal
forebrain cells responsible for arousal (Figure 9.12),
acting by metabotropic synapses that produce an effect
Basal forebrain
lasting hours (Basheer, Rainnie, Porkka-Heiskanen,
Ramesh, & McCarley, 2001). When people are deprived
of sleep, the accumulating adenosine produces pro-
longed sleepinessa phenomenon known as sleep
debt.
Figure 9.12 Basal forebrain
Caffeine, a drug found in coffee, tea, and many soft
The basal forebrain is the source of many excitatory axons
drinks, increases arousal by blocking adenosine re-
(releasing acetylcholine) and inhibitory axons (releasing
ceptors (Rainnie, Grunze, McCarley, & Greene, 1994).
GABA) that regulate arousal of the cerebral cortex.
It also constricts the blood vessels in the brain, thereby
decreasing its blood supply. (Abstention from caffeine
after repeated use can increase blood flow to the brain
enough to cause a headache.) The message: Just as you
might use caffeine to try to keep yourself awake, you
might try decreasing your caffeine intake if you have
STOP & CHECK
trouble sleeping.
Prostaglandins are additional chemicals that pro- 4. What would happen to the sleepwake schedule of
mote sleep, among other functions. Like adenosine, someone who took a drug that blocked GABA?
prostaglandins build up during the day until they pro- 5. Why do most antihistamines make people drowsy?
voke sleep, and they decline during sleep (Ram et al.,
6. What would happen to the sleepwake schedule of
1997; Scamell et al., 1998). In response to infection, the
someone who lacked orexin?
immune system produces more prostaglandins, result-
ing in the sleepiness that accompanies illness. 7. How does caffeine increase arousal?
Table 9.1 summarizes the effects of some key brain Check your answers on page 285.
areas on arousal and sleep.
Geniculate
P
Pons
Sleep period
3-hr
Difficulty
Sleep Disorders phase advance
staying asleep
Sleep Apnea
Alan Williams/Alamy
could quit if necessary. (Stressors take a greater toll
when they are unpredictable and uncontrollable.) Dur-
ing a few days of sleep deprivation, rats show increased
body temperature, metabolic rate, and appetite, indi-
A European swift. cating that the body is working harder than usual. With
still longer sleep deprivation, the immune system be-
gins to fail, the animal loses its resistance to infection,
It picks an altitude where the air is not too cold, accepts and brain activity decreases (Everson, 1995; Recht-
the risk of being blown far from home, and awakens schaffen & Bergmann, 1995). However, it is difficult to
the next morning to resume its chase of flying insects separate the effects of sleep deprivation from those of
(Bckman & Alerstam, 2001). Perhaps it, like dolphins, the various kinds of prods that were necessary to keep
sleeps on one side of its brain at a time, but we wont the animals awake.
know until someone figures out how to measure the
EEG of a bird in flight.
Sleep and Memory
When people learn something and then get tested the
next day, their performance is often better the second
STOP & CHECK day than the first, but only if they get adequate sleep
during the night (Stickgold, James, & Hobson, 2000;
1. Some fish live in caves or the deep ocean with no Stickgold, Whidbee, Schirmer, Patel, & Hobson, 2000).
light. What might one predict about their sleep? (The obvious message to students: When you are study-
ing, get enough sleep.) When people practice some-
Check your answer on page 292.
thing repeatedly, sometimes their performance deteri-
orates during the day, but it recovers if they get a short
nap (Mednick et al., 2002). These results imply that
sleep enhances memory. Sleep also helps people re-
Restorative Functions of Sleep analyze their memories: In one study, people who had
Even if the original function of sleep was to conserve just practiced a complex task were more likely to per-
energy, sleep undeniably serves additional functions ceive a hidden rule (an aha experience) after a period
today. One way to examine the restorative functions of sleep than after a similar period of wakefulness (Wag-
is to observe the effects of sleep deprivation. People ner, Gais, Haider, Verleger, & Born, 2004).
who have gone without sleep for a week or more, ei- In several studies, researchers recorded brain ac-
ther as an experiment or as a publicity stunt, have re- tivity as people learned a motor skillsimilar to the
ported dizziness, impaired concentration, irritability, skills you might learn in a video gameand then mon-
hand tremors, and hallucinations (Dement, 1972; L. C. itored brain activity during sleep. Later, they recorded
Johnson, 1969). Astronauts in space follow irregular brain activity while these people slept and found in-
work schedules, sleep in spacesuits under weightless- creased activity in the same areas that had been acti-
ness, and for a variety of other reasons have trouble vated while these participants had been learning the
sleeping as much as usual. On long trips, they experi- skill. Furthermore, the amount of activity in those
ence unpleasant mood, decreased alertness, and im- areas during sleep correlated highly with the improve-
paired performance (Mallis & DeRoshia, 2005). People ment in skill seen the next day (Huber, Ghilardi, Mas-
working during the winter in Antarctica sleep poorly simini, & Tononi, 2004; Maquet et al., 2000; Peigneux
and feel depressed (Palinkas, 2003). Even one night of et al., 2004). Similar results have been reported for
sleeplessness temporarily increases activity of the im- birds learning to sing (Dergnaucourt, Mitra, Fehr,
mune system (Matsumoto et al., 2001). That is, you Pytte, & Tchernichovski, 2005). All these results point
react to sleep deprivation as if you were ill. to sleep as a time when memories are strengthened.
Chapter Ending
Key Terms and Activities
Terms
activation-synthesis hypothesis endogenous circannual rhythm orexin (or hypocretin) (p. 278)
(p. 290) (p. 266) paradoxical sleep (p. 276)
adenosine (p. 280) free-running rhythm (p. 271) periodic limb movement disorder
alpha wave (p. 275) insomnia (p. 282) (p. 284)
basal forebrain (p. 278) jet lag (p. 272) PGO wave (p. 281)
caffeine (p. 280) K-complex (p. 275) pineal gland (p. 271)
cataplexy (p. 283) locus coeruleus (p. 278) polysomnograph (p. 275)
clinico-anatomical hypothesis melatonin (p. 271) pontomesencephalon (p. 278)
(p. 291) narcolepsy (p. 283) prostaglandin (p. 280)
endogenous circadian rhythm night terror (p. 284) rapid eye movement (REM) sleep
(p. 266) (p. 276)
non-REM (NREM) sleep (p. 277)
Suggestions for
Further Reading http://www.thomsonedu.com
Dement, W. C. (1992). The sleepwatchers. Stanford, Go to this site for the link to ThomsonNOW, your one-stop study
CA: Stanford Alumni Association. Fascinating, shop, Take a Pre-Test for this chapter, and ThomsonNOW will
entertaining account of sleep research by one of generate a Personalized Study Plan based on your test results.
its leading pioneers. The Study Plan will identify the topics you need to review and
direct you to online resources to help you master these topics.
Foster, R. G., & Kreitzman, L. (2004). Rhythms of life.
You can then take a Post-Test to help you determine the con-
New Haven, CT: Yale University Press. Nontech-
cepts you have mastered and what you still need work on.
nical discussion of research on circadian rhythms.
Moorcroft, W. H. (2003) Understanding sleep and
dreaming. New York: Kluwer. Excellent review of
research on many aspects of sleep and dreams.
Refinetti, R. (2005). Circadian physiology (2nd ed.).
Boca Raton, FL: CRC Press. Marvelous summary
of research on circadian rhythms and the rele-
vance to human behavior.
Websites
to Explore
You can go to the Biological Psychology Study
Center and click these links. While there, you
can also check for suggested articles available
on InfoTrac College Edition. The Biological Psychol- Here is a step-by-step view of the stages of sleep.
ogy Internet address is:
http://psychology.wadsworth.com/book/kalatbiopsych9e/
Exploring Biological
Psychology CD
Sleep Rhythms (learning module)
Sleep Cycle (video)
Explore the progression through the stages of sleep.
Critical Thinking (essay questions)
Chapter Quiz (multiple-choice questions)
W
Thought Questions
hat is life? Life can be defined in different ways
Module 10.3 depending on whether our interest is medical,
Hunger legal, philosophical, or poetic. Biologically, what is nec-
How the Digestive System Influences essary for life is a coordinated set of chemical reac-
Food Selection tions. Not all chemical reactions are alive, but all life
Short- and Long-Term Regulation of Feeding has precisely regulated chemical reactions.
Brain Mechanisms Every chemical reaction in the body takes place
Eating Disorders in a water solution at a rate that depends on the iden-
In Closing: The Multiple Controls of Hunger tity and concentration of molecules in the water, the
Summary temperature of the solution, and the presence of con-
Answers to Stop & Check Questions taminants. Much of our behavior is organized to keep
Thought Question the right chemicals in the right proportions and at the
right temperature.
Terms
Suggestions for Further Reading
Websites to Explore
Exploring Biological Psychology CD
ThomsonNOW
295
Module 10.1
Temperature Regulation
AP
has to recruit more and more fast-twitch muscle fibers
Companies will freeze a dead body with the prospect that
to remain active at the risk of rapid fatigue. Birds and
future technologies can restore the person to life.
mammals, in contrast, keep their muscles warm at all
times, regardless of air temperature, and therefore stay
constantly ready for vigorous activity. In other words,
water expands when it freezes, ice crystals would tear we eat a great deal to support our high metabolism so
apart blood vessels and cell membranes, killing the that even when the weather is cold, we can still run as
animal. fast and far as possible.
Ordinarily, amphibians and reptiles avoid that Why did mammals evolve a body temperature of
risk by burrowing underground or finding other shel- 37C (98F) instead of some other value? From the
tered locations. However, some frogs, fish, and insects standpoint of muscle activity, we gain an advantage
survive through winters in northern Canada where by being as warm as possible. A warmer animal has
even the underground temperature approaches 40C warmer muscles and therefore runs faster and with less
(which is also 40F). How do they do it? Some insects fatigue than a cooler animal. However, we have trade-
and fish stock their blood with glycerol and other anti- offs. To get even hotter than 37 would require still more
freeze chemicals at the start of the winter (Liou, Tocilj, energy. Furthermore, beyond about 40 or 41C, pro-
Davies, & Jia, 2000). Wood frogs actually do freeze, but teins begin to break their bonds and lose their useful
they have several mechanisms to reduce the damage. properties. Birds body temperatures are in fact about
They start by withdrawing most fluid from their organs 41C (105F).
and blood vessels and storing it in extracellular spaces. It is possible to evolve proteins that are stable at
Therefore, ice crystals have room to expand when they higher temperatures; indeed, odd microscopic animals
do form, without tearing the blood vessels and cells. called thermophiles survive in water close to its boiling
Also, they have chemicals that cause ice crystals to point (Hoffman, 2001). However, to do so, they need
form gradually, not in chunks. Finally, they have such many extra bonds to stabilize their proteins. The en-
extraordinary blood-clotting capacity that they can zymatic properties of proteins depend on the proteins
quickly repair any blood vessels that do rupture (Storey flexible structure, so making them rigid enough to with-
& Storey, 1999). stand high temperatures decreases their versatility and
As you may have heard, some people have had usefulness (Somero, 1996).
their bodies frozen after death, in hopes that scientists Reproductive cells require a somewhat cooler en-
will discover a cure for their disease and a way to bring vironment than the rest of the body (Rommel, Pabst, &
a frozen body back to life. What do you think? If you McLellan, 1998). Birds lay eggs and sit on them, instead
had enough money, would you choose this route to of developing them internally, because the birds in-
possible life after death? ternal temperature is too hot for an embryo. Similarly,
My advice is dont bother. The wood frogs that sur- in most male mammals, the scrotum hangs outside the
vive after freezing begin by dehydrating their organs body because sperm production requires a cooler tem-
and blood vessels. Unless a person underwent similar perature than the rest of the body. (A man who wears
dehydrationbefore dying!ice crystals are sure to his undershorts too tight keeps his testes too close to
tear up blood vessels and cell membranes throughout the body, overheats them, and therefore produces fewer
the body. Repairing all those membranes sounds pretty healthy sperm cells.) Pregnant women are advised to
close to impossible. avoid hot baths and anything else that might overheat
a developing fetus.
Anterior commissure
Dorsomedial
Lateral hypothalamus
hypothalamus
(behind plane of view)
Posterior hypothalamus
Anterior hypothalamus
Preoptic area
Ventromedial
Arcuate nucleus
Optic chiasm hypothalamus
Answers to
STOP & CHECK
Module 10.1 Questions
In Closing: Combining Physiological
1. Homeostasis is a set of processes that keep certain
and Behavioral Mechanisms body variables within a fixed range. Allostasis is an
One of the key themes of this module has been the re- adjustment of that range, increasing it or decreas-
dundancy of mechanisms. Your body has various phys- ing it as circumstances change. (p. 300)
pressure has not changed anywhere in your body, you An animal with osmotic thirst has an increased
need fluid. Your heart has trouble pumping blood up preference for pure water, but one with hypovolemic
to the head, and nutrients do not flow as easily as usual thirst cant drink much pure water without diluting its
into the cells. Your body will react with hormones that body fluids and changing their osmotic pressure. The
constrict blood vessels. Vasopressin is one such hor- animal therefore increases its preference for slightly
mone; another is angiotensin II. When blood volume salty water (Stricker, 1969). If the animal is offered both
drops, the kidneys release the enzyme renin, which pure water and salt, it alternates between them to yield
splits a portion off angiotensinogen, a large protein in an appropriate mixture. If sufficient salt is not readily
the blood, to form angiotensin I, which other enzymes available, it shows a strong craving for salty tastes. This
convert to angiotensin II. Like vasopressin, angioten- preference, known as sodium-specific hunger, devel-
sin II constricts the blood vessels, compensating for ops automatically as soon as the need exists, even in in-
the drop in blood pressure (Figure 10.8). fant animals (Leshem, 1999; Richter, 1936). In contrast,
Angiotensin II also helps trigger thirst in conjunc- specific hungers for other vitamins and minerals have
tion with receptors that detect blood pressure in the to be learned by trial and error (Rozin & Kalat, 1971).
large veins. However, this thirst is different from os- You may have noticed this phenomenon yourself. A
motic thirst because you need to restore your body flu- woman around the time of menstruation, or anyone
ids, including the salts, and not just water. This kind who has sweated heavily, finds that salty snacks taste
of thirst is known as hypovolemic (HI-po-vo-LEE-mik) especially good.
thirst, meaning thirst based on low volume. When an- Sodium-specific hunger depends partly on hor-
giotensin II reaches the brain, it stimulates neurons in mones (Schulkin, 1991). When the bodys sodium re-
areas adjoining the third ventricle (Fitts, Starbuck, & serves are low, the adrenal glands produce the hormone
Ruhf, 2000; Mangiapane & Simpson, 1980; Tanaka et al., aldosterone (al-DOSS-ter-one), which causes the kid-
2001). Those neurons send axons to the hypothalamus, neys, salivary glands, and sweat glands to retain salt
where they release angiotensin II as their neurotrans- (Verrey & Beron, 1996). Aldosterone and angioten-
mitter (Tanaka, Hori, & Nomura, 2001). That is, the sin II together change the properties of the neurons in
neurons surrounding the third ventricle both respond the nucleus of the tractus solitarius, part of the taste
to angiotensin II and release it. This example suggests system (see Figure 7.19), such that they begin reacting
that the connection between a neurotransmitter and to salt in nearly the same way they would to sugar
its function is not at all arbitrary; the brain uses a chem- (McCaughey & Scott, 2000).
ical that was already performing a related function else- Table 10.1 summarizes the differences between
where in the body. osmotic thirst and hypovolemic thirst.
? 316%
Battersby, 1997
Short- and Long-Term
Regulation of Feeding Figure 10.13 Chewing gum from about 4500 B.C.
The gum, made from birch-bark tar, has small tooth marks
Eating is far too important to be entrusted to just one indicating that it was chewed by a child or adolescent.
mechanism. Your brain gets messages from your mouth,
stomach, intestines, fat cells, and elsewhere to regu-
late your eating. If necessary, could you become satiated without
tasting your food? In one experiment, college students
consumed lunch five days a week by swallowing one
Oral Factors end of a rubber tube and then pushing a button to pump
Youre a busy person, right? If you could get all the nu- a liquid diet into the stomach (Jordan, 1969; Spiegel,
trition you needed by swallowing a pill, would you do 1973). (They were paid for participating.) After a few
it? Most of us would not. Never mind how much time days of practice, each person established a consistent
we would save; we like to eat. In fact, many people pattern of pumping in a constant volume of the liquid
like to taste and chew even when they are not hungry. each day and maintaining a constant body weight. Most
Figure 10.13 shows a piece of 6,500-year-old chewing found the untasted meals unsatisfying, however, and
gum made from birch-bark tar. The tooth marks indi- reported a desire to taste or chew something ( Jordan,
cate that a child or teenager chewed it. Anthropolo- 1969).
Insulin
(intermediate-term Melanocortin Neurons in the
satiety signal) paraventricular nucleus
Satiety-motive of the hypothalamus
CCK Output to other areas, including
(short-term cerebral cortex and brainstem.
satiety signal) Output increases feeding.
Lateral
hypothalamus Medial Areas of the Hypothalamus
Neuroscientists have known since the 1940s that a large
Ventromedial hypothalamus
lesion centered on the ventromedial hypothalamus
Third ventricle
(VMH) leads to overeating and weight gain (see Fig-
Figure 10.20 The lateral hypothalamus, ure 10.20). Some people with a tumor in that area have
ventromedial hypothalamus, and paraventricular gained more than 10 kg (22 lb) per month (Al-Rashid,
hypothalamus 1971; Killeffer & Stern, 1970; Reeves & Plum, 1969).
The side view above indicates the plane of the coronal Rats with similar damage sometimes double or triple
section of the brain below. (Source: After Hart, 1976) their weight (Figure 10.23). Eventually, body weight
Stage 1. Aphagia and Stage 2. Anorexia. Rat eats a Stage 3. Adipsia. The rat Stage 4. Near-recovery. The rat eats
adipsia. Rat refuses all food small amount of palatable eats enough to stay alive, enough to stay alive, though at a
and drink; must be force- foods and drinks sweetened though at a lower-than- lower-than-normal body weight. It
fed to keep it alive. water. It still does not eat normal body weight. It still drinks plain water, but only at meal-
enough to stay alive. refuses plain water. times to wash down its food. Under
slightly stressful conditions, such as
in a cold room, the rat will return
to an earlier stage of refusing food
and water.
