Biological

Psychology
An Introduction to Behavioral,
Cognitive, and Clinical Neuroscience
SEVENTH EDITION

S. MARC BREEDLOVE NEIL V. WATSON

Michigan State University Simon Fraser University

Sinauer Associates, Inc. Publishers • Sunderland, Massachusetts

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd III 3/1/13 1:51 PM

 Brief Contents

Chapter 1 Biological Psychology: Scope and Outlook 1

PART I Biological Foundations of Behavior 21
Chapter 2 Functional Neuroanatomy: The Nervous System and Behavior 23
Chapter 3 Neurophysiology: The Generation, Transmission, and Integration
of Neural Signals 59
Chapter 4 The Chemistry of Behavior: Neurotransmitters and
Neuropharmacology 91
Chapter 5 Hormones and the Brain 125
PART II Evolution and Development of the Nervous System 155
Chapter 6 Evolution of the Brain and Behavior 157
Chapter 7 Life-Span Development of the Brain and Behavior 185
PART III Biological Foundations of Behavior 221
Chapter 8 General Principles of Sensory Processing, Touch, and Pain 223
Chapter 9 Hearing, Vestibular Perception, Taste, and Smell 255
Chapter 10 Vision: From Eye to Brain 291
Chapter 11 Motor Control and Plasticity 327
PART IV Regulation and Behavior 359
Chapter 12 Sex: Evolutionary, Hormonal, and Neural Bases 361
Chapter 13 Homeostasis: Active Regulation of the Internal Environment 393
Chapter 14 Biological Rhythms, Sleep, and Dreaming 423
PART V Emotions and Mental Disorders 455
Chapter 15 Emotions, Aggression, and Stress 457
Chapter 16 Psychopathology: Biological Basis of Behavioral Disorders 491
PART VI Cognitive Neuroscience 523
Chapter 17 Learning and Memory 525
Chapter 18 Attention and Higher Cognition 561
Chapter 19 Language and Hemispheric Asymmetry 597
© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured
or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd VI 3/4/13 3:52 PM

 Contents

Biological Psychology:
1 Scope and Outlook 1
Human or Machine? 1
The Brain Is Full of Surprises 2
What Is Biological Psychology? 2
Five Viewpoints Explore the Biology of Behavior 3
BOX 1.1 We Are All Alike, and We Are All Different 5
Three Approaches Relate Brain and Behavior 6
Neuroplasticity: Behavior Can Change the Brain 7
Biological Psychologists Use Several Levels of Analysis 10
A Preview of the Book: Relations between Brain and Behavior 11
Neuroscience Contributes to Our Understanding of Human
Disorders 12
Animal Research Makes Vital Contributions 13
The History of Research on the Brain and Behavior Begins
in Antiquity 14
BOX 1.2 Bigger Better? The Case of the Brain and Intelligence 17
The Cutting Edge Neuroscience Is Advancing at a
Tremendous Rate 19
Visual Summary 20

PART

I Biological Foundations of Behavior 21

2 Functional Neuroanatomy:
The Nervous System and Behavior 23
A Stimulating Experience 23
The Nervous System Is Composed of Cells 24

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd VII 3/1/13 1:51 PM

 VIII CONTENTS

BOX 2.1 Neuroanatomical Methods Provide Ways The Chemistry of Behavior:

4
to Make Sense of the Brain 26
Neurotransmitters and
The Nervous System Consists of Central and Neuropharmacology 91
Peripheral Divisions 34
BOX 2.2 Three Customary Orientations for Viewing The Birth of a Pharmaceutical Problem
the Brain and Body 40 Child 91
The Brain Is Described by Both Structure and Many Chemical Neurotransmitters Have Been
Function 43 Identified 92
Specialized Support Systems Protect and Nourish Neurotransmitter Systems Form a Complex Array in
the Brain 47 the Brain 94
Brain Imaging Techniques Reveal the Structure and The Effects of a Drug Depend on Its Site of Action
Function of the Living Human Brain 49 and Dose 98
BOX 2.3 Isolating Specific Brain Activity 51 Drugs Affect Each Stage of Neural Conduction and
Synaptic Transmission 104
The Cutting Edge Two Heads Are Better
Than One 54 Drugs That Affect the Brain Can Be Divided into
Functional Classes 107
Visual Summary 56
Drug Abuse Is Pervasive 116
Neurophysiology: The BOX 4.1 The Terminology of Substance-Related
Generation, Transmission, Disorders 117
3 and Integration of Neural
Signals 59
The Cutting Edge The Needle and the
Damage Undone 121
The Laughing Brain 59 Visual Summary 123

Electrical Signals Are the Vocabulary of the Nervous

System 60 Hormones and the
BOX 3.1 Changing the Channel 67 5 Brain 125
BOX 3.2 Electrical Synapses Work with No Life-Threatening Lethargy 125
Time Delay 71
Hormones Have Many Actions in the Body 125
Synapses Cause Graded, Local Changes in the
Postsynaptic Membrane Potential 71 Hormones Have a Variety of Cellular
Actions 131
Synaptic Transmission Requires a Sequence of
Events 76 BOX 