Prefrontal cortex
(food-seeking behaviors)
Hypothalamus
Nucleus of the
tractus solitarius
(NTS)
Control rat
300
200
Food intake (grams)
30
Yoav Levy/Phototake
10
10 20 30 40 50 60
Operation Days
(a) (b)
Chapter Ending
Key Terms and Activities
Terms
agouti-related peptide (AgRP) homeostasis (p. 297) poikilothermic (p. 297)
(p. 314) homeothermic (p. 297) preoptic area/anterior
aldosterone (p. 305) hypovolemic thirst (p. 305) hypothalamus (POA/AH)
allostasis (p. 297) (p. 300)
insulin (p. 310)
angiotensin II (p. 305) set point (p. 297)
lactase (p. 308)
anorexia nervosa (p. 319) sham-feeding (p. 310)
lactose (p. 308)
arcuate nucleus (p. 313) sodium-specific hunger (p. 305)
lateral hypothalamus (p. 314)
basal metabolism (p. 297) splanchnic nerve (p. 310)
lateral preoptic area (p. 304)
bulimia nervosa (p. 320) subfornical organ (SFO) (p. 304)
leptin (p. 312)
carnivore (p. 308) supraoptic nucleus (p. 304)
melanocortin (p. 314)
cholecystokinin (CCK) (p. 310) vagus nerve (p. 310)
negative feedback (p. 297)
conditioned taste aversion vasopressin (also known as
neuropeptide Y (NPY) (p. 314)
(p. 308) antidiuretic hormone, ADH)
omnivore (p. 308) (p. 303)
cytokines (p. 301)
osmotic pressure (p. 303) ventromedial hypothalamus
duodenum (p. 310)
osmotic thirst (p. 304) (VMH) (p. 315)
ghrelin (p. 313)
OVLT (p. 304)
glucagon (p. 310)
paraventricular nucleus (PVN)
herbivore (p. 308) (pp. 304, 313)
Gisolfi, C. V., & Mora, F. (2000). The hot brain: Survival, Go to this site for the link to ThomsonNOW, your one-stop study
temperature, and the human body. Cambridge, shop, Take a Pre-Test for this chapter, and ThomsonNOW will
MA: MIT Press. Discusses research on tempera- generate a Personalized Study Plan based on your test results.
ture regulation. The Study Plan will identify the topics you need to review and
direct you to online resources to help you master these topics.
Stricker, E., & Wooes, S. (Eds.). (2004). Neurobiology of
You can then take a Post-Test to help you determine the con-
food and fluid intake (2nd ed.). New York: Kluwer.
cepts you have mastered and what you still need work on.
Collection of scholarly reviews of the literature.
Widmaier, E. P. (1998). Why geese dont get obese (and
we do). New York: W. H. Freeman. Lighthearted
and often entertaining discussion of the physiol-
ogy of eating, thirst, and temperature regulation.
Websites
to Explore
You can go to the Biological Psychology Study
Center and click these links. While there, you
can also check for suggested articles available
on InfoTrac College Edition. The Biological Psychol-
ogy Internet address is:
http://psychology.wadsworth.com/book/kalatbiopsych9e/
A video interview presents a patient with anorexia nervosa.
Internet Mental Health: Anorexia Nervosa
http://www.mentalhealth.com/dis/p20-et01.html
Exploring Biological
Psychology CD
Pathways from the Lateral Hypothalamus (animation)
Hypothalamic Control of Feeding (animation)
Anorexia Patient: Susan (video)
Chapter Quiz (multiple-choice questions)
Critical Thinking (essay questions)
Terms
Suggestions for Further Reading
Websites to Explore
Exploring Biological Psychology CD
W hat good is sex? Well, yes, I know: We enjoy it.
But why did we evolve to reproduce sexually
instead of individually? In some species, including
ThomsonNOW one kind of lizard, the female makes an egg with two
copies of each chromosome instead of one. She then
doesnt have to wait for a male; her egg simply starts
dividing and differentiating. In many ways, it would
be more efficient if you could reproduce without sex.
You wouldnt have to find a mate, and you could make
babies that are entirely like yourself, instead of just
50%. We wouldnt go to all the bother of courtship and
mating if it didnt provide an advantage.
Biologists are not entirely settled on the advantages
of sexual reproduction, and you are welcome to gener-
ate your own ideas. Currently, the dominant hypothe-
sis is that sexual reproduction increases variation and
thereby enables quick evolutionary adaptations to any
change in the environment (Colgreave, 2002; Goddard,
Opposite: Humans may be the only species that plans Godfray, & Burt, 2005).
parenthood, but all species have a strong biological drive In this chapter, we shall consider questions about
that can lead to parenthood. sexual reproduction that we often ignore or take for
Source: Art Wolfe. granted. We also consider some of the ways in which
being male or female influences our behavior.
325
Module 11.1
Sex and Hormones
Female Male
Fallopian Ejaculatory Seminal
tube duct vesicle
Gonad
Wolffian Prostate
duct
Uterus Vas
Mllerian deferens
duct
Vagina Urethra (testis) Gonad (ovary)
Epididymis
Urethra
a
b
c
Which of the lines at the d Men
left has the same angle as e
f
the one at the right? g
Ovum
becomes
FSH
fertilized
Progesterone
Estradiol
or is
FSH
Estradiol
discarded
iol
Estrad
Ovum
Follicle Follicle
Corpus
luteum
Ovary Ovary Ovary Ovary
0.3
fertility move womens mate preferences toward men
who look and act more masculine.
0.2
E X T E N S I O N S A N D A P P L I C AT I O N S
Premenstrual Syndrome
0.1
Some women experience anxiety, irritability, and de-
pression during the days just before menstruation, an
experience known as premenstrual syndrome (PMS)
15 10 5 0
Days before menstruation or premenstrual dysphoric disorder. The terms syn-
drome and disorder imply a medical problem requir-
Figure 11.8 Female-initiated sexual activities ing medical treatment and are therefore inappropriate
during the monthly cycle in most cases. Nevertheless, the terms are widely used.
The top graph shows autosexual activities (masturbation Because PMS occurs at a time of major hormonal
and sexual fantasies); the bottom graph shows female- changes, it seems reasonable to explore the possible
initiated activities with a male partner. Intrusive birth- relationship between hormones and PMS. Just before
control methods are diaphragm, foam, and condom; menstruation, estradiol and progesterone levels de-
nonintrusive methods are IUD and vasectomy. Note crease, while levels of cortisol (an adrenal hormone)
that women other than pill users increase self-initiated sex increase. However, women with PMS have about the
activities when their estrogen levels peak. (Source: From same fluctuations in these hormones as women with-
Rise in female-initiated sexual activity at ovulation and its out PMS (Schmidt, Nieman, Danaceau, Adams, & Ru-
suppression by oral contraceptives, by D. B. Adams, A. R. Gold, binow, 1998). If anything, women with PMS may have
and A. D. Burt, 1978, New England Journal of Medicine, 299, weaker fluctuations: Throughout the menstrual cycle,
pp. 11451150. Reprinted by permission of The New England they have steadier levels of estradiol, progesterone, and
Journal of Medicine.) norepinephrine than do other women (I. Blum et al.,
2004).
Much research interest focuses on the metabo-
Gold, & Burt, 1978; Udry & Morris, 1968) (Figure 11.8). lism of progesterone. Progesterone is metabolized into
According to another study, women rate an erotic video several other chemicals, including allopregnanolone,
as more pleasant and arousing if they watch it during which modifies GABA synapses, which control anxi-
the periovulatory period than if they watch it at other ety and stress responses. Several studies have found
times (Slob, Bax, Hop, Rowland, & van der Werff ten that women with PMS have normal levels of proges-
Bosch, 1996). These effects are small, though. terone but lower than normal levels of allopregnano-
Sex hormones also influence womens attention lone, particularly during the premenstrual period (Fol-
to sex-related stimuli. Women in one study were asked lesa et al., 2000; Monteleone et al., 2000; Rapkin et al.,
to look at facial photos on a screen and classify each 1997).
as male or female as quickly as possible. They made the
pendent at a later stage. If a female that has never been later phase, experience maintains the maternal behav-
pregnant is left with some baby rats, she ignores them ior even though the hormones start to decline (Rosen-
at first but gradually becomes more attentive. (Because blatt, 1970).
the babies cannot survive without parental care, the Are hormones important for human parental be-
experimenter must periodically replace them with new, havior? Hormonal changes are necessary for a woman
healthy babies.) After about 6 days, the adoptive mother to nurse a baby, but otherwise, hormonal changes are
builds a nest, assembles the babies in the nest, licks not necessary to prepare anyone for infant care. After
them, and does everything else that a normal mother all, both men and never-pregnant women can adopt
would, except nurse them. This experience-dependent children and be excellent parents. It is possible that
behavior does not require hormonal changes and oc- hormonal changes facilitate or increase some aspects
curs even in rats that have had their ovaries removed of human parental behavior, but research data are not
(Mayer & Rosenblatt, 1979; Rosenblatt, 1967). That is, available on this point.
humans are not the only species in which a mother can
adopt young without first going through pregnancy.
An important influence from being with babies is
that the mother becomes accustomed to their odors.
STOP & CHECK
Infant rats release some chemicals that stimulate the
mothers vomeronasal organ, which responds to pher- 10. What factors are responsible for maternal behavior
omones (see Chapter 7). We might imagine that evolu- shortly after rats give birth? What factors become
tion would have equipped infants with pheromones more important in later days?
that elicit maternal behavior, but actually, their phero-
Check your answers on page 337.
mones interfere with maternal behavior by stimulat-
ing aggressive behaviors (Sheehan, Cirrito, Numan, &
Numan, 2000). For a mother that has just gone through
pregnancy, this interference does not matter; her hor-
mones have primed her medial preoptic area so strongly Module 11.1
that it overrides competing impulses. A female with- In Closing: Reproductive Behaviors
out hormonal priming, however, rejects the young until
she has become familiar with their smell (Del Cerro
and Motivations
et al., 1995). A mother rat licks her babies all over shortly after their
Why do mammals need two mechanisms for ma- birth, and that stimulation is essential for their sur-
ternal behaviorone hormone-dependent and one vival. Why does she do it? Presumably, she does not
not? In the early phase, hormones compensate for the understand that licking will help them. She licks be-
mothers lack of familiarity with the young. In the cause they are covered with a salty fluid that tastes good
Girls' toys
Boys' toys
Neutral toys
Testicular Feminization
Certain individuals with an XY chromosome pattern
have the genital appearance of a female. This condition
is known as androgen insensitivity, or testicular femi-
nization. Although such individuals produce normal
amounts of androgens (including testosterone), they with a very small penis for various reasons not limited
lack the androgen receptor that enables it to activate to androgen insensitivity. How should they be reared?
genes in a cells nucleus. Consequently, the cells are Beginning in the 1950s, medical doctors began recom-
insensitive to androgens, and development proceeds mending that all intersex people be reared as girls,
as if the level of testosterone and related hormones was using surgery if necessary to make their genitals look
very low. This condition occurs in various degrees, re- more feminine (Dreger, 1998). The reason was that it
sulting in anatomy that ranges from a smaller than av- is easier to reduce an enlarged clitoris to normal size
erage penis to genitals like those of a normal female. than expand it to penis size. If necessary, surgeons can
In some cases, no one has any reason to suspect the per- build an artificial vagina or lengthen a short one. After
son is anything other than a normal female, until pu- the surgery, the child looks female. Physicians and psy-
berty. Then, in spite of breast development and broad- chologists assumed that any child who was consis-
ening of the hips, menstruation does not begin because tently reared as a girl would fully accept that identity.
the body has internal testes instead of ovaries and a And she lives happily ever after, right? Not neces-
uterus. (The vagina is short and leads to nothing.) Also, sarily. Many intersexes have complained about their
pubic hair is sparse or absent because it depends on treatment. A surgically created or lengthened vagina
androgens in females as well as males (Figure 11.13). may be satisfactory to a male partner, but it provides
no sensation to the woman and requires almost daily
Issues of Gender Assignment attention to prevent it from scarring over. Many inter-
sexes wish they had their original abnormal penis/
and Rearing clitoris instead of the mutilated, insensitive structure
Many girls with CAH and related conditions are born left to them by a surgeon. A few3 of 53 people in one
looking only slightly masculinized, but some look as studyask to be reassigned from female to male
much male as female. Some genetic males are born (Zucker et al., 1996). Moreover, intersexes resent being
Capiluppi, 2004; Hamer, Hu, Magnuson, Hu, & Patta- and other hypotheses, if indeed genes have much to
tucci, 1993). For example, uncles and cousins on the do with sexual orientation.
mothers side were more likely to be homosexual than
uncles and cousins on the fathers side. These results
suggest a gene on the X chromosome, which a man nec-
Hormones
essarily receives from his mother. However, other stud- Sexual orientation is clearly not related to adult hor-
ies have not replicated these results, and the current mone levels. Most homosexual men have testosterone
status is inconclusive (Bailey et al., 1999; Rice, Ander- and estrogen levels similar to those for heterosexual
son, Risch, & Ebers, 1999). men; most lesbian women have hormone levels simi-
If certain genes promote a homosexual orientation, lar to those for heterosexual women. A more plausi-
why hasnt evolution selected strongly against those ble hypothesis is that sexual orientation depends on
genes, which decrease the probability of reproduc- testosterone levels during a sensitive period of brain
tion? One hypothesis is that homosexual people help development (Ellis & Ames, 1987). In studies of ani-
their brothers or sisters rear children and thereby per- mals ranging from rats to pigs to zebra finches, males
petuate genes that the whole family shares (LeVay, that were exposed to much-decreased levels of testos-
1993). However, survey data indicate that homosex- terone early in life have as adults shown sexual inter-
ual men are no more likely than heterosexuals to help est in other males (Adkins-Regan, 1988). Females ex-
their nephews or nieces (Bobrow & Bailey, 2001). In- posed to extra testosterone during that period show
deed, many are quite estranged from their relatives. A an increased probability of attempting to mount sex-
second hypothesis is that certain genes sometimes lead ual partners in the way that males typically do (Fig-
to homosexuality (perhaps in men homozygous for the ure 11.15). However, in many of these studies, the an-
gene) but in other men produce advantages, such as imals not only acted somewhat like the opposite sex
increased tendency to form friendships and alliances but also looked like them.
(Rahman & Wilson, 2003). So far, this hypothesis has Homosexual and heterosexual humans look like
not been tested. each other in most ways but differ in subtle regards.
According to another hypothesis, genes that pro- On the average, the bones of the arms, legs, and hands
duce homosexuality in males are in some way advan- are longer in heterosexual men than in homosexual
tageous to their female relatives, increasing their prob- men and longer in homosexual women than in hetero-
ability of reproducing and therefore spreading the sexual women. That is, in this regard, homosexual men
genes. The results of one study support this hypoth- are partly feminized and homosexual women are
esis. Homosexual mens mothers, and aunts on the partly masculinized (Martin & Nguyen, 2004). The
mothers side, had more children on the average than length of those bones begins to differ between boys and
other women did (Camperio-Ciani et al., 2004). Obvi- girls early in lifebefore pubertyso the differences
ously, we need more research to decide among these probably relate to prenatal hormones.
Drner, 1974
later-born homosexual men tend to be shorter than av-
erage (Bogaert, 2003a). However, this hypothesis seems
to predict that if one son is homosexual, all later sons
Figure 11.15 A female rat mounting a male will be also, and that prediction is incorrect. If this hy-
The female was injected with androgens during an early pothesis is on the right track, it will need to be refined.
sensitive period; the male was castrated at birth and injected Laboratory research has shown that prenatal stress
with androgens at adulthood. can alter sexual development. In several experiments,
rats in the final week of pregnancy had the stressful
experience of confinement in tight Plexiglas tubes for
Researchers have examined brain structures that more than 2 hours each day under bright lights. In some
tend to be larger in heterosexual men than heterosex- cases, they were given alcohol as well. These rats
ual women. The results are complex. In some ways, ho- daughters looked and acted approximately normal.
mosexual men are shifted partly in the female-typical The sons, however, had normal male anatomy but, at
direction but not in other ways. Similarly, on the aver- adulthood, often responded to the presence of another
age, homosexual womens brains are slightly shifted in male by arching their backs in the typical rat female
the male direction in some ways but not others (Rah- posture for sex (I. L. Ward, Ward, Winn, & Bielawski,
man & Wilson, 2003). 1994). Most males that were subjected to either pre-
Consider another task: Experimenters repeatedly natal stress or alcohol developed male sexual behavior
present a loud noise and measure the startle response. in addition to these female sexual behaviors, but those
On some trials, they present a weaker noise just before who were subjected to both stress and alcohol had de-
the loud noise; the first noise decreases the startle re- creased male sexual behaviors (I. L. Ward, Bennett,
sponse to the louder one. The decrease is called pre- Ward, Hendricks, & French, 1999).
pulse inhibition. Prepulse inhibition is ordinarily Prenatal stress and alcohol may alter brain devel-
stronger in men than in women. In this regard, homo- opment through several routes. Stress releases endor-
sexual men do not differ significantly from heterosex- phins, which can antagonize the effects of testosterone
ual men, but homosexual women are slightly shifted in on the hypothalamus (O. B. Ward, Monaghan, & Ward,
the male direction compared to heterosexual women 1986). Stress also elevates levels of the adrenal hor-
(Rahman, Kumari, & Wilson, 2003). mone corticosterone, which decreases testosterone re-
Overall, what do these results indicate? Homosex- lease (O. Ward, Ward, Denning, French, & Hendricks,
ual women appear to have been masculinized, on the 2002; M. Williams, Davis, McCrea, Long, & Hennessy,
average, in some ways and not others. Homosexual 1999). The long-term effects of either prenatal stress
men appear to be shifted in the female direction in or alcohol include several changes in the structure of
some ways and not others. These results cannot make the nervous system, making the affected males anat-
sense in terms of an overall increase or decrease in omy closer to that of females (Nosenko & Reznikov,
prenatal exposure to testosterone. They might, how- 2001; I. Ward, Romeo, Denning, & Ward, 1999).
ever, indicate altered amounts of testosterone at par- Although the relevance of these results to humans
ticular stages of development. They might also indi- is debatable, they prompted investigators to exam-
cate that certain brain areas have altered sensitivities ine possible effects of prenatal stress on humans. One
to testosterone (Rahman & Wilson, 2003). Bear in mind approach is to ask the mothers of homosexual men
that each of the results varies from one individual to whether they experienced any unusual stress during
another, and there are no doubt many factors influenc- pregnancy. Three surveys compared mothers of homo-
ing sexual orientation. sexual sons to mothers of heterosexual sons. In two of
the three, the mothers of homosexual sons recalled
more than average stressful experiences during their
Prenatal Events pregnancies (Bailey, Willerman, & Parks, 1991; Ellis,
The probability of a homosexual orientation is higher Ames, Peckham, & Burke, 1988; Ellis & Cole-Harding,
among men with older brothers than among oldest sons. 2001). However, these studies relied on womens mem-
Brain Anatomy
On the average, mens brains differ from
womens in several ways, including the Image not available due to copyright restrictions
sizes of several parts of the hypothala-
mus (Swaab, Chung, Kruijver, Hofman,
& Ishunina, 2001). Do the brains of ho-
mosexual men differ from those of het-
erosexual men?
Results vary from one brain area to
another. The anterior commissure (see
p. 91) is, on the average, larger in hetero-
sexual women than in heterosexual men;
in homosexual men, it is at least as large
as in women, perhaps even slightly larger
(Gorski & Allen, 1992). The implications
of this difference are unclear, as the an-
terior commissure has no known relationship to sex- that the difference between heterosexual men and the
ual behavior. The suprachiasmatic nucleus (SCN) is other two groups is fairly large and that the cause of
also larger in homosexual men than in heterosexual death (AIDS versus other) has no clear relationship to
men (Swaab & Hofman, 1990). Recall from Chapter 9 the results. LeVay (1993) later examined the hypothal-
that the SCN controls circadian rhythms. How might amus of a homosexual man who died of lung cancer;
a difference in the SCN relate to sexual orientation? The he had a small INAH-3, like the homosexual men who
answer is not clear, but male rats that are deprived of died of AIDS.
testosterone during early development also show ab- A later study partly replicated these general trends.
normalities in the SCN, and their preference for male Researchers found that the INAH-3 nucleus was slightly
or female sexual partners varies with time of day. They larger in heterosexual than homosexual men, although
make sexual advances toward both male and female in this study the homosexual mens INAH-3 nucleus
partners early in their active period of the day but was larger than that of heterosexual women (Byne
mostly toward females as the day goes on (Swaab, Slob, et al., 2001). Among heterosexual men or women, the
Houtsmuller, Brand, & Zhou, 1995). Does human sex- INAH-3 nucleus was larger in those who were HIV neg-
ual orientation fluctuate depending on time of day? ative than those who were HIV positive, but even if we
No research has been reported. look only at HIV+ men, we still find a difference in the
The most suggestive studies concern the third inter- hypothalamus between heterosexual and homosexual
stitial nucleus of the anterior hypothalamus (INAH-3), men. Figure 11.18 displays the means for the five
which is generally more than twice as large in hetero- groups. On microscopic examination of the INAH-3,
sexual men as in women. This area corresponds to researchers found that heterosexual men had larger
part of the sexually dimorphic nucleus, which is larger neurons than homosexual men but about the same
in male than female rats. Simon LeVay (1991) exam- number. (Neither this study nor LeVays earlier study
ined INAH-3 in 41 people who had died between the included homosexual females.)
ages of 26 and 59. Of these, 16 were heterosexual men, Interpreting these studies is another matter, how-
6 were heterosexual women, and 19 were homosexual ever. One possibility is that brain differences predis-
men. All of the homosexual men, 6 of the 16 hetero- posed some men to become homosexual and others to
sexual men, and 1 of the 6 women had died of AIDS. become heterosexual. Another possibility is that dif-
LeVay found that the mean volume of INAH-3 was ferent kinds of sexual activity produce changes in the
0.12 mm3 in heterosexual men, 0.056 mm3 in hetero- size of the adult hypothalamic neurons. Some brain
sexual women, and 0.051 mm3 in homosexual men. areas do grow or shrink in adults as a result of hormonal
Figure 11.16 shows typical cross-sections for a hetero- or behavioral influences (Cooke, Tabibnia, & Breed-
sexual man and a homosexual man. Figure 11.17 shows love, 1999). A further limitation is that we do not know
the distribution of volumes for the three groups. Note the role of INAH-3 in human sexual behavior.
0.15 0.08
0.06
0.04
0.10
0.02
0
Hetero Hetero Homo Hetero Hetero
0.05 male male male female female
HIV HIV+ HIV+ HIV HIV+
Summary
STOP & CHECK
1. Many of the mating habits of people make sense in
terms of increasing the probability of passing on our
6. What kind of experience in early development can genes. If we saw the same behaviors in nonhumans,
cause a male rat to develop sexual responsiveness we probably assume a genetic, evolved basis. How-
to other males and not to females? How does that ever, in humans, we cannot assume a genetic basis
experience probably produce its effects? because people may have learned these behaviors
7. What is the relationship between birth order and and preferences. (p. 339)
homosexuality? 2. People can develop ambiguous genitals or genitals
8. What evidence argues against the hypothesis that that dont match their chromosomal sex for several
homosexual men have been feminized in general reasons. One is congenital adrenal hyperplasia, in
or that homosexual women have been masculinized which a genetic defect in cortisol production leads
in general? to overstimulation of the adrenal gland and there-
Check your answers on page 350. fore extra testosterone production. When that con-
dition occurs in a female fetus, she becomes partly
masculinized. (p. 341)
3. On the average, girls with a history of congenital
Module 11.2 adrenal hyperplasia show more interest in boy-
typical toys than other girls do, and during adoles-
In Closing: We Are Not All the Same
cence and young adulthood, they continue to show
When Alfred Kinsey conducted the first massive sur- partly masculinized interests. These trends appar-
veys of human sexual behavior, he found that people ently relate to the influence of prenatal hormones.
varied enormously in their frequency of sexual acts, but (p. 342)
Terms
activating effect (p. 327) hermaphrodite (p. 341) progesterone (p. 326)
alpha-fetoprotein (p. 329) impotence (p. 332) sensitive period (p. 328)
androgen (p. 326) intersex (or pseudohermaphrodite) sex-limited genes (p. 326)
androgen insensitivity (or (p. 341) sexually dimorphic nucleus
testicular feminization) (p. 343) luteinizing hormone (LH) (p. 333) (p. 329)
congenital adrenal hyperplasia menstrual cycle (p. 332) SRY gene (p. 327)
(CAH) (p. 341) Mllerian duct (p. 327) steroid hormones (p. 326)
estradiol (p. 326) organizing effect (p. 327) testis (p. 327)
estrogen (p. 326) ovary (p. 328) testosterone (p. 327)
follicle-stimulating hormone (FSH) periovulatory period (p. 333) Wolffian duct (p. 327)
(p. 332)
premenstrual syndrome (PMS)
gender identity (p. 341) (p. 334)
Terms
Suggestions for Further Reading
Websites to Explore
S uppose researchers have discovered a new spe-
cieslets call it species Xand psychologists
begin testing its abilities. They let X touch a blue tri-
angle that is extremely hot. X makes a loud sound and
Exploring Biological Psychology CD
backs away. Then someone picks up the blue triangle
ThomsonNOW
(with padded gloves) and starts moving with it rapidly
toward X. As soon as X sees this happening, it makes
the same sound, turns, and starts moving rapidly away.
Shall we conclude that it feels the emotion of fear?
If you said yes, now let me add: I said this was a
new species, and so it is, but its a new species of robot,
Opposite: Even a static photo or statue can convey strong not animal. Do you still think X feels emotions?
emotion, as in this statue, The Angry Boy. With movement, If such behavior isnt adequate evidence for emo-
we can express even more. tion in a robot, is it adequate evidence for an animal?
Source: Gunnar Strom Biological researchers therefore concentrate mostly on
emotional behaviors, which are observable, even if the
emotions behind them are not.
353
Module 12.1
What Is Emotion?
Celiac
ganglion Thoracic
nerves
(12 pairs)
Lumbar
nerves
(5 pairs)
(Most ganglia
near spinal cord) Sacral
Pelvic nerves
nerve (5 pairs)
The James-Lange theory leads to two predictions: blood from your head toward the ground, except that
People with weak autonomic or skeletal responses your autonomic nervous system increases your heart
should feel less emotion, and causing or increasing rate and constricts the veins in your head. In someone
someones responses should enhance an emotion. Lets with pure autonomic failure, those reflexes do not
consider the evidence. occur. To avoid fainting, people have to learn to stand
up slowly. Also, people with this condition have no
Is Physiological Arousal changes in heart rate, blood pressure, or sweating dur-
ing any kind of psychological stress or physical chal-
Necessary for Emotions? lenge. According to the James-Lange theory, we would
People with damage to the spinal cord are paralyzed expect them to report no emotions. In fact, they report
from the level of the damage downward. People who the same emotions as anyone else and have little diffi-
are unable to move their arms and legs certainly can- culty identifying what emotion a character in a story
not attack or run away. Most such people report that would probably experience (Heims, Critchley, Dolan,
they feel emotions about the same as before their in- Mathias, & Cipolotti, 2004). However, they report that
jury (Cobos, Snchez, Prez, & Vila, 2004). This find- they feel their emotions much less intensely than before
ing indicates that emotions do not depend on feedback the onset of the disease (Critchley, Mathias, & Dolan,
from muscle movements. However, paralysis does not 2001). Presumably, when they report emotions, they
affect the autonomic nervous system, so it remains are reporting the cognitive aspect: Yes, Im angry; this
possible that emotional feelings depend on feedback is a situation that calls for anger. But they do not feel
from autonomic responses. much anger. Their decreased emotional feeling is con-
In people with an uncommon condition called pure sistent with predictions from the James-Lange theory.
autonomic failure, output from the autonomic nervous
system to the body fails, either completely or almost Is Physiological Arousal
completely. Heart beat and other activities continue to
occur, but the nervous system no longer regulates them.
Sufficient for Emotions?
One effect occurs when people stand up. When you According to the James-Lange theory, emotional feel-
suddenly stand up, gravity and inertia would pull the ings result from our bodys actions. If your heart started
(a)
Figure 12.7
Testosterone levels
for men convicted
Armed
Rape Burglary of various crimes
robbery
Men convicted of
rape and murder have
higher testosterone
levels, on the average,
Higher testosterone than men convicted
Murder Drug offenses Intermediate testosterone of burglary or drug
offenses. (Source: Based
Lower testosterone
on Dabbs, Carr, Frady, &
Riad, 1995)
Nonhuman Animals
Much of the earliest evidence came from studies on
mice. Luigi Valzelli (1973) found that isolating male
mice for 4 weeks increased their aggressive behav-
ior and decreased their serotonin turnover. Turnover
is the amount of release and resynthesis of a neuro-
transmitter by presynaptic neurons. That is, a brain
with low serotonin turnover releases relatively small
amounts of serotonin and therefore does not need to
resynthesize much of it. Serotonin turnover can be
low even if the brains total serotonin content is nor-
mal. Researchers infer turnover from the concentration
of 5-hydroxyindoleacetic acid (5-HIAA), a serotonin
metabolite, in the blood, cerebrospinal fluid (CSF), or
Delgado, 1981
urine. High levels of 5-HIAA imply much serotonin
release and turnover.
Comparing different genetic strains of mice, Val-
Figure 12.8 Effects of stimulation in the zelli and his colleagues found the lowest serotonin
medial hypothalamus turnover in strains that fought the most (Valzelli &
Stimulation in some brain areas can lead to a full attack; Bernasconi, 1979). Social isolation does not decrease
in this case the result was undirected growling and facial serotonin turnover in female mice in any genetic strain,
expressions, a mere fragment of a normal attack. and it does not make the females aggressive.
In a fascinating study, investigators measured
5-HIAA levels in 2-year-old male monkeys living in a
natural environment and then observed their behavior
was talking to his wife. He would then interpret
closely. The monkeys in the lowest quartile for 5-HIAA,
something she said as an insult, throw her against
and therefore the lowest serotonin turnover, were the
the wall and attack her brutally for 5 to 6 minutes.
most aggressive, had the greatest probability of attack-
After one of these attacks, he would go to sleep
ing larger monkeys, and incurred the most injuries.
for a half hour and wake up feeling refreshed.
Most of them died by age 6, whereas all monkeys with
Eventually, he was taken to a hospital, where
high serotonin turnover survived (Higley et al., 1996).
epileptic activity was found in the temporal lobes
Female monkeys with low 5-HIAA levels are also likely
of his cerebral cortex. For the next 7 months, he
to get injured and die young (Westergaaard, Cleveland,
was given a combination of anti-anxiety drugs,
Trenkle, Lussier, & Higley, 2003).
antiepileptic drugs, and other medications. None
If most monkeys with low turnover die young, why
of these treatments reduced his violent behavior.
hasnt natural selection eliminated the genes for low
Psychiatrists had previously treated him for 7 years
serotonin turnover? One possibility is that evolution se-
without apparent effect. Eventually, he agreed to
lects for an intermediate amount of aggression and anx-
a surgical operation to destroy a small part of the
iety (Trefilov, Berard, Krawczak, & Schmidtke, 2000).
amygdala on both sides of the brain. Afterwards,
The most fearless animals get into fights and die young,
he had no more episodes of rage, although he
but those with excessively high fears have other prob-
continued to have periods of confusion and dis-
lems. A study of laboratory rats found that those show-
ordered thinking.
ing the greatest fearmeasured by hesitance to explore
Temporal lobe epilepsy is uncommon, and most a new environmentdied of natural causes earlier than
people with this kind of epilepsy do not show aggres- other rats (Cavigelli & McClintock, 2003). A likely hy-
sive behaviors. Consequently, this brain abnormality pothesis is that they overreact to stressful events and
accounts for only a small percentage of human aggres- thereby impair their health.
sive acts. Another explanation sees aggressiveness as a high-
risk, high-payoff strategy: Most monkeys with 5-HIAA
Serotonin Synapses die young, but the survivors often achieve a dominant
status within their troop. In contrast, those with high
and Aggressive Behavior 5-HIAA survive but remain submissive (Fairbanks et al.,
Several lines of evidence link aggressive behavior to 2004). A dominant male has the opportunity to im-
low serotonin release. Nearly all the evidence relies on pregnate females beginning at an earlier age than other
Humans
Many studies have found low serotonin turnover in
people with a history of violent behavior, including
people convicted of arson and other violent crimes
(Virkkunen, Nuutila, Goodwin, & Linnoila, 1987) and Never Attempted Attempted
people who attempt suicide by violent means (G. L. attempted once again
Brown et al., 1982; Edman, sberg, Levander, & Schal- suicide
ling, 1986; Mann, Arango, & Underwood, 1990; Pandey
Figure 12.9 Levels of 5-HIAA in the CSF of
et al., 1995; Spreux-Varoquaux et al., 2001). Serotonin
depressed people
turnover varies by 5% to 10% from one time of year
Measurements for the two suicide-attempting groups were
to another; a study in Belgium found higher suicide
taken after the first attempt. Low levels of 5-HIAA indicate
rates in spring, when serotonin turnover was lowest,
low serotonin turnover. (Source: Based on results of Roy, DeJong,
and lower rates in fall and winter, when serotonin
& Linnoila, 1989)
turnover was highest (Maes et al., 1995).
A study of children and adolescents with a his-
tory of aggressive behavior found that those with the
lowest serotonin turnover were most likely to get into with less active forms of this enzyme are more likely
trouble for additional aggressive behavior during the than other people to report frequent anger and aggres-
following 2 years (Kruesi et al., 1992). Follow-up stud- sion (Hennig, Reuter, Netter, Burk, & Landt, 2005; Ru-
ies on people released from prison found that those jescu et al., 2002) and more likely than others to make
with lower serotonin turnover had a greater probability violent suicide attempts (Abbar et al., 2001). Another
of further convictions for violent crimes (Virkkunen, relevant gene controls the serotonin transporterthe
DeJong, Bartko, Goodwin, & Linnoila, 1989; Virkkunen, protein in the membrane of the presynaptic neuron,
Eggert, Rawlings, & Linnoila, 1996). A study of people which reabsorbs serotonin after its release and use. The
who had survived suicide attempts found that low short form of this gene, linked to decreased activity
serotonin turnover levels predicted additional suicide of the transporter, theoretically should increase sero-
attempts within the next 5 years (Roy, DeJong, & Lin- tonin transmission by prolonging the presence of sero-
noila, 1989) (Figure 12.9). However, although each of tonin in the synapse after its release. Behaviorally, this
these relationships is statistically reliable, blood tests gene is linked to increased anxiety, which usually de-
do not enable anyone to identify dangerous people with presses aggressive behavior (Gross & Hen, 2004; Hariri
confidence and then take preventive measures. et al., 2002).
It is possible to alter serotonin synthesis by changes Still another gene controls production of the en-
in diet. Neurons synthesize serotonin from tryptophan, zyme monoamine oxidase (MAO), which breaks down
an amino acid found in proteins, though seldom in large serotonin into inactive chemicals. People who produce
amounts. Tryptophan crosses the blood-brain barrier by less of this enzyme, and who were also mistreated as
an active transport channel that it shares with phenyl- children, are prone to violence and antisocial behav-
alanine and other large amino acids. Thus, a diet high iors (Caspi et al., 2002). This result is interesting in its
in other amino acids impairs the brains ability to syn- emphasis on the combination of both genetics and so-
thesize serotonin. One study found that many young cial environment. People with either the genetic or en-
men on such a diet showed an increase in aggressive vironmental influence, but not both, were only mildly
behavior a few hours after eating (Moeller et al., 1996). prone to violence. Theoretically, however, the result
Under the circumstances, it would seem prudent for is puzzling because decreased MAO should increase
anyone with aggressive or suicidal tendencies to re- serotonin levels and therefore presumably decrease
duce consumption of aspartame (NutraSweet), which aggression.
is 50% phenylalanine, and maize (American corn), Another problem: We shall get to depression in
which is high in phenylalanine and low in tryptophan Chapter 15, but you might already know from other
(Lytle, Messing, Fisher, & Phebus, 1975). readings that depression is linked to low serotonin
Might some of the variation in serotonin activity, activity. Nevertheless, most depressed people are not
and therefore violence, relate to genetics? People vary violent. If some treatment suddenly lowered your sero-
in the gene that controls tryptophan hydroxylase, the tonin level, would you at once become violent, de-
enzyme that converts tryptophan into serotonin. People pressed, or what? When researchers use drugs or diet to
STOP & CHECK Peoples choices of activities depend in part on how easily
they develop anxiety.
Happy Sad
Surprised Disgusted
Microdialysis
Silica tubing
In microdialysis, an investigator implants into the brain In Glue
a double fluid-filled tube with a thin membrane tip across
which chemicals can diffuse. The experimenter slowly Polyethylene
delivers some fluid through one tube, while an equal tubing
amount of fluid exits through the other tube and brings
with it some of the brain chemicals that have diffused
across the membrane. In this manner, researchers dis-
cover which neurotransmitters are released during par-
ticular behaviors (e.g., Stanley, Schwartz, Hernandez,
Leibowitz, & Hoebel, 1989). Stainless steel
tubing
Active
Porous
region
Source: Reprinted with permission from Dr. Juan Dominguez membrane
Anxiety-Reducing Drugs
A little anxiety can be a useful thing by alerting us to
danger. However, some people have exaggerated fears
and excessive reactions to frightening events, partly for Resident
genetic reasons (Garpenstrand, Annas, Ekblom, Ore- Protective
land, & Fredrikson, 2001; Gratacs et al., 2001; Kend- cage wire
ler, Myers, Prescott, & Neale, 2001). As you might mesh
guess, people with excessive fears have a hyperactive
amygdala. One study found that children with panic Intruder
disorder or generalized anxiety disorder had exagger-
ated amygdala responses to photos showing fearful
faces (K. M. Thomas et al., 2001). Figure 12.15 Procedure for giving male
Drugs intended to control anxiety alter activity at rats anxiety
amygdala synapses. One of the amygdalas main excita- Male rats were placed in a cage, protected from the resident
tory neuromodulators is CCK (cholecystokinin), which male. Later, the protection was removed. For some, the
increases anxiety, and the main inhibitory transmitter resident male was removed at the same time, so the intruder
is GABA, which inhibits anxiety. was safe. For others, the protection was removed but the
Here is a good experiment showing the role of CCK: resident male was left to attack the intruder. Those who
Male intruder rats were placed in a protected area were attacked (and inevitably defeated) developed great
within a resident male rats cage for 30 minutes at a anxiety when again placed in the same location.
time, at which point the protective barrier was removed
(Figure 12.15). For some of the intruders, the resident
male was removed at the same time as the barrier. For that had been defeated four times by the resident male
the others, the protection was removed but not the res- showed clear signs of anxiety when again placed into
ident. Those intruders were always attacked and de- that males cage (motionlessness, defensive postures,
feated. (The home field advantage is enormous for and squeals); his prefrontal cortex showed an enor-
rats. A resident male almost always defeats an intruder.) mous increase in release of CCK. However, if he was
After four repetitions of this experience, the research- given a drug that blocked his CCK type B receptors,
ers implanted a cannula into the intruder males brain he showed no anxiety, even when placed in the dan-
and used microdialysis (Methods 12.1) to measure gerous cage (C. Becker et al., 2001). The brain also has
brain chemistry. The result was that an intruder male CCK type A receptors, less abundant, with different
Neuron membrane
Cl Cl
E X T E N S I O N S A N D A P P L I C AT I O N S
Alcohol as an Anxiety Reducer
Ethyl alcohol, the beverage alcohol, has behavioral ef- they were sober and try to drive home when they were
fects similar to those of benzodiazepines. It decreases still somewhat impaired. Furthermore, giving such a
anxiety and behavioral inhibitions. Moreover, a com- pill to alcoholics could easily backfire. Alcoholics gen-
bination of alcohol and benzodiazepines depresses erally drink to get drunk; a pill that decreased their
body activities and brain functioning more severely feeling of intoxication would probably lead them to
than either drug alone would. (A combination of both drink even more. Ro15-4513 reverses the behavioral
can be fatal.) Furthermore, alcohol, benzodiazepines, effects of moderate alcohol doses, but a large dose can
and barbiturates all exhibit the phenomenon of cross- still be a health hazard or even fatal (Poling, Schlinger,
tolerance: An individual who has used one of the & Blakely, 1988). For these reasons, Ro15-4513 is used
drugs enough to develop a tolerance to it will show a only in experimental laboratories.
partial tolerance to other depressant drugs as well.
Alcohol promotes the flow of chloride ions through
the GABAA receptor complex, just as benzodiazepines
do (Suzdak et al., 1986), probably by facilitating the
binding of GABA to its receptors. Alcohol influences
STOP & CHECK
the brain in other ways as well, but the effects on GABA
are responsible for alcohols anti-anxiety and intoxi- 9. What would be the effect of benzodiazepines on
cating effects. Drugs that block the effects of alcohol someone who had no GABA?
on the GABAA receptor complex also block most of
Check your answer on page 375
alcohols behavioral effects. One experimental drug,
known as Ro15-4513, is particularly effective in this re-
gard (Suzdak et al., 1986). Besides affecting the GABAA
receptor complex, Ro15-4513 blocks the effects of al-
cohol on motor coordination, its depressant action on Module 12.2
the brain, and its ability to reduce anxiety (H. C. Becker, In Closing: Doing Something
1988; Hoffman, Tabakoff, Szabo, Suzdak, & Paul, 1987;
Ticku & Kulkarni, 1988) (Figure 12.17).
About Emotions
Could Ro15-4513 be useful as a sobering-up pill Today, we understand the physiology of violence and
or as a treatment to help people who want to stop drink- fearfulness far better than we used to, but clearly, we
ing alcohol? Hoffman-LaRoche, the company that dis- have a long way to go. If we do make major advances,
covered it, eventually concluded that the drug would what then? Imagine we could take a blood sample
be too risky. People who relied on the pill might think measuring 5-HIAA or whateverplus an fMRI scan
Leukocytes
The most important elements of the immune system are
ACTH the leukocytes, commonly known as white blood cells
(through blood) (Kiecolt-Glaser & Glaser, 1993; OLeary, 1990).
We distinguish several types of leukocytes, includ-
ing B cells, T cells, and natural killer cells (Figure 12.19):
B cells, which mature mostly in the bone marrow, se-
Adrenal crete antibodies, which are Y-shaped proteins that at-
cortex
tach specifically to particular kinds of antigen, just as
a key fits a lock. Every cell has surface proteins called
antigens (antibody-generator molecules), and your
own bodys antigens are as unique as your finger-
Cortisol prints. The B cells recognize the self antigens, but
when they find an unfamiliar antigen, they attack the
Figure 12.18 The hypothalamus-pituitary-adrenal cell. This kind of attack defends the body against
cortex axis viruses and bacteria; it also causes rejection of organ
Prolonged stress leads to the secretion of the adrenal transplants, unless physicians take special steps to
hormone cortisol, which elevates blood sugar and increases minimize the attack. After the body has made anti-
metabolism. These changes help the body sustain prolonged bodies against a particular intruder, it remembers
activity but at the expense of decreased immune system the intruder and quickly builds more of the same
activity. kind of antibody if it encounters that intruder again.
T cells mature in the thymus gland. Several kinds
of T cells attack intruders directly (without secreting
antibodies), and some help other T cells or B cells
Many researchers refer to cortisol as a stress hor-
to multiply.
mone and use measurements of cortisol level as an
Natural killer cells, another kind of leukocytes, at-
indication of someones recent stress level. One prob-
tack tumor cells and cells that are infected with vi-
lem in that regard is that cortisol levels rise and fall
ruses. Whereas each B or T cell attacks a particular
during the day, so measurements taken from different
kind of foreign antigen, natural killer cells non-
people at different times of day are not comparable
specifically attack all intruders.
(Bremner et al., 2003).
Cortisol helps the body mobilize its energies to In response to an infection, leukocytes and other
fight a difficult situation, but the effects depend on cells produce small proteins called cytokines (e.g., in-
amount. For example, moderate levels of cortisol im- terleukin-1, or IL-1), which combat infections and also
prove attention and memory formation (Abercrombie, communicate with the brain to elicit appropriate be-
Kalin, Thurow, Rosenkranz, & Davidson, 2003), and haviors (Maier & Watkins, 1998). Cytokines are the im-
either higher or lower levels impair them (Kuhlmann, mune systems way of telling the brain that the body
Piel, & Wolf, 2005; Mizoguchi, Ishige, Takeda, Aburada, is ill. Although cytokines do not cross the blood-brain
& Tabira, 2004). Temporary increases in cortisol en- barrier, they stimulate receptors on the vagus nerve
hance activity of the immune system, helping it fight (see Figure 4.9, p. 89), which relays a message to hypo-
illnesses (Benschop et al., 1995), but prolonged in- thalamic and hippocampal cells that release cytokines
creases impair immune activity. To see why, we start themselves. You might think of it as sending a fax,
with an overview of the immune system. where the original material doesnt travel but tells the
recipient to make a copy. The release of cytokines in
the brain produces symptoms of illness, such as fever,
The Immune System sleepiness, lack of energy, lack of appetite, and loss of
The immune system consists of cells that protect the sex drive. In other words, cytokines are responsible for
body against such intruders as viruses and bacteria. what Selye called the general adaptation syndrome.
B Some B cells
B cell attaches
memory differentiate
to a bacterium,
cells into memory cells
leaving the B cell prepared to attack
bacterium's
the same antigen
antigen
exposed
Secretions of cytokines
Helper T cell Helper
causes this T cell
B cell to divide
Plasma cell
Circulating antibodies
attach to the antigen
Some B cells and mark it for destruction
become plasma cells
that secrete antibodies
specific to this antigen
Note that these symptoms of illness are what the over the immune system. The study of this relationship,
body does, by way of cytokines, in response to an in- called psychoneuroimmunology, deals with the ways
fection. Most people think of fever and fatigue as some- in which experiences, especially stressful ones, alter the
thing the illness did to them; in fact, fever and fatigue immune system, and how the immune system in turn
are among the strategies we have evolved for fighting influences the central nervous system (Ader, 2001).
the illness. As discussed in Chapter 10, a moderate fever Stress affects the immune system in several ways.
helps fight many infections. Sleep and inactivity are In response to a stressful experience, the nervous sys-
ways of conserving energy so that the body can devote tem activates the immune system to increase its produc-
more energy to its immune attack against the intruders. tion of natural killer cells, the leukocyte scavengers, and
the secretion of cytokines (Segerstrom & Miller, 2004).
Even fear or anger enhances the immune system re-
STOP & CHECK sponse, temporarily (Mayne, 1999). During the stressful
period of final exams, many college students have in-
creased immune system activity (Liu et al., 2002). The
1. What kind of cell releases cytokines, and how do elevated cytokine levels help combat infections, but they
cytokines affect brain activity? also trigger the brain to produce the same symptoms
2. What behavioral changes do cytokines stimulate? as if one were ill. Rats subjected to inescapable shocks
show symptoms resembling illness, including fever,
Check your answers on page 380.
sleepiness, and decreased appetite. The same is true for
people who are under great stress, such as those who are
dreading some public speech they have to give (Maier &
Effects of Stress on the Immune System Watkins, 1998). In short, if you have been under a great
Contrary to assumptions biologists used to hold, we deal of stress and start to feel ill, one possibility is that
now know that the nervous system has much control your symptoms are reactions to the stress itself.
Terms
ACTH (p. 376) cytokines (p. 377) leukocyte (p. 377)
antibody (p. 377) diazepam-binding inhibitor (DBI) limbic system (p. 357)
antigen (p. 377) or endozepine (p. 372) natural killer cell (p. 377)
B cell (p. 377) GABAA receptor complex (p. 372) panic attack (p. 356)
barbiturate (p. 372) general adaptation syndrome posttraumatic stress disorder
(p. 376) (PTSD) (p. 379)
Behavioral Activation System
(BAS) (p. 358) HPA axis (p. 376) psychoneuroimmunology (p. 378)
Behavioral Inhibition System (BIS) 5-hydroxyindoleacetic acid pure autonomic failure (p. 355)
(p. 358) (5-HIAA) (p. 364)
startle reflex (p. 366)
behavioral medicine (p. 376) immune system (p. 377)
stress (p. 376)
benzodiazepine (p. 372) intermittent explosive disorder
T cell (p. 377)
(p. 363)
cortisol (p. 376) turnover (p. 364)
James-Lange theory (p. 354)
cross-tolerance (p. 373)
Websites
to Explore
You can go to the Biological Psychology Study http://www.thomsonedu.com
Center and click these links. While there, you Go to this site for the link to ThomsonNOW, your one-stop study
can also check for suggested articles available shop, Take a Pre-Test for this chapter, and ThomsonNOW will
on InfoTrac College Edition. The Biological Psychol- generate a Personalized Study Plan based on your test results.
ogy Internet address is: The Study Plan will identify the topics you need to review and
http://psychology.wadsworth.com/book/kalatbiopsych9e/ direct you to online resources to help you master these topics.
You can then take a Post-Test to help you determine the con-
Stress-Related Links cepts you have mastered and what you still need work on.
http://www.stressless.com/AboutSL/StressLinks.cfm
383
Module 13.1
Learning, Memory, Amnesia,
and Brain Functioning
Salivation
Conditioned Conditioned
stimulus (CS) response (CR)
(a) Classical conditioning
Froot
Loops
Reinforcement
Increased
probability
of same response
Shock
Response:
Rat enters one arm Punishment
of the maze
Shift to
different
(b) Operant conditioning response
and quote on pages inside the back cover.) Lashley was Lashley reasoned that if learning depends on new
searching for the engramthe physical representation or strengthened connections between two brain areas,
of what has been learned. (A connection between two a knife cut somewhere in the brain should interrupt
brain areas would be a possible example of an engram.) that connection and abolish the learned response. He
UCR CS
(a)
Figure 13.3 View of rat brain from above, showing
cuts that Lashley made in the brains of various rats
He found that no cut or combination of cuts interfered with
a rats memory of a maze. (Source: Adapted from Lashley, 1950)
UCS
trained rats on a variety of mazes and a brightness dis- The Modern Search for the Engram
crimination task and then made one or more deep cuts
Richard F. Thompson and his colleagues used a sim-
in varying locations in their cerebral cortices (Lashley,
pler task than Lashleys and sought the engram of mem-
1929, 1950) (Figure 13.3). However, no knife cut sig-
ory not in the cerebral cortex but in the cerebellum.
nificantly impaired the rats performances. Evidently,
Thompson and his colleagues studied classical condi-
the types of learning that he studied did not depend on
tioning of eyelid responses in rabbits. They presented
connections across the cortex.
first a tone (CS) and then a puff of air (UCS) to the cor-
Lashley also tested whether any portion of the cere-
nea of the rabbits eye. At first, a rabbit blinked at the air
bral cortex is more important than others for learning.
puff but not at the tone; after repeated pairings, classi-
He trained rats on mazes before or after he removed
cal conditioning occurred and the rabbit blinked at the
large portions of the cortex. The lesions impaired per-
tone also. Investigators recorded the activity in vari-
formance, but the amount of retardation depended
ous brain cells to determine which ones changed their
more on the amount of brain damage than on its loca-
responses during learning.
tion. Learning and memory apparently did not rely on
Thompson and other investigators consistently
a single cortical area. Lashley therefore proposed two
found changes in cells of one nucleus of the cerebel-
principles about the nervous system:
lum, the lateral interpositus nucleus (LIP). At the start
equipotentialityall parts of the cortex contribute of training, those cells showed little response to the
equally to complex behaviors such as learning; any tone, but as learning proceeded, their responses in-
part of the cortex can substitute for any other. creased (Thompson, 1986). However, the fact that a
A B C D E F
Corpus callosum
Cerebral cortex
Lateral
interpositus
nucleus of
cerebellum
% CRs 60 % CRs 60
40 40
20 20
0 0
Session 1 2 3 4 5 6 Session 1 2 3 4 5 6
Responses in intact animal Responses after LIP recovers activity Responses after red nucleus recovers activity
100 100 100
80 80 80
60 60 60
% CRs % CRs % CRs
40 40 40
20 20 20
0 0 0
Session 1 2 3 4 5 6 Session 1 2 3 4 Session 1 2 3 4
you dont have to forget something old to make room Probably not long. How long would you remember,
for it. The score is 64 to 57 with 2 minutes and 15 seconds
Short-term memories fade quickly unless you re- left to play? Providing a meaningful context makes
hearse them. For example, if you read the letter se- the material easier to remember, even though it is still
quence DZLAUV and then something distracts you a temporary memory.
so that you cannot rehearse those letters, your chance With short-term memory, once you have forgot-
of repeating them correctly is fairly good 5 seconds ten something, it is lost. For example, if you read
later, less after 10 seconds, and poor at 20 seconds CYXGMBF and then forget it, you probably wont
or beyond (Peterson & Peterson, 1959). In contrast, gain from the hint starts with CY. With long-term
you can recall many long-term memories that you memory, however, you might think you have forgot-
havent thought about in years. Note, however, that ten something and yet find that some hint helps you
the research demonstrating rapid loss of short-term reconstruct it. For example, try naming all your high
memory has dealt almost entirely with meaningless school teachers. After you have named all you can,
material. How long would you remember the num- you will be able to name still more if someone shows
ber sequence 64-57-2-15 without constant rehearsal? you photos and tells you the teachers initials.
Olfactory bulb
Horizontal section
Thalamus through left
Olfactory tract posterior cerebral
cortex
Mamillary body
Hippocampus
Left temporal lobe
Hippocampus
(a) (b)
on anyone else. Sometimes strokes or other disorders The Hippocampus and Spatial Memory
damage the hippocampus, but seldom on both sides A second hypothesis is that the hippocampus is espe-
of the brain and never without also damaging other cially important for spatial memories. Electrical record-
nearby structures. Researchers therefore supplement ings indicate that many neurons in a rats hippocampus
the human studies by examining laboratory animals. are tuned to particular spatial locations, responding
best when an animal is in a particular place (OKeefe
The Hippocampus and & Burgess, 1996) or looking in a particular direction
Declarative Memory (Dudchenko & Taube, 1997; Rolls, 1996a). In a large
One hypothesis is that the hippocampus is critical enough environment, a given cell responds strongly
for declarative memory, especially episodic memory in several locations (Hafting, Fyhn, Molden, Moser, &
(Squire, 1992). How could we test this hypothesis with Moser, 2005). Ordinarily, a given cell responds in the
nonhumans, who cannot declare anything? What same way whenever the rat is in a particular environ-
could they do that would be the equivalent of describ- ment. If we move the rat to a new environment or
ing an episode? change the current environmentfor example, widen-
Here is one attempt to examine episodic memory: ing the cagethe various cells establish new patterns
A rat digs food out of five piles of sand, each with a dif- of response to different locations. That is, they re-map
ferent odor. Then it gets a choice between two of the their new environment (Leutgeb et al., 2005; Moita,
odors and is rewarded if it goes toward the one it Rosis, Zhou, LeDoux, & Blair, 2004). Younger rats re-
smelled first. Intact rats learn to respond correctly, ap- adjust their hippocampal maps faster than older rats
parently demonstrating memory of not only what they and more quickly learn to find their way to important
smelled but also when they smelled it. Memory of a places within the environment (Rosenzweig, Redish,
specific event qualifies as episodic, at least by a broad McNaughton, & Barnes, 2003). When people perform
definition. Rats with hippocampal damage do poorly spatial tasks, such as imagining the best route between
on this task (Fortin, Agster, & Eichenbaum, 2002; Kes- one friends house and another, fMRI results show en-
ner, Gilbert, & Barua, 2002). hanced activity in the hippocampus (Kumaran & Ma-
In the delayed matching-to-sample task, an ani- guire, 2005). All these results suggest that the hippo-
mal sees an object (the sample) and then, after a delay, campus might be particularly important for spatial
gets a choice between two objects, from which it must memory.
choose the one that matches the sample. In the de- In one study, researchers conducted PET scans on
layed nonmatching-to-sample task, the procedure is the brains of London taxi drivers as they answered ques-
the same except that the animal must choose the ob- tions such as, Whats the shortest legal route from the
ject that is different from the sample (Figure 13.7). In Carlton Tower Hotel to the Sherlock Holmes Museum?
both cases, the animal must remember which object (London taxi drivers are well trained and answer with
was present on this occasion, thereby showing what impressive accuracy.) Answering these route questions
we might call a declarative mem-
ory, perhaps an episodic memory.
Hippocampal damage strongly im-
pairs performance in many cases
(Zola et al., 2000). Unfortunately Delay
for theorists, performance varies
enormously with changes in pro-
cedure. For example, if the de-
layed matching- (or nonmatching-)
to-sample task uses constantly
changing varieties of objects, then
hippocampal damage greatly im-
Monkey lifts sample object to get food. Food is under the new object.
pairs performance. However, if re-
searchers use the same two objects Figure 13.7 Procedure for delayed nonmatching-to-sample task
Size of Hippocampus
Habitat Relative to Rest of Brain Spatial Memory Color Memory
Pinyon jay Lives at fairly high altitude; Second largest Second best Slightly better
depends on stored food
to survive the winter.
Scrub jay Stores some food but Smaller Less good Slightly worse
less dependent on it.
Tom Vezo/The Wildlife Collection
Mexican jay Stores some food but Smaller Less good Slightly better
less dependent on it.
Tangle
Sensitization in Aplysia
If you experience an unexpected, intense pain, you
Much of the research on Aplysia concerns changes probably react more strongly than usual to loud sounds,
in behavior as a result of experience. Some of those sharp pinches, and other sudden stimuli in the next few
changes may seem simple, and it is a matter of defini- days. This phenomenon is sensitization, an increase
tion whether we call them learning or use the broader in response to mild stimuli as a result of previous ex-
term plasticity. One commonly studied behavior is the posure to more intense stimuli. Similarly, a strong stim-
withdrawal response: If someone touches the siphon, ulus almost anywhere on Aplysias skin can intensify
mantle, or gill of an Aplysia (Figure 13.17), the animal later withdrawal responses to a touch.
vigorously withdraws the irritated structure. Investi- Researchers have traced sensitization to changes at
gators have traced the neural path from the touch re- identified synapses (Cleary, Hammer, & Byrne, 1989;
Dale, Schacher, & Kandel, 1988; Kandel & Schwartz, of the action potential; the exit of potassium restores
1982), as shown in Figure 13.19. Strong stimulation the neuron to its usual polarization. When serotonin
anywhere on the skin excites a particular facilitating blocks the potassium channels, the effect is a pro-
interneuron, which releases serotonin (5-HT) onto the longed action potential in the presynaptic cell and
presynaptic terminals of many sensory neurons. These therefore more transmitter release. If the sensitizing
are called presynaptic receptors. When serotonin at- stimulus occurs repeatedly, the sensory neuron syn-
taches to these receptors, it closes potassium channels thesizes new proteins that produce long-term sensiti-
in the membrane. As you will zation (C. H. Bailey, Giustetto, Huang, Hawkins, &
recall from Chapter 2, po- Kandel, 2000).
tassium flows out of the Facilitating
The research on Aplysia shows us that changes
neuron after the peak interneuron in synaptic activity can produce behavioral plas-
ticity, and in 2000, Eric Kandel won a Nobel
Prize for this work. The Aplysia research
Input from skin
receptors that were
also shows that even sensitization, widely
strongly stimulated regarded as a fairly simple process,
Excitatory
interneuron
depends on interactions among sev-
eral neurons and their synapses
Facilitating interneuron releases (Bailey et al., 2000).
serotonin at presynaptic receptors.
Result is to block potassium channels and Muscle that
thus prolong release of neurotransmitter retracts gill
Sensory Motor
neuron neuron
Gill
Siphon
Na+ and
Much Na+ Ca++ enter Much Na+
enters enters
Dendrite, much depolarized
Presynaptic Changes
The changes just described occur in the post- Figure 13.22 One way in which LTP occurs
synaptic neuron. In many cases, LTP depends In some cases, the dendrite makes new branches, which attach to
on changes in the presynaptic neuron instead or branches of the same axon, thus increasing the overall stimulation.
in addition. Extensive stimulation of a postsyn- (Source: Based on Toni, Buchs, Nikonenko, Bron & Muller, 1999)
Chapter Ending
Key Terms and Activities
Terms
Alzheimers disease (p. 399) consolidation (p. 397) implicit memory (p. 392)
amnesia (p. 389) cooperativity (p. 408) Korsakoffs syndrome (p. 398)
AMPA receptor (p. 408) declarative memory (p. 392) lateral interpositus nucleus (LIP)
amyloid beta protein 42 (Ab42) delayed matching-to-sample task (p. 386)
(p. 400) (p. 394) long-term depression (LTD) (p. 408)
anterograde amnesia (p. 390) delayed nonmatching-to-sample long-term memory (p. 387)
associativity (p. 408) task (p. 394) long-term potentiation (LTP)
central executive (p. 389) delayed response task (p. 389) (p. 408)
classical conditioning (p. 384) engram (p. 385) mass action (p. 386)
conditioned response (CR) episodic memory (p. 390) Morris water maze task (p. 395)
(p. 384) equipotentiality (p. 386) NMDA receptor (p. 409)
conditioned stimulus (CS) (p. 384) explicit memory (p. 392) operant conditioning (p. 384)
confabulation (p. 399) habituation (p. 406) phonological loop (p. 389)
configural learning (p. 397) Hebbian synapse (p. 405) plaque (p. 400)
Suggestion for
Further Reading http://www.thomsonedu.com
Eichenbaum, H. (2002). The cognitive neuroscience Go to this site for the link to ThomsonNOW, your one-stop study
of memory. New York: Oxford University Press. shop, Take a Pre-Test for this chapter, and ThomsonNOW will
Thoughtful treatment of both the behavioral and generate a Personalized Study Plan based on your test results.
physiological aspects of memory. The Study Plan will identify the topics you need to review and
direct you to online resources to help you master these topics.
You can then take a Post-Test to help you determine the con-
cepts you have mastered and what you still need work on.
Websites
to Explore
You can go to the Biological Psychology Study
Center and click these links. While there, you
can also check for suggested articles available
on InfoTrac College Edition. The Biological Psychol-
ogy Internet address is:
http://psychology.wadsworth.com/book/kalatbiopsych9e/
Exploring Biological
Psychology CD
Classical Conditioning (video)
Amnesia and Different Types of Memory (video)
Amnestic Patient (video)
Implicit Memories (Try It Yourself)
Alzheimers Patient (video)
Long-Term Potentiation (Try It Yourself)
Neural Networks and Me (video)
Chapter Quiz (multiple-choice questions)
Critical Thinking (essay questions)
B
Dyslexia
In Closing: Language and the Brain iological explanations of vision, hearing, and
Summary movement are fairly detailed. Explanations of
Answers to Stop & Check Questions motivations, emotions, and memory are less precise,
Thought Questions mainly because researchers can less precisely describe
or measure the behaviors themselves. Language,
Module 14.3 thought, and attention are again difficult to measure,
Attention much less explain physiologically. Nevertheless, they
Alterations in Brain Responses have been integral topics for neuroscience since its ear-
Neglect liest days, beginning with Paul Brocas report in the
Attention-Deficit Hyperactivity Disorder 1860s that speech depends on part of the left frontal
In Closing: Attending to Attention cortex.
Summary Although research on the biology of cognition is
Answers to Stop & Check Questions difficult, many of the results are fascinating. After dam-
Thought Question age to the corpus callosum, which connects the two
hemispheres, people act as if they have two fields of
Terms awarenessseparate minds, you might say. With
Suggestions for Further Reading damage to certain areas of the left hemisphere, people
Websites to Explore lose their language abilities, while remaining unim-
Exploring Biological Psychology CD paired in other ways. People with damage to parts of
ThomsonNOW the right hemisphere ignore the left side of their body
and the left side of the world. Studies of such people
offer clues about how the brain operates and raise
stimulating questions.
Opposite: Language may have evolved from our tendency
to make gestures.
Source: Daly & Newton/Getty Images
415
Module 14.1
Lateralization of Function
Eric Schrader/www.phutu.com
J P Fruchet/Getty Images
(a) (b)
Figure 14.2 Two views of the corpus callosum The auditory system is organized differently. Each
The corpus callosum is a large set of axons conveying ear sends the information to both sides of the brain be-
information between the two hemispheres. (a) A sagittal cause any part of the brain that contributes to localiz-
section through the human brain. (b) A dissection (viewed ing sounds must receive input from both ears. How-
from above) in which gray matter has been removed to ever, when the two ears receive different information,
expose the corpus callosum. each hemisphere does pay more attention to the ear
on the opposite side (Hugdahl, 1996).
Olfactory
bulbs
Optic
chiasm
Blood
To left Optic To right vessels
(a) (b)
HATBAND
Hippocampus
Anterior commissure
Hippocampal commissures
Corpus callosum
Thalamus
bookturn three pagesthen somehow the left likely to say it was himself. When he saw it in the left
hand will pick up two pages and youre at page 5, visual field (right hemisphere), he usually thought it
or whatever. Its better to let it go, pick it up with was the other person (Turk et al., 2002).
the right hand, and then turn to the right page. In other situations, the hemispheres learn to co-
With your right hand, you correct what the left operate. A split-brain person who was tested with
has done. (p. 211) the apparatus shown in Figure 14.4 used an interest-
ing strategy to answer a yesno question about what
Such conflicts are more common soon after surgery he saw in the left visual field. Suppose an experi-
than later. The corpus callosum does not heal, but the menter flashes a picture in the left visual field and
brain learns to use the smaller connections between asks, Was it green? The left (speaking) hemisphere
the left and right hemispheres (Myers & Sperry, 1985). takes a guess: Yes. That guess might be correct. If
The left hemisphere somehow suppresses the right not, the right hemisphere, which knows the correct
hemispheres interference and takes control in some answer, makes the face frown. (Both hemispheres con-
situations. However, even then, the hemispheres show trol facial muscles on both sides of the face.) The left
differences of opinion if we test carefully enough. hemisphere, feeling the frown, says, Oh, Im sorry, I
In one study, researchers used computers to morph meant no.
photos to look partly like a split-brain person and partly In another experiment, a split-brain patient saw
like another familiar person. Then they asked this split- two words flashed at once, one on each side. He was
brain person to identify each photo, after viewing it then asked to draw a picture of what he had read. Each
briefly in one visual field or the other. When he saw it hemisphere saw a full word, but the two words could
in the right visual field (left hemisphere), he was more combine to make a different word. For example,
(a) (b)
Ornstein (1997), the left hemisphere focuses more on though the effects are small. Here is something you can
details and the right hemisphere more on overall pat- try yourself: Tap with your right hand as many times
terns. For example, in one study, people with intact as you can in a short period of time. Rest and repeat
brains examined visual stimuli such as the one in Fig- with your left hand. Then repeat with each hand while
ure 14.8, in which many repetitions of a small letter talking. The Online Try It Yourself activity Hemisphere
compose a different large letter. When they were asked Control will keep track of your totals. For most right-
to identify the small letters (in this case, B), activity handers and many left-handers, talking decreases the
increased in the left hemisphere, but when they were tapping rate with the right hand more than O N L I N E
asked to identify the large overall letter (H), activity with the left hand (Kinsbourne & McMur-
was greater in the right hemisphere (Fink et al., 1996). ray, 1975). Evidently, it is more difficult to try it
do two things at once when both activities yourself
depend on the same hemisphere.
B B
B B
B B STOP & CHECK
B B
BB B B B BB
7. Which hemisphere is dominant for each of the fol-
B B lowing in most people: speech, emotional inflection
B B of speech, interpreting other peoples emotional
B B expressions, spatial relationships, perceiving overall
B B patterns?
Check your answers on page 428.
Figure 14.8 Stimulus to test analytical and
holistic perception
When people were told to name the large composite letter,
they had more activity in the right hemisphere. When told
to name the small component letters, they had more activity Development of Lateralization
in the left hemisphere. (Source: Based on Fink, Halligan, et al., and Handedness
1996)
Because most peoples language depends primarily on
Hemispheric Specializations the left hemisphere, it is natural to ask whether the
hemispheres differ anatomically. If so, is the differ-
in Intact Brains ence present before speech develops or does it develop
The differences between the two hemispheres also can later? What is the relationship between handedness
be demonstrated in people without brain damage, al- and hemispheric dominance for speech?
Anterior
Planum
temporale
Planum
Location of cut temporale
Left Right
Posterior
Answers to
STOP & CHECK
Questions
1. According to Klars hypothesis, both twins have
two genes for random handedness. Although one
of them randomly became right-handed and the
other left-handed, they will both pass their random
handedness genes to their children, and both twins
should have an equal percentage of left-handed chil-
dren. The data confirm that prediction (Klar, 2005).
(p. 417)
Common Chimpanzees
After many unsuccessful attempts to teach chimpanzees
to speak, researchers achieved better results by teach-
ing them American Sign Language or other visual sys-
Courtesy of Ann Premack
Duane Rumbaugh
Rumbaugh, Duane Rumbaugh,
and their associates tried to teach
a female bonobo named Matata
to press symbols that lit when Figure 14.11 Language tests for Kanzi, a bonobo (Pan paniscus)
touched; each symbol represents He listens to questions through the earphones and points to answers on a board.
a word (Figure 14.11). Although The experimenter with him does not know what the questions are or what answers
Matata made little progress, her are expected. (Source: From Georgia State Universitys Language Research Center, operated
infant son Kanzi learned just with the Yerkes Primate Center of Emory.)
by watching her. When given a
chance to use the symbol board, he
quickly excelled. Soon researchers noticed that Kanzi haps bonobos have more language potential than com-
understood a fair amount of spoken language. For ex- mon chimpanzees. A second explanation is that Kanzi
ample, whenever anyone said the word light, Kanzi and Mulika began language training when young, un-
would flip the light switch. By age 512, he understood like the chimpanzees in most other studies. A third
about 150 English words and could respond to such reason pertains to the method of training: Perhaps learn-
unfamiliar spoken commands as Throw your ball in ing by observation and imitation promotes better under-
the river and Go to the refrigerator and get out a standing than the formal training methods of previous
tomato (Savage-Rumbaugh, 1990; Savage-Rumbaugh, studies (Savage-Rumbaugh et al., 1992).
Sevcik, Brakke, & Rumbaugh, 1992). Since then, Kanzi For more information about bonobos, visit this web-
and his younger sister have demonstrated language site: http://www.blockbonobofoundation.org/blinks.htm
comprehension comparable to that of a 2- to 212-year-
old child (Savage-Rumbaugh et al., 1993):
They understand more than they can produce. STOP & CHECK
They use symbols to name and describe objects even
when they are not requesting them.
They request items that they do not see, such as bub- 1. How does common chimpanzees use of symbols
bles (I want to play with the bubble-blower) or car differ from language?
trailer (drive me in the car to the trailer). 2. What are three likely explanations for why the bono-
They occasionally use the symbols to describe past bos made more language progress than common
events. Kanzi once pressed the symbols Matata bite chimpanzees?
to explain the cut that he had received on his hand Check your answers on page 441.
an hour earlier.
They frequently make original, creative requests,
such as asking one person to chase another while
he watches. Nonprimates
Why have Kanzi and Mulika developed such im- What about nonprimate species? Elephants learn to im-
pressive skills where other chimpanzees failed? Per- itate the sounds they hear, including the vocalizations
David Carter
use sounds meaningfully. She kept Alex
in a stimulating environment and taught
him to say words in conjunction with spe- Figure 14.12 Language tests for Alex, an African gray parrot
cific objects. First, she and the other train- Alex has apparently learned to converse about objects in simple English
ers would say a word many times and for example, giving the correct answer to What color is the circle? He
then offer rewards if Alex approximated receives no food rewards.
the same sound. Here is an excerpt from
a conversation with Alex early in train-
ing (Pepperberg, 1981, p. 142): Relying on language is not always helpful. Pepper-
berg put Alex and three other gray parrots on perches;
Pepperberg: Pasta! (Takes pasta.) Pasta! (Alex stretches each had a chain of large plastic links from the perch
from his perch, appears to reach for pasta.) to an almond on the bottom. (Almonds are favorite
Alex: Pa! foods for parrots.) Alex and another language-trained
Pepperberg: Better . . . what is it? parrot repeatedly told the experimenter, Want nut.
When she declined to bring it to them, they gave up.
Alex: Pah-ah.
Two other parrots, untrained in language, used their
Pepperberg: Better! claws to pull up the chain until they reached the al-
Alex: Pah-ta. mond (Pepperberg, 2004) (Figure 14.13).
Pepperberg: Okay, heres the pasta. Good try. What do we learn from studies of nonhuman lan-
guage abilities? At a practical level, we gain insights
Although pasta was used in this example, Pepperberg into how best to teach language to those who do not
generally used toys. For example, if Alex said paper, learn it easily, such as people with brain damage or
wood, or key, she would give him what he asked children with autism. At a more theoretical level, these
for. In no case did she reward him with food for say- studies point out the ambiguity of our concept of lan-
ing paper or wood. guage: We cannot decide whether chimpanzees or par-
Alex gradually learned to give spoken answers to rots have language unless we define language more
spoken questions. He was shown a tray of 12 small ob- precisely.
jects and then asked such questions as What color is Because bonobos show some potential for language
the key? (answer: green) and What object is gray? learning, apparently human language evolved from a
(answer: circle). In one test, he correctly answered precursor that was present in the ancient ancestor from
39 of 48 questions. Even many of his incorrect answers which both species derived. What we still dont know
were almost correct. In one case, he was asked the is what that precursor ability was doing. Do bonobos,
color of the block and he responded with the color of and perhaps other primates, have language-type abili-
the rock (Pepperberg, 1993). He also can answer ques- ties that they use for communication? Perhaps so; the
tions of the form How many blue key? in which he ability for spoken language probably evolved from early
has to examine objects, count the blue keys among ob- communication by gestures (Corballis, 1999). (Gestures
jects of two shapes and two colors, and then say the are still important for communication; ask someone
answer, ranging from one to six (Pepperberg, 1994). what a spiral is and then watch their hands move!)
And what an elephant is, it is one of the Figure 14.14 A drawing and
animals. And what the elephant does, it a description of an elephant
lives in the jungle. It can also live in the by a young woman with Williams
zoo. And what it has, it has long gray
ears, fan ears, ears that can blow in the syndrome
wind. It has a long trunk that can pick The labels on the drawing were provided
up grass, or pick up hay . . . If theyre by the investigator based on what the
in a bad mood it can be terrible . . . If
woman said she was drawing. (Source:
the elephant gets mad it could stomp; it
could charge, like a bull can charge. From Williams syndrome: An unusual
They have long big tusks. They can neuropsychological profile, by U. Bellugi, P. P.
damage a car . . . it could be dangerous. Wang, and T. L. Jernigan, In S. H. Broman and
When theyre in a pinch, when theyre
in a bad mood it can be terrible. You J. Grafman, Eds., Atypical Cognitive Deficits in
dont want an elephant as a pet. You Developmental Disorders. Copyright 1987
want a cat or a dog or a bird . . . Lawrence Erlbaum. Reprinted by permission.)
Brocas aphasia Very poor Mostly nouns and verbs; omits prepositions Impaired if the meaning depends
and other grammatical connectives on complex grammar
Wernickes aphasia Unimpaired Grammatical but often nonsensical; has Seriously impaired
trouble finding the right word, especially
names of objects
Correct identification
mal readers and for people with dyslexia.
For people with this abnormality,
of letters
an effective treatment might be to teach 50
them to attend to just one word at a time.
Some children and adults with dyslexia
have been told to place over the page that
they are reading a sheet of paper with a
window cut out of it that is large enough
to expose just one word. In 3 months, 15
dyslexic children improved their read- 10 0 10
Left Distance from fixation point Right
ing skills by 1.22 grade levels (Geiger,
Lettvin, & Fahle, 1994). Four dyslexic Figure 14.17 Identification of a letter at various distances from
adults also made spectacular progress; the fixation point
one advanced from a third-grade to a Normal readers identify a letter most accurately when it is closest to the
tenth-grade reading level in 4 months fixation point, and their accuracy drops steadily as letters become more
(Geiger et al., 1992). After about the remote from that point. Many people with dyslexia show a small impairment
first 3 weeks of practice, they no longer for letters just to the right of the fixation point, yet they are substantially
needed the special cut-out sheet of paper. more accurate than normal readers are in identifying letters 5 to 10 degrees
One final twist: Of the four adults to the right of fixation. (Source: Reprinted from Task-Determined Strategies of
with dyslexia who went through this Visual Process, by G. Geiger, J. Y. Lettvin, & U. Zegarra-Moran, 1992, Cognitive Brain
process, three decided that they would Research, 1, pp. 3952, 1992, with kind permission of Elsevier Science-NL, Sara
rather return to being dyslexic! While Burgerhartstraat 25, 1055 KV Amsterdam, The Netherlands.)
dyslexic, they could attend to several
tasks at once, such as talking to some-
one, listening to news on the radio, creating a work of reasons. Language is not simply a by-product of over-
art, and so forth. When they learned to read one word all intelligence, but it is hardly independent of other
at a time, they found themselves able to perform only intellectual functions either.
one task at a time, and they missed their old way of
life. In short, their reading skills were tied to their
overall attentional strategies.
For more information about dyslexia, visit this
Summary
website: http://www.bda-dyslexia.org.uk 1. Chimpanzees can learn to communicate through
gestures or nonvocal symbols, although their out-
put does not closely resemble human language.
STOP & CHECK Bonobos have made more language progress than
common chimpanzees because of species differ-
ences, early onset of training, and different train-
10. What are four hypotheses about the causes of ing methods. (p. 429)
dyslexia? 2. An African gray parrot has shown surprising lan-
Check your answer on page 441. guage abilities, with a brain organized differently
from that of primates. (p. 430)
3. The hypothesis that language emerged as a by-
product of overall intelligence or brain size faces
major problems: Some people have full-sized brains
Module 14.2 but impaired language, and people with Williams
In Closing: Language and the Brain syndrome have nearly normal language despite men-
Perhaps the best summary of dyslexia is also the best tal retardation. (p. 432)
summary of language impairments in general: Lan- 4. People are specialized to learn language easily, but
guage and reading are sufficiently complicated that the nature of that specialization is not yet clear.
people can become impaired in many ways for many (p. 434)
Terms
anomia (p. 437) epilepsy (p. 419) productivity (p. 429)
anterior commissure (p. 426) focus (p. 419) Rasmussens encephalopathy
aphasia (p. 435) hippocampal commissure (p. 426) (p. 427)
attention-deficit hyperactivity language acquisition device spatial neglect (p. 443)
disorder (ADHD) (p. 444) (p. 434) split-brain people (p. 420)
attentional blink task (p. 445) lateralization (p. 417) stop signal task (p. 445)
Brocas aphasia (p. 435) nonfluent aphasia (p. 435) visual field (p. 418)
Brocas area (p. 435) object naming latency task Wada Test (p. 421)
choice delay task (p. 445) (p. 421) Wernickes aphasia (p. 437)
corpus callosum (p. 417) optic chiasm (p. 418) Wernickes area (p. 437)
dichotic listening task (p. 421) planum temporale (p. 425) Williams syndrome (p. 433)
dyslexia (p. 438) poverty of the stimulus argument
(p. 434)
A
In Closing: The Biology of Mood Swings
Summary re mental illnesses really illnesses, analogous to
Answers to Stop & Check Questions tuberculosis or influenza? Or are they normal
Thought Question reactions to abnormal experiences? They are not ex-
actly either. They are outcomes that combine biologi-
Module 15.3 cal predispositions with experiences, and to control
Schizophrenia them, we need a good understanding of both aspects.
Characteristics In this chapter, the emphasis is strongly on the
Genetics biological components of mental illnesses; Biological
The Neurodevelopmental Hypothesis Psychology is, after all, the title of the book. But this
Neurotransmitters and Drugs emphasis does not imply that other influences are
In Closing: The Fascination of Schizophrenia unimportant.
Summary
Answers to Stop & Check Questions
Thought Questions
Terms
Suggestions for Further Reading
Websites to Explore
Exploring Biological Psychology CD
ThomsonNOW
451
Module 15.1
Substance Abuse
and Addictions
Sensitization of the
Nucleus Accumbens
As people become addicted to something, it dominates
their attention, and the nucleus accumbens responds
more strongly to it. That is, the nucleus accumbens
becomes sensitized. Repeated use of cocaine increases
its ability to release dopamine in the nucleus accum-
bens, also its ability to activate part of the right pre-
frontal cortex, and the individuals tendency to seek
the drug (Robinson & Berridge, 2001; Volkow et al.,
2005). Meanwhile, the person responds less than nor-
mal to other incentives, including sex. According to
one hypothesis, the prefrontal cortex ordinarily sends
stimulatory input to the nucleus accumbens to sup-
port any potentially reinforcing activity. However, re-
peated drug use increases the background inhibition
in the prefrontal cortex, such that only the strongest
stimuli (i.e., the addictive substances) can get through.
comes. Gambling and video games occupy peoples at- Everything else is filtered out (Kalivas, Volkow, & Sea-
tention by the hour, but the participants usually dont mans, 2005).
smile. Many drug addicts say that the drug no longer When and how does the brain become sensitized
provides much pleasure, even though they continue to an addictive substance? Researchers gave rats an
to be obsessed with obtaining it. opportunity to learn to press a lever to inject them-
To account for these observations, Kent Berridge selves with heroin. Then they let some of the rats self-
and Terry Robinson (1998) distinguished between administer heroin during a withdrawal state, while
liking and wanting. According to their view, activ- others went through withdrawal without any heroin.
ity in the nucleus accumbens relates to wanting. To de- At a later time, when rats went through withdrawal a
velop an addiction is to increase how much you want second time, all the rats had an opportunity to press a
something, not necessarily how much you like it. lever to try to get heroin, although at this time the lever
Something you want monopolizes your attention. Ad- was inoperative. Although both groups of rats pressed
dictive drugs have a tremendous ability to dominate a the lever, those that had self-administered heroin dur-
users attention and cravings, even when the drug ex- ing the previous withdrawal state pressed far more fre-
perience is not pleasant (Berridge & Robinson, 1995). quently (Hutcheson, Everitt, Robbins, & Dickinson,
To evaluate this idea, researchers studied mice 2001). Evidently, receiving an addictive drug during a
with a mutation that increases and prolongs the effects withdrawal period is a powerful experience that pro-
of dopamine. To test how much the mice like sweet duces sensitization. In effect, the userrat or human
tastes, researchers watched the facial expressions. Mice learns that the drug relieves the distress caused by drug
make distinctive mouth movements when they eat withdrawal and produces heightened effects at that
something tastynot exactly a smile, but a distinctive time. Both humans and rats that have abstained from a
expression nevertheless. The mice with increased drug show heightened seeking of the drugi.e., crav-
dopamine showed no more expression of liking than ingduring periods of stress or after any reminder of
did other mice. However, when they had an opportu- the drug (Ciccocioppo, Martin-Fardon, & Weiss, 2004;
nity to run down an alley to get a sweet food, they left Ghitza, Fabbricatore, Prokopenko, Pawlak, & West,
the start box faster than other mice, paused less in 2003; Kruzich, Congleton, & See, 2001).
Genetics
Recall that early-onset Parkinsons disease has a clear STOP & CHECK
genetic basis; late-onset Parkinsons disease does not.
The same is true for Alzheimers disease. With Hunt-
5. Which type of alcoholism has a stronger genetic
ingtons disease, people with many C-A-G repeats on
basis? Which type has earlier onset?
the gene for the huntingtin protein develop symptoms
early in life, whereas those with fewer repeats develop 6. Name three ways a gene could influence alcoholism.
them later or not at all. Similarly, the genetic basis is Check your answers on page 458.
strong for early-onset alcoholism, especially in men.
Several Few or
who will none who
become will become
alcoholics alcoholics
later
James W. Kalat
7. What are two ways in which sons of alcoholics differ,
on average, from sons of nonalcoholics?
Check your answer on page 458.
Figure 15.4 Robin Kalat (the authors teenage
daughter) finds an alcohol vending machine in
Tokyo in 1998
Restrictions against buying alcohol were traditionally weak
in a country where most people cannot quickly metabolize
Medications to Combat acetaldehyde and therefore drink alcohol only in moderation.
Substance Abuse However, in 2000, Japan banned public alcohol vending
machines.
Many people who are trying to overcome substance
abuse join Alcoholics Anonymous, Narcotics Anony-
mous, or similar organizations, and others see psycho- The drug disulfiram, which goes by the trade name
therapists. Because many people do not respond well Antabuse, antagonizes the effects of acetaldehyde de-
to any of these treatments, researchers have been seek- hydrogenase by binding to its copper ion. Like many
ing medications that might decrease the cravings. Many drugs, its effects were discovered by accident. The
possibilities are still in the experimental phase (Vocci, workers in one rubber-manufacturing plant found that
Acri, & Elkashef, 2005). Here we briefly consider Anta- when they got disulfiram on their skin, they developed
buse and methadone. a rash (Schwartz & Tulipan, 1933). If they inhaled it,
they couldnt drink alcohol without getting sick. Soon
therapists tried using disulfiram (Antabuse) as a drug,
Antabuse hoping that alcoholics would associate alcohol with
After someone drinks ethyl alcohol, enzymes in the illness and stop drinking.
liver metabolize it to acetaldehyde, a poisonous sub- Most studies find that Antabuse is only moder-
stance. An enzyme, acetaldehyde dehydrogenase, then ately effective (Hughes & Cook, 1997). When it works,
converts acetaldehyde to acetic acid, a chemical that it supplements the alcoholics own commitment to
the body can use as a source of energy: stop drinking. By taking a daily pill and imagining
the illness that could follow a drink of alcohol, the
Acetaldehyde
person reaffirms a decision to abstain. In that case, it
dehydrogenase
doesnt matter whether the pill really contains Ant-
Ethyl alcohol Acetaldehyde Acetic acid
abuse or not; someone who never drinks will not ex-
People with a weaker gene for acetaldehyde de- perience the threatened illness (Fuller & Roth, 1979).
hydrogenase metabolize acetaldehyde more slowly. Those who drink in spite of taking the pill become
If they drink much alcohol, they accumulate acetal- ill, but unfortunately, they are as likely to quit taking
dehyde, which can produce flushing of the face, in- the pill as to quit drinking alcohol. Antabuse treat-
creased heart rate, nausea, headache, abdominal pain, ment is more effective if friends make sure the person
impaired breathing, and damage to internal organs. As takes the pill daily (Azrin, Sisson, Meyers, & Godley,
you might guess, people who cant metabolize acetal- 1982).
dehyde are unlikely to drink much alcohol. About half
of the people in China and Japan have a gene that slows
acetaldehyde metabolism; probably for that reason, al-
Methadone
cohol abuse has historically been uncommon in those Heroin is an artificial substance invented in the 1800s
countries (Tu & Israel, 1995) (Figure 15.4). In other as a safer alternative for people who were trying to
words, this gene affects alcohol abuse by altering liver quit morphine (an opiate drug). Some physicians even
enzymes. recommended that people using alcohol switch to
Probability of major
depression episode
Hudson et al., 2003).
The risk of depression is particularly elevated .30 s/l
among relatives of women with early-onset depres-
sionthat is, beginning before age 30 (Bierut et al., l/l
.20
1999; Kendler, Gardner, & Prescott, 1999; Lyons et al.,
1998). Compare this pattern to alcoholism, where the
.10
risk is highest among relatives of men with early-onset
alcoholism.
Although the predisposition to depression un- .00
0 1 2 3 4+
doubtedly depends on many genes, several have been
Number of stressful life events
identified. A particular form of one gene leads to an
80% decrease in the brains ability to produce the neuro- Figure 15.6 Genetics, stress, and depression
transmitter serotonin. A study found that gene in 9 of The probability of depression increases for people
87 people with major depression, and only 3 of 219 peo- reporting higher numbers of stressful experiences in
ple in a control group. Furthermore, 2 of those 3 had the previous 5 years; however, the rate of increase
mild depression (X. Zhang et al., 2005). Although it depends on their genetics. (Reprinted with permission
is clear that most people with depression do not have from A. Caspi, et al., Influence of life stress on depression:
this gene, people who do have it are at high risk for Moderation by a polymorphism in the 5-HTT gene, Science,
depression. 301, pp. 386389 (15). 2003 AAAS)
Another gene of interest is the one controlling the
serotonin transporter protein (Pezawas et al., 2005).
That protein controls the ability of an axon to reabsorb
serotonin after its releasethat is, the ability to recycle
Hormones
it for further use. The effect of that gene on depression Depression occurs in episodes rather than constantly.
interacts with peoples experiences. Investigators ex- It lasts months, alleviates for months or years, and then
amined the serotonin transporter genes of 847 people, returns. One likely trigger is stress, which releases cor-
identifying them in terms of two types: the short tisol, as described in Chapter 12. Cortisol readies the
type and the long type. They also asked each partici- body for action, but prolonged high levels can exhaust
pant to record certain highly stressful events from age the bodys energies, impair sleep, impair the immune
21 until age 26. Those events included financial set- system, and set the stage for an episode of depression.
backs, changes of job or housing, divorce, and so forth. The role of sex hormones is less certain. Most
Figure 15.6 shows the results. For people with two women feel some emotional distress for a day or two
short forms of the gene, increasing numbers of stress- after giving birth, and about 20% experience postpar-
ful experiences led to a big increase in the probability tum depressionthat is, depression after giving birth.
of depression. For those with two long forms, stress- Most women recover fairly quickly without treatment,
ful events hardly increased the risk of depression at but about 0.1% enter a serious, long-lasting depression
all. Those with one short and one long gene were in- (Hopkins, Marcus, & Campbell, 1984). Postpartum de-
termediate. In other words, the short form of the gene pression is more common among women who have also
by itself does not lead to depression, nor does a series suffered major depression at other times and among
of stressful events, but a combination of both is haz- women who experience severe discomfort around the
ardous (Caspi et al., 2003). The gene probably alters the time of menstruation (Bloch, Rotenberg, Koren, & Klein,
way people react to stress. 2005).
One study found that after a drug-induced drop in
estradiol and progesterone levels, women with a his-
tory of postpartum depression suddenly show new
STOP & CHECK symptoms of depression, whereas other women do not
(M. Bloch et al., 2000). That is, some women are more
vulnerable to depression than others, and hormonal
1. What is the relationship between depression and the changes can trigger an episode of depression for the
short form of the gene controlling the serotonin trans- vulnerable women. Another study found that estradiol
porter protein? supplements relieved depression in many middle-aged
Check your answer on page 469. women going through menopause (Soares, Almeida,
Joffe, & Cohen, 2001).
&
Bode & Ludwig, 1997
STOP CHECK
Bipolar Disorder
Image not available due to copyright restrictions
Depression can be either unipolar or bipolar. People
with unipolar disorder vary between normality and
one poledepression. People with bipolar disorder
formerly known as manic-depressive disorderalter-
nate between two polesdepression and its opposite,
mania. Mania is characterized by restless activity, ex-
citement, laughter, self-confidence, rambling speech,
and loss of inhibitions. Some manic people are dan-
gerous to themselves and others. Figure 15.12 shows
the brains increase in glucose use during mania and
its decrease during depression (Baxter et al., 1985).
People who have full-blown episodes of mania are
said to have bipolar I disorder. People with bipolar II
disorder have much milder manic phases, called hy-
pomania, which are characterized mostly by agitation
or anxiety. About 1% of people have at least a mild case
of bipolar disorder at some time in life, with an average
age of onset in the early 20s (Craddock & Jones, 1999).
In addition to the mood swings, most people with bi- control, and impairments of verbal memory (Quraishi
polar disorder have attention deficits, poor impulse & Frangou, 2002).
Body temperature
and SAD people 37.5
People with seasonal
Patients with SAD are phase- affective disorder
delayed while most other patients 37C
with depression are phase-advanced.
36.5
evening (Eastman, Young, Fogg, Liu, & Meaden, 1998; to definitely because it is difficult to do research on
Lewy et al., 1998; Terman, Terman, & Ross, 1998). The human moods. At each stage in life, brain structure and
most likely explanations are that bright light affects chemistry alter ones reactions to events, and events in
serotonin synapses and alters circadian rhythms. Re- turn alter the brain, affecting the reaction to the next
gardless of how it works, it produces substantial ben- event. Studying an adult brain has the same challenge
efits. Many therapists recommend bright light therapy as watching only a few minutes in the middle of a com-
for nonseasonal depression as well. Research on that plex movie and trying to guess how events led up to
possibility has been sparsepossibly because drug that point.
companies are not interested in sponsoring itbut
so far, the results indicate that bright light relieves
depression at least as well as drugs or psychotherapy
and possibly better. Bright light is less expensive than
Summary
the other therapies and produces its benefits more 1. People with major depression find that almost noth-
rapidly, often within 1 week (Kripke, 1998). Combin- ing makes them happy. Depression occurs as a se-
ing bright light with another form of therapy mag- ries of episodes. (p. 459)
nifies the benefits (Kripke, 1998; Loving, Kripke, &
2. Depression shows a strong family tendency, espe-
Shuchter, 2002).
cially for relatives of women with early-onset de-
The following website provides much information
pression. (p. 460)
about light therapy and biological rhythms: http://
www.sltbr.org/ 3. One gene has been shown to increase the proba-
bility of depression only among people who have
had stressful experiences. (p. 460)
4. Depression is associated with decreased activity
STOP & CHECK in the left hemisphere of the cortex. (p. 461)
5. Four kinds of antidepressant drugs are in wide use.
11. What are the advantages of light treatment compared Tricyclics block reuptake of serotonin and cate-
to antidepressant drugs? cholamines but produce strong side effects. SSRIs
Check your answer on page 469. block reuptake of serotonin. MAOIs block an en-
zyme that breaks down catecholamines and sero-
tonin. Atypical antidepressants are a miscella-
neous group with diverse effects. (p. 462)
6. The antidepressants alter synaptic activity quickly,
Module 15.2 but their effects on behavior build up over weeks.
In Closing: The Biology of Mood Swings (p. 463)
Do you feel sad because of events that have happened 7. The behavioral effects of antidepressant drugs prob-
to you or because of your brain chemistry? According ably depend on two slow changes in the brain. The
to biological psychologists, that is a meaningless ques- drugs increase the release of BDNF, which pro-
tion. Your experiences are changes in your brain state; motes neuron growth and survival. They also de-
you cannot have one without the other. The better ques- sensitize autoreceptors and thereby increase re-
tion is: Are some people more likely than others to be- lease of neurotransmitters. (p. 463)
come depressed because of a preexisting condition in 8. Other therapies for depression include psycho-
their brain structure or chemistry? The answer to that therapy, electroconvulsive therapy, and altered
one is probably. It is difficult to get from probably sleep patterns. (p. 464)
Uncles/aunts 2%
Adopted Children Who
Nephews/nieces 4% Develop Schizophrenia
Grandchildren 5% When an adopted child develops schizophrenia, it is
more common among his or her biological relatives
Half-siblings 6% than adopting relatives. One Danish study found schiz-
ophrenia in 12.5% of the immediate biological relatives
Children 13%
and none of the adopting relatives (Kety et al., 1994).
Children of schizophrenic mothers, Note in Figure 15.14 that children of a schizophrenic
adopted by nonschizophrenic mothers 17% mother have a moderately high probability of schizo-
Siblings 9% phrenia, even if adopted by nonschizophrenic parents.
These results suggest a genetic basis, but they are
DZ twins 17% also consistent with a prenatal influence. Consider a
pregnant woman with schizophrenia. True, she passes
Parents 6%
her genes to her child, but she also provides the pre-
MZ twins 48%
natal environment, which may be less than healthy.
Many women with schizophrenia have poor nutrition,
Children of two smoke and drink heavily, and fail to get medical care
schizophrenic parents 46%
during pregnancy. If their children develop schizophre-
nia, we cannot be sure that the influence is genetic.
Figure 15.14 Probabilities of
developing schizophrenia
People with a closer genetic relationship to someone
Efforts to Locate a Gene
with schizophrenia have a higher probability of developing There are reasons for skepticism about a strong role for
it themselves. (Source: Based on data from Gottesman, 1991) genetics. One of the biggest is that people with schizo-
and those in gray appeared normal in all studies van, Menon, Lim, & Pfefferbaum, 1997). Most patients
(Honea, Crow, Passingham, & Mackay, 2005). Note with schizophrenia show deficits of memory and atten-
that the strongest deficits were in the left temporal and tion similar to those of people with damage to the tem-
frontal areas of the cortex. Note also that most corti- poral or prefrontal cortex (Park, Holzman, & Goldman-
cal areas showed mild abnormalities in at least one or Rakic, 1995) (Methods 15.1).
two studies. At a microscopic level, the most reliable finding
Furthermore, the ventricles (fluid-filled spaces is that cell bodies are smaller than normal, especially
within the brain) are larger than normal in people with in the hippocampus and prefrontal cortex (Pierri,
schizophrenia (Wolkin et al., 1998; Wright et al., 2000) Volk, Auh, Sampson, & Lewis, 2001; Rajkowska, Sele-
(Figure 15.16). The increased size of the ventricles im- mon, & Goldman-Rakic, 1998; Selemon, Rajkowska, &
plies less space taken by brain cells. Signs of brain Goldman-Rakic, 1995; Weinberger, 1999).
damage are especially common in people who had a Lateralization also differs from the normal pattern.
history of complications during pregnancy or at birth In most people, the left hemisphere is slightly larger
(Stefanis et al., 1999). than the right, especially in the planum temporale of
The areas with consistent signs of abnormality the temporal lobe, but in people with schizophrenia,
include some that mature slowly, such as the dorso- the right planum temporale is equal or larger (Kasai
lateral prefrontal cortex (Berman, Torrey, Daniel, & et al., 2003; Kwon et al., 1999). People with schizo-
Weinberger, 1992; Fletcher et al., 1998; Gur, Cowell, phrenia have lower than normal overall activity in the
et al., 2000). As you might predict, people with schiz- left hemisphere (Gur & Chin, 1999) and are more likely
ophrenia perform poorly at working memory tasks, than other people to be left-handed (Satz & Green,
which depend on the prefrontal cortex (Goldberg, Wein- 1999). All these results suggest a subtle change in early
berger, Berman, Pliskin, & Podd, 1987; Spindler, Sulli- brain development.
METHODS 15.1
The Wisconsin Card Sorting Task
Neuropsychologists use many behavioral tests to mea- sort them by a different rule, such as number. Shifting to
sure the functioning of the prefrontal cortex. One is the a new rule requires suppressing the old one and evokes
Wisconsin Card Sorting Task. A person is handed a shuf- activity in the prefrontal cortex (Konishi et al., 1998).
fled deck of cards that differ in number, color, and shape People with damage to the prefrontal cortex can sort
of objectsfor example, three red circles, five blue tri- by whichever rule is first, but then they have trouble
angles, four green squares. First the person is asked to shifting to a new rule. People with schizophrenia have
sort them by one rule, such as separate them by color. the same difficulty.
Then the rule changes, and the person is supposed to
Terms
acetaldehyde (p. 456) dopamine hypothesis of schizophrenia (p. 470)
acetic acid (p. 456) schizophrenia (p. 478) seasonal affective disorder (SAD)
acute (p. 470) electroconvulsive therapy (ECT) (p. 467)
(p. 464) season-of-birth effect (p. 474)
alcoholism (or alcohol
dependence) (p. 454) glutamate hypothesis of second-generation antipsychotics
schizophrenia (p. 478) (p. 479)
Antabuse (p. 456)
hallucination (p. 470) selective serotonin reuptake
antipsychotic (or neuroleptic) drug
(p. 477) lithium (p. 467) inhibitors (SSRIs) (p. 462)
atypical antidepressants (p. 463) major depression (p. 459) self-stimulation of the brain
mania (p. 466) (p. 452)
bipolar disorder (or manic-
depressive disorder) (p. 466) mesolimbocortical system (p. 479) substance abuse (p. 452)
bipolar I disorder (p. 466) monoamine oxidase inhibitors substance-induced psychotic
(MAOIs) (p. 462) disorder (p. 478)
bipolar II disorder (p. 466)
negative symptom (p. 470) tardive dyskinesia (p. 479)
Borna disease (p. 461)
neurodevelopmental hypothesis thought disorder (p. 471)
butyrophenone (p. 477)
(p. 473) tricyclic (p. 462)
chlorpromazine (p. 477)
phencyclidine (PCP) (p. 478) Type I alcoholism (p. 454)
chronic (p. 470)
phenothiazine (p. 477) Type II alcoholism (p. 454)
concordance (p. 472)
positive symptom (p. 470) unipolar disorder (p. 466)
delusion (p. 470)
postpartum depression (p. 460)
differential diagnosis (p. 471)
Exploring Biological
Psychology CD An interview shows how depression robs someone of the joy
of living.
Understanding Addiction (video)
CNS Stimulants (animation)
Major Depressive Disorder: Barbara (two videos)
Antidepressant Drugs (animation)
Bipolar Disorder: Mary (three videos)
Bipolar Disorder: Etta (two videos)
Chapter Quiz (multiple-choice questions)
Critical Thinking (essay questions)
http://www.thomsonedu.com
Go to this site for the link to ThomsonNOW, your one-stop study
shop, Take a Pre-Test for this chapter, and ThomsonNOW will This video highlights research on nicotine addiction.
generate a Personalized Study Plan based on your test results.
know a little about chemistry. If you have taken a high Table A.1 The Elements That Compose
school or college course and remember the material Almost All of the Human Body
reasonably well, you should have no trouble with the Percentage by Weight
chemistry in this text. If your knowledge of chemistry Element Symbol in Human Body
is pretty hazy, this appendix will help. (If you plan to
take other courses in biological psychology, you should Oxygen O 65
study as much biology and chemistry as possible.) Carbon C 18
Hydrogen H 10
Nitrogen N 3
Elements and Compounds Calcium Ca 2
If you look around, you will see an enormous variety Phosphorus P 1.1
of materialsdirt, water, wood, plastic, metal, cloth, Potassium K 0.35
glass, your own body. Every object is composed of a
Sulfur S 0.25
small number of basic building blocks. If a piece of
wood catches fire, it breaks down into ashes, gases, Sodium Na 0.15
and water vapor. The same is true of your body. An in- Chlorine Cl 0.15
vestigator could take those ashes, gases, and water and Magnesium Mg 0.05
break them down by chemical and electrical means
Iron Fe Trace
into carbon, oxygen, hydrogen, nitrogen, and a few
other materials. Eventually, however, the investigator Copper Cu Trace
arrives at a set of materials that cannot be broken down Iodine I Trace
further: Pure carbon or pure oxygen, for example, can- Fluorine F Trace
not be converted into anything simpler, at least not by
Manganese Mn Trace
ordinary chemical means. (High-power bombardment
with subatomic particles is another story.) The matter Zinc Zn Trace
we see is composed of elements (materials that can- Selenium Se Trace
not be broken down into other materials) and com- Molybdenum Mo Trace
pounds (materials made up by combining elements).
485
Periodic Table of the Elements
Alkali
486
Period Metals Noble Gases
1 18
IA Alkaline VIIA
Earth
1 Metals Halogens 2
1 H He
hydrogen 2 13 14 15 16 17 helium
1.008 IIA IIIA IVA VA VIA VIIA 4.003
3 4 5 6 7 8 9 10
2 Li Be B C N O F Ne
lithium beryllium boron carbon nitrogen oxygen fluorine neon
6.941 9.012 10.81 12.011 14.007 16.0 18.999 20.179
11 12 Transition Elements 13 14 15 16 17 18
3 Na Mg Al Si P S Cl Ar
sodium magnesium 3 4 5 6 7 8 9 10 11 12 aluminum silicon phosphorous sulfur chlorine argon
22.99 24.305 IIIB IVB VB VIB VIIB VIIIB VIIIB VIIIB IB IIB 26.982 28.085 30.974 32.060 35.453 39.948
19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36
4 K Ca Sc Ti V Cr Mn Fe Co Ni Cu Zn Ga Ge As Se Br Kr
potassium calcium scandium titanium vanadium chromium manganese iron cobalt nickel copper zinc gallium germanium arsenic selenium bromine krypton
39.098 40.08 44.955 47.90 50.941 51.996 54.938 55.847 58.933 58.70 63.546 65.38 69.72 72.59 74.922 78.96 79.904 83.80
55 56 57 72 73 74 75 76 77 78 79 80 81 82 83 84 85 86
6 Cs Ba La Hf Ta W Re Os Ir Pt Au Hg Tl Pb Bi Po At Rn
cesium barium lanthanum hafnium tantalum tungsten rhenium osmium iridum platinum gold mercury thallium lead bismuth polonium astatine radon
132.905 137.33 138.906 178.49 180.948 183.85 186.207 190.20 192.22 195.09 196.967 200.59 204.37 207.20 208.980 (209) (210) (222)
87 88 89 104 105 106 107 108 109 110 111 112 113 114 115 116 117 118
7 Fr Ra Ac Rf Db Sg Bh Hs Mt Ds Rg Uub Uut Uuq Uup Uuh Uus Uuo
francium radium actinium rutherfordium dubnium seaborgium bohrium hassium meitnerium darmstadtium roentgenium ununbium ununtrium ununquadium ununpentium ununhexium ununseptium ununoctium
(223) 226.025 (227) (261) (262) (266) (264) (269) (268) (271) (272) (285) (284) (289) (288) (292) (?) (?)
58 59 60 61 62 63 64 65 66 67 68 69 70 71
Lanthanides Ce Pr Nd Pm Sm Eu Gd Tb Dy Ho Er Tm Yb Lu
6 cerium praseodymium neodymium prometheum samarium europium gadolinium terbium dysprosium holmium erbium thulium ytterbium lutetium
140.12 140.908 144.24 (145) 150.40 151.96 157.25 158.925 162.50 164.93 167.26 168.934 173.04 174.97
H H
HCH HCH HC CH
H H The Role of ATP
H H O H O H O H O
+H N C C N C C N C C N C C
H R H R H R H R O
(a protein)
O H H H H H H H H H H H H H H H H H
C C C C C C C C C C C C C C C C C C H
HO H H H H H H H H H H H H H H H H H
Stearic acid
(a fat)
HO Adenosine
NH2
OH C
HO N C N
CHCH2NHCH3 Epinephrine CH
Phosphates HC C N
HO N
O O O
Ribose
O P O P O P O CH2 O
HO CH2CH2NH2 Serotonin
(5-hydroxytryptamine) O O O C H H C
N H H
H C C
OH OH
OH O
AMP
CCH2CH2CHC Glutamate
ADP
OH NH3+OH
ATP
491
compliance with the Society Policy and are required General de Salud en Materia de Investigacion para la
to verify that they have done so when submitting ab- Salud of the Secretaria de Salud (published on Jan. 6,
stracts for presentation at the Annual Meeting or manu- 1987). Similarly, in addition to complying with the
scripts for publication in The Journal of Neuroscience. laws and regulations of their home countries, Foreign
Adherence to the Society Policy is also an important Members of the Society should adhere to the official
step toward receiving help from the Society in respond- Society Policy outlined here.
ing to questions about a members use of animals in
research. A complete description of the Societys policy
and procedures for defending members whose research Recommended References
comes under attack is given in the Societys Handbook
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and Use of Laboratory Animals (revised Sept. 1986).
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Office for Protection from Research Risks, NIH,
relevance to human or animal health, the advancement
6100 Executive Blvd., Suite 3B01-MSC 7507, Rock-
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ville, MD 20892-7507.
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Mexico must comply with the Reglamento de la Ley ber required to obtain valid results.
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Name Index
Abbar, M., 365 Almqvist, E., 259, 260 Arnold, A. P., 13, 163, 327 Barnes, B. M., 287
Abbott, D. F., 369 Alnwick, K. A., 334 Arnold, S. E., 476 Barnes, C. A., 394
AbdelMalik, P., 473 Alonso, J.-M., 174 Aron, A., 357 Barnes, J., 463
Abell, C. W., 454 Altar, C. A., 465 Arseneau, L. M., 297 Baron-Cohen, S., 369, 433
Abercrombie, H. C., 377 Amanzio, M., 212 Arvanitogiannis, A., 73 Baron, E., 159
Abi-Dargham, A., 478 Amaral, D. G., 398399 Arvidson, K., 216 Baron, R., 159
Aboitz, F., 438 Amateau, S. K., 328 sberg, M., 365 Barone, F. C., 139
Abraham, N. M., 221 Ambady, N., 368 Ascherio, A., 257 Barr, H. M., 454
Abraham, W. C., 408 Ambrosini, P. J., 446 Aserinsky, Eugene, 276 Barratt-Fornell, A., 219
Abrams, R. A., 389 Amedi, A., 132, 133 Asghar, S. J., 71 Barrot, M., 73
Abrams, W., 141 American Psychiatric Assal, G., 423 Bartel, P., 286
Aburada, M., 377 Association, 452, 459, 470 Aston-Jones, G., 271 Bartels, A., 179
Acri, J., 456 Ames, M. A., 346, 347 Astur, R. S., 330 Bartke, A., 328
Acuna-Goycolea, C., 314 Amidzic, O., 109 Attwell, D., 138 Bartko, J., 365
Adamec, R. E., 361 Amiry-Moghaddam, M., 36 Atzei, A., 143 Barton, L., 328
Adams, D. B., 333334 Amlaner, C. J., 287 Au, T. K., 434 Barton, R. A., 96, 97
Adams, L. F., 334 Amoore, J. E., 222 Audinat, E., 35 Bartoshuk, L. M., 212, 216,
Adams, R. B., Jr., 368 Amsterdam, J. D., 461 Auer, R. N., 138 217, 218
Adams, R. D., 399 Andersen, J. L., 235 Augath, M., 109 Barua, L. A., 394
Adams, W., 474 Andersen, R. A., 244 Auh, S., 475 Basbaum, A. I., 63, 210, 212
Ader, R., 378 Andersen, S. L., 73 Axel, R., 125 Basheer, R., 280
Adkins, E. K., 338 Andersen, T. S., 102 Axel, Richard, 222 Basil, J. A., 396
Adkins-Regan, E., 346 Anderson, A. K., 369 Ayers, G., 335 Basma, A., 256
Adler, E., 218 Anderson, A. W., 178 Azrin, N. H., 456 Bassett, J., 473
Adler, N. T., 331, 338 Anderson, C., 346 Bassetti, C. L., 277
Adolphs, R., 369, 370, 423 Anderson, D. J., 199 Babich, F. R., 405 Basso, A., 426
Adrien, J., 282 Anderson, L. A., 463 Babiloni, C., 210 Bastian, J., 44
Agerbo, E., 471 Anderson, M. E., 250 Babinet, C., 124 Bastien, C., 275
Aggleton, J. P., 394 Anderson, S. W., 359 Babkoff, H., 266 Bates, E., 436, 437
Aglioti, S., 143, 219, 417, 444 Andersson, D., 330 Babor, T. F., 454 Battersby, S., 309
Agster, K. L., 394 Andersson, K.-E., 332 Backlund, E.-O., 258 Baudry, M., 367
Agyei, Y., 345 Andreasen, N. C., 105, 329, Bckman, J., 288 Baulac, S., 419
Aharon, I., 452 470, 473, 476 Baddeley, A. D., 384, 389, 433 Baum, A., 379
Aharon, L., 452, 453 Andreasen, Nancy, 470, 471 Baer, J. S., 454 Baum, M. J., 224, 331
Ahlskog, J. E., 317 Andres, K. H., 206 Baghdoyan, H. A., 281 Bax, C. M., 334
Ahmed, I. I., 328 Andrew, D., 208, 213 Bailey, C. H., 407 Baxter, L. R., Jr., 466
Ahn, S. J., 310 Andrews, T. J., 166 Bailey, J. M., 345, 346, 347 Baxter, M. G., 369
Airaksinen, M. S., 129 Angelucci, A., 124, 155 Bailey, K. G. D., 437 Bayley, P. J., 392
Akbarian, S., 478 Angulo, M. C., 35 Baillet, S., 442 Baylis, G. C., 176, 177
Al-Rashid, R. A., 315 Annas, P., 371 Baird, A. A., 368 Baylor, D. A., 128
Albertini, R. J., 130 Antanitus, D. S., 35, 36 Baker, B. N., 199 Beal, M., 261
Alberts, J. R., 337 Antle, M. C., 272 Baker, G. B., 290 Bear, M. F., 185, 186
Albrecht, D. G., 176 Antoun, N., 358 Baker, J., 284 Bechara, A., 74, 359, 360
Albright, T. D., 30 Anzenberger, G., 335 Baker, L., 14 Beck, Max, 341
Aldinio, C., 141 Aosaki, T., 93 Bakker, J., 329 Becker, C., 371
Aldrich, M. S., 283 Apfelbach, R., 372 Balda, R. P., 396 Becker, H. C., 373
Aldridge, J. W., 453 Apostolakis, E. M., 331 Baldwin, A. E., 411 Becker, J. T., 116
Aleman, A., 471 Applebaum, S., 223 Ball, W. A., 281 Beebe, D. W., 283
Alerstam, T., 288 Appolinario, J. C., 319 Ballard, P. A., 256 Beeman, M. J., 420, 423
Alexander, A. L., 368 Araneda, R. C., 223 Balleine, B. W., 328 Beersma, G. M., 273
Alexander, J. T., 314 Arango, V., 365 Balthazart, J., 329 Behl, C., 331
Alkire, M. T., 114 Arduino, C., 212 Banich, M. T., 424 Behrmann, M., 178
Allen, C. B., 408 Arendt, D., 157, 417 Bao, S., 132, 133 Belin, P., 201
Allen, J. J. B., 395 Arendt, J., 271, 272 Baquet, Z. C., 129 Bell, E. C, 71
Allen, J. S., 115, 200, 329 Arey, L. B., 170 Bar, M., 176 Bello, N. T., 74
Allen, L. S., 348 Arkin, A. M., 284 Barash, S., 250 Bellugi, U., 433, 435
Alliger, R., 470 Arman, A. C., 226 Barbour, D. L., 200 Ben-Shoshan, R., 225
Allis, C. D., 270 Armony, J. L., 368 Barchas, J., 289 Bencsics, C., 138139
Allison, T., 178, 287 Armstrong-James, M., 131 Bargmann, C. I., 126 Benedetti, F., 212
Almazen, M., 433 Armstrong, S. M., 271 Barinaga, M., 138 Benes, F. M., 125, 476
Almeida, O. P., 460 Arnason, B. G. W., 258 Barker, P. B., 435 Benjamin, J., 68
Almli, C. R., 315 Arndt, S., 470 Barnea, G., 223 Benn, S. C., 129
550
Bennett, A. L., 347 Blzquez, C., 401 Bressler, K., 209 Buzski, G., 138
Bennett, E. L., 131 Bleuler, Eugen, 470 Bridgeman, B., 171 Byl, N. N., 135
Bennett, K., 340 Blindt, H. S., 394 Bridges, R. S., 335 Byne, W., 348, 349
Benoit, S. C., 314 Bliss, T. V. P., 408, 410 Brightman, M. W., 37 Byrne, J. H., 406
Benschop, R. J., 377 Bloch, G. J., 328, 331 Brinkmann, R. R., 259, 260 Byrne, M., 471
Benson, D. F., 101, 177 Bloch, M., 460 Broca, Paul, 109
Benson, M. D., 400 Blonder, L. X., 178 Brockmole, J. R., 181 Caamao-Isorna, F., 257
Berard, J., 364 Blossom-Stach, C., 437 Brodie, M. S., 71 Caccappolo-van Vliet, E., 439
Berdoy, M., 367 Blum, D., 24 Brody, A. L., 464 Cade, J. F., 467
Berenbaum, S. A., 342, 343 Blum, I., 334 Bron, C. R., 410 Cadoret, R. J., 362
Berger, R. J., 286 Blum, K., 67 Bronson, R. T., 335 Cagniard, B., 453
Berger-Sweeney, J., 123 Blumenthal, J., 446 Brooks, D. C., 281 Cahalan, M. D., 44
Berglund, H., 225 Boatman, D., 427 Brooks, D. J., 242, 256 Cahill, L., 398, 411, 417
Bergmann, B. M., 288 Bobrow, D., 346 Brooks, J. H., 363 Cahn, J., 141
Berkhof, J., 459 Bode, L., 461, 462 Brotchie, P., 244 Cahn, W., 476
Berliner, D. L., 224, 225 Boder, E., 438 Broughton, R., 284 Cai, X. J., 314
Berlucchi, G., 143, 219, 417, Boeve, B. F., 284 Brown, A. S., 474 Caicedo, A., 218
421 Bogaert, A. F., 347 Brown, G. L., 365 Cajochen, C., 271
Berman, A. J., 144 Bohbot, V. D., 395 Brown, G. M., 282 Calder, A. J., 358
Berman, K. F., 475 Bohlin, G., 342 Brown, J., 454 Caldwell, J. A., 267
Bernasconi, S., 364 Bohman, M., 454 Brown, J. R., 335 Caldwell, J. D., 335
Bernati, T., 443 Bohn, M. C., 258 Brown, M. J., 15 Calhoun, J., 443
Bernhardt, P. C., 363 Boivin, D. B., 273 Brown, T. L., 111 Calicott, J. H., 473
Bernstein, J. J., 140 Bolaos, C. A., 73 Brownell, K. D., 319 Calignano, A., 75
Berntsen, D., 398 Bonhoeffer, T., 185, 410 Browning, M. D., 410 Callaway, E. M., 179
Berntson, G. G., 278 Bonifati, V., 254 Bruce, Kristi, 341 Camargo, L. A. A., 304
Beron, J., 305 Bontempi, B., 397 Bruesch, S. R., 170 Cameron, H. A., 379
Berridge, C. W., 278 Bookheimer, S. Y., 390 Brugge, J. F., 199 Cameron, N. M., 130
Berridge, K. C., 453 Booth, F. W., 234 Brugger, P., 426 Caminiti, R., 425
Berridge, Kent, 453 Borbly, A. A., 289 Brning, J. C., 313 Campbell, K., 275
Berry, M. J., 182 Born, J., 288, 290 Brunswick, N., 439 Campbell, S., 268
Berson, D. M., 273 Borodinsky, L. N., 59 Bruss, J., 115, 200, 329 Campbell, S. B., 460
Berti, A., 247 Bortz, R., 286 Bruyer, R., 426 Campbell, S. S., 287
Berton, O., 73 Borziex, M.-G., 141 Bryant, R. A., 379 Camperio-Ciani, A., 345346
Bertram, L., 400 Bosetti, F., 467 Bryden, M. P., 426 Campfield, L. A., 312, 319
Bessler, A., 445 Boshart, J., 433 Bryk, K., 342 Canepari, M., 235
Betarbet, R., 256 Boston, J. D., 461 Bubash, S., 405 Canli, T., 368
Bever, T. G., 429 Bouchard, T. J., Jr., 14 Buchanan, T. W., 369 Cannon, M., 473
Beverley, J. L., 297 Boucher, T. J., 212 Buchs, P.-A., 410 Cannon, W. B., 297
Bhatia, K. P., 479 Boucsein, K., 369 Buck, L. B., 218, 223 Cantallops, I., 410
Bialek, W., 182 Boudin, H., 61 Buck, Linda, 222 Cantalupo, C., 425
Bialystok, E., 434 Bouffard, M., 201 Buck, R., 358, 423 Cao, Y. Q., 209
Biben, M., 361 Boutrel, B., 282 Bucy, P. C., 100 Capiluppi, C., 345346
Bica, L. A., 151 Bowles, C. A., 379 Budinger, T. F., 114 Caramazza, A., 435
Biederman, I., 176 Bowles, S., 20 Budney, A. J., 74 Carden, D., 179
Biederman, J., 73, 130 Bowmaker, J. K., 158, 163 Buell, S. J., 125 Cardno, A. G., 472
Bielawski, D., 347 Boycott, B. B., 167 Buen, T.-V., 397 Carello, C., 439
Biernaskie, J., 146 Boyeson, M. G., 139 Buka, S. L., 474 Carey, D. P., 172, 244245
Bierut, L. J., 460 Boynton, G. M., 226 Bulsara, K. R., 141 Carlezon, W. A., Jr., 73
Bigler, E. D., 114 Bozarth, M. A., 71 Bult-Ito, A., 271 Carlisle, H. J., 298
Bilkey, D. K., 396 Bracha, H. S., 426 Bunch, S. T., 258 Carlson, J. R., 195
Billington, C. J., 314 Bradwejn, J., 372 Bundgaard, M., 36 Carlsson, A., 59
Binart, N., 335 Braithwaite, V. A., 329 Burgess, C. E., 473 Carlyon, R. P., 444
Binder, G. K., 36 Brakke, K. E., 430 Burgess, N., 394 Carmichael-Olson, H., 130
Bini, L., 464 Brammer, M. J., 471 Burgess, P. W., 399 Caron, M. G., 62, 72, 478
Bioulac, B., 267 Branch, B. J., 328 Burk, C., 365 Carpenter, G. A., 266
Birbaumer, N., 107 Branco, T., 45 Burk, J. A., 395 Carpino, A., Jr., 272
Bischof, M., 277 Brand, T., 348 Burke, D., 347 Carr, T. S., 363
Bischofberger, J., 62, 125 Brandeis, D., 445 Burn, D. J., 256 Carreiras, M., 421
Bizzi, E., 281 Brandon, J. G., 131 Burn, P., 312, 319 Carrier, B., 212
Bjrklund, A., 257 Brandt, J., 258 Burnett, A. L., 332 Carrigan, T., 141
Blackless, M., 342 Brandt, T., 205 Burns, M. M., 161 Carro, E., 132
Blackwell, A., 436, 437 Brasted, P. J., 258 Burr, D. C., 181 Carroll, E., 171
Blagrove, M., 291 Braun, A. R., 245, 281 Burrell, B., 105 Carruth, L. L., 327
Blair, H. T., 367, 394 Braun, C., 107 Burt, A., 325 Carter, A. G., 75
Blairy, S., 356 Braus, H., 89 Burt, A. D., 333334 Carter, C. S., 332
Blake, H. T., 225 Braverman, E. R., 67 Burton, H., 132 Caspi, A., 365, 460
Blake, R., 180, 186, 225 Bray, G. A., 319 Burton, L. A., 424 Caspy, T., 266
Blake, S.-H., 8 Breedlove, S. M., 129, 326, Burton, R. F., 297, 319 Cassia, V. M., 184, 185
Blakely, E., 373 327, 348 Burwell, R. D., 270 Cassone, V. M., 269
Blakemore, C., 181, 259 Breer, H., 223 Bush, G., 357 Castellanos, F. X., 446
Blakemore, S., 291 Breiter, H. C., 452 Bushnell, M. C., 212 Castelucci, V., 406
Blakemore, S.-J., 207 Bremner, J. D., 377 Buss, D. M., 19, 339, 340 Castelucci, V. F., 407
Blakeslee, Sandra, 143, 144 Brennan, P. A., 130, 362 Butelman, H. R., 454 Castiello, U., 242
Blanchard, R., 347 Brent, H. P., 189, 190 Butler, P. C., 331 Castillo, V., 75
Blanke, O., 205 Breslin, P. A., 224 Butler, S. R., 179 Castner, S. A., 389
Note: Italicized page numbers refer to Activation-synthesis hypothesis view that of the hypothalamus that regulate
figures, illustrations, and tables. during dreams, various parts of the feeding, 314
cortex are activated by the input Alarm stage of stress, 376
Ab42. See Amyloid beta protein arising from the pons plus whatever Alcohol, 454456, 456
Abducens nerve, 88 stimuli are present in the room, and and anxiety, 372, 373, 373
Ablation removal of a structure, 109 the cortex synthesizes a story to make and cerebellum, 248
Absolute refractory period time immedi- sense of all the activity, 290291 dependence on, 67, 303, 398, 454456,
ately after an action potential, when Active transport protein-mediated 455
the sodium gates close and the mem- process that expends energy to pump and eating regulation, 318
brane cannot produce an action chemicals from the blood into the fetal exposure to, 130, 130
potential in response to stimulation brain, 37 and infant reflexes, 237
of any intensity, 44 Acuity, 155 and schizophrenia, 476
Accessory nerve, 88 Acute conditions conditions having a and sleep, 283
Acetaldehyde toxic substance produced sudden onset and a strong possibility Alcoholism (alcohol dependence) the
by the metabolism of alcohol, 456 of ending quickly, 470 inability to quit drinking or to limit
Acetic acid chemical that the body uses as Adaptation decreased response to a stim- intake of alcohol in spite of strong
a source of energy, 456 ulus as a result of recent exposure to intentions to do so, 67, 303, 398,
Acetylcholine chemical similar to an it, 217, 221, 224 454456, 455
amino acid, except that the NH2 Addictive behavior. See Substance abuse Aldosterone adrenal hormone that causes
group has been replaced by an Adenosine breakdown product of AMP the kidneys to conserve sodium
N(CH3)3 group; a neurotransmitter: that forms during metabolic activity; when excreting urine, 64, 305
and Alzheimers disease, 401, 411 a neuromodulator that inhibits the Allied reflexes, 237
and antidepressant drugs, 462 basal forebrain cells that promote Allopregnanolone, 334
and arousal, 278 arousal and wakefulness, 60, 280, All-or-none law principle stating that
inactivation/reuptake of, 6667 454 the size, amplitude, and velocity of
and muscles, 234 Adenosine triphosphate (ATP) a com- the action potential are independent
and parasympathetic nervous system, 87 pound that stores energy; also used of the intensity of the stimulus that
receptors for, 62, 63 as a neuromodulator, 488489, 489 initiated it, 44
release of, 62 ADH (antidiuretic hormone). See Allostasis adaptive way in which the
and sleep, 281282 Vasopressin body changes its set points in re-
synthesis of, 60, 61, 61 ADHD. See Attention-deficit/hyperactivity sponse to changes in its life or
Acetylcholinesterase enzyme that breaks disorder changes in the environment, 297
acetylcholine into acetate and Adoption studies, 14. See also specific Alpha-fetoprotein protein that binds with
choline, 6667 research subjects estrogen in the bloodstream of many
Across-fiber pattern principle notion that Adrenal cortex, 64, 376377 immature mammals, 329
each receptor responds to a wide Adrenal hormones, 64, 341, 376377, Alpha wave rhythm of 8 to 12 brain
range of stimuli and contributes to 379380 waves per second, generally associ-
the perception of every stimulus in Adrenaline, 58 ated with relaxation, 275, 276
its system, 215, 218 Adrenal medulla, 64 Alprazolam (Xanax), 372
ACTH (adrenocorticotropic hormone) Adrenocorticotrophic hormone. See ACTH Altruistic behavior behavior that benefits
hormone that stimulates the human Aerobic process one that uses oxygen someone other than the individual
adrenal cortex to release cortisol and during the activity, 234 engaging in the behavior, 1920
the rat adrenal gland to release corti- Afferent axon neuron that brings informa- Alzheimers disease condition character-
costerone, 64, 66, 341, 376 tion into a structure, 3334, 34 ized by memory loss, confusion,
Action potential rapid depolarization and Affinity tendency of a drug to bind to a depression, restlessness, hallucina-
slight reversal of the usual polariza- particular type of receptor, 71 tions, delusions, sleeplessness, and
tion caused by stimulation beyond African gray parrot, 431, 431, 432 loss of appetite, 399401, 400, 401,
the threshold, 4247, 42, 43, 45 2-AG (sn-2 arachidonylglycerol) chemical 411
and audition, 198199 that is produced in large quantities and basal forebrain, 278, 401
and color vision, 157 by the brain and that attaches to and nucleus basalis, 93
and movement, 250 cannabinoid receptors, 67, 75, 212 and somatosensory cortex, 209
and muscles, 234, 235 Age: Amacrine cells, 154, 154, 166
and myelin sheaths, 4647, 46 and Alzheimers disease, 400 Amblyopia. See Lazy eye
and neurotransmitter release, 61 and circadian rhythms, 267, 268 Amino acids acids containing an amine
propagation of, 4546, 45 and hippocampus, 393 group, 12, 37, 60, 65, 222, 365
and somatosensation, 206 and mate selection, 339340 Amisulpride, 479
and taste, 217218 and sleep, 289, 289 Amnesia memory loss, 398402
and vestibular sensation, 205 Aggressive behavior. See Attack behaviors and Alzheimers disease, 399401, 400,
Activating effect temporary effect of a Agonist drug that mimics or increases the 401
hormone on behavior or anatomy, effects of a neurotransmitter, 71 and hippocampus, 389393, 391
occurring only while the hormone is Agouti-related peptide (AgRP) inhibitory and Korsakoffs syndrome, 37,
present, 327, 331335 neurotransmitter found in the areas 398399
Nancy C.
Andreasen
Being a scientist and a William C.
clinician is a double priv- Dement
ilege. We actually get The average person
paid to spend our time would not, at first blush,
asking both scientific and pick watching people
clinical questions that sleep as the most appar-
everyone would like to ask and have answered, and people ent theme for a spine-tingling scientific adventure thriller.
grant us the trust of sharing their most intimate thoughts and However, there is a subtle sense of awe and mystery
experiences with us. surrounding the short death we call sleep.
Masao Ito
Brains seem to be built
on several principles such
that numerous neurons
interact with each other
through excitation and
Roger W. Sperry
inhibition, that synaptic (19131994)
plasticity provides mem- When subjective values
ory elements, that multi- have objective conse-
layered neuronal networks quences . . . they become
bear a high computational power, and that combination of part of the content of science. . . . Science would become the
neuronal networks, sensors and effectors constitutes a neu- final determinant of what is right and true, the best source
ral system representing a brain function. Thus, Hebbian and authority available to the human brain for finding ulti-
tradition has provided a very successful paradigm in modern mate axioms and guideline beliefs to live by, and for reach-
neuroscience, but we may have to go beyond it in order to ing an intimate understanding and rapport with the forces
understand the entire functions of brains. that control the universe and created man.
David Hubel
Brain science is difficult
and tricky, for some rea-
son; consequently one Patricia S.
should not believe a Goldman-Rakic
result (ones own or any- The question of how the
one elses) until it is brain organizes its sub-
proven backwards and forwards or fits into a framework so systems to produce integrated behavior is perhaps the
highly evolved and systematic that it couldnt be wrong. most challenging that can be posed.
Eric R. Kandel
Charles Scott
The questions posed by
Sherrington
higher cognitive processes
(18571952) such as learning and
A rainbow every morning memory are formidable,
who would pause to look and we have only begun
at? The wonderful which to explore them. Al-
comes often or is plenti- though elementary as-
fully about us is soon pects of simple forms of
taken for granted. That is practical enough. It allows us to get learning have been accessible to molecular analysis in
on with life. But it may stultify if it cannot on occasion be invertebrates, we are only now beginning to know a bit
thrown off. To recapture now and then childhoods wonder, about the genes and proteins involved in more complex,
is to secure a driving force for occasional grown-up thoughts. hippocampus based, learning processes of mammals.
Frank A. Beach
Carla J. Shatz (19111988)
The functioning of the Grant money comes
brain depends upon the from taxes; taxes come
precision and patterns of from a lot of folks who
its neural circuits. How dont have much money.
is this amazing computa- Spend that money wisely.
tional machine assembled To what degree should
and wired during devel- my choice of research
opment? The biological work be governed by
answer is so much more human needs, by social
wonderful than antici- imperatives, and how
pated! The adult precision is sculpted from an early imprecise am I going to justify spending all of my energies on any re-
pattern by a process in which connections are verified by the search that does not bear directly on pressing human prob-
functioning of the neurons themselves. Thus, the developing lems? . . . The solution, or rationalization, that I have finally
brain is not simply a miniature version of the adult. come up with is that it is a perfectly worthwhile way of
Moreover, the brain works to wire itself, rather than assem- spending ones life to do your level best to increase human
bling itself first and then flipping a switch, as might happen knowledge, and it is not necessary nor is it always even desir-
in the assembly of a computer. This kind of surprise in scien- able to be constrained by possible applicability of what you
tific discovery opens up new vistas of understanding and find to immediate problems. This may sound very peculiar to
possibility and makes the process of doing science infinitely some young people, but it is a value judgement which I my-
exciting and fascinating. self have made and which I can live with.
Walter B. Cannon
(18711945)
As a matter of routine I
have long trusted uncon-
scious processes to serve
me. . . . [One] example I
may cite was the interpre-
tation of the significance
of bodily changes which
Jerre Levy occur in great emotional
Despite the quite amaz- excitement, such as fear
ing progress of the last and rage. These changes
half century in neuro- the more rapid pulse, the
scientific understanding, deeper breathing, the increase of sugar in the blood, the secre-
we are still, in my view, tion from the adrenal glandswere very diverse and seemed
as distant now as ever in unrelated. Then, one wakeful night, after a considerable col-
knowing what questions to ask about how and why brains lection of these changes had been disclosed, the idea flashed
make minds. It is simply evading the issue to say, as some through my mind that they could be nicely integrated if con-
philosophers do, that our mental experiences are just the ceived as bodily preparations for supreme effort in flight or in
inside view of the stuff we measure on the outside. Why is fighting.
the inside view so utterly different from our external mea-
surements? Even if we specified all the critical spatiotempo-
ral neural dynamics that were necessary and sufficient for
a given mental experience, this would not tell us why those Larry R. Squire
dynamics give rise to any experience at all. . . . Nature will Memory is personal and
answer if we ask the right questions. evocative, intertwined
with emotion, and it pro-
vides us with a sense of
who we are. During the
past two decades there
has been a revolution in
our understanding of
what memory is and
what happens in the
brain when we learn and
remember. At the beginning of the 21st century, one has the
sense that memory may be the first mental faculty that will
be understandable in terms of molecules, cells, brain sys-
tems, and behavior. Yet, even with all the progress, there can
Curt P. Richter
be no doubt that the study of the brain is still a young sci-
(18941988) ence, rich with opportunity for the student and beginning
I enjoy research more scientist. This is a good time to hear about the promise and
than eating. excitement of neuroscience. The best is yet to come.
Torsten Wiesel
Neural connections can
be modulated by envi-
ronmental influences
during a critical period
Karl S. Lashley
of postnatal develop- (18901958)
ment. . . . Such sensitiv- Psychology is today
ity of the nervous system a more fundamental
to the effects of experience may represent the fundamental science than neurophysi-
mechanism by which the organism adapts to its environ- ology. By this I mean that
ment during the period of growth and development. the latter offers few
principles from which we may predict or define the normal
organization of behavior, whereas the study of psychological
processes furnishes a mass of factual material to which the
laws of nervous action in behavior must conform.
Ralph
Adolphs
Will a better under-
standing of the social
brain lead to a better Rita
understanding of social Levi-Montalcini
behavior? And can such Many years later, I often
knowledge ultimately be asked myself how we
used to help our species could have dedicated
negotiate and survive in ourselves with such
the vastly complex social world it has helped create? To ap- enthusiasm to solving
proach such questions, social neuroscientists will need to this small neuroembryo-
establish dialogues with other disciplines in the social and logical problem while
behavioral sciences, and to be highly sensitive to the public German armies were advancing throughout Europe, spread-
consequences of the data they generate. ing destruction and death wherever they went and threaten-
ing the very survival of Western civilization. The answer lies
in the desperate and partially unconscious desire of human
beings to ignore what is happening in situations where full
awareness might lead one to self-destruction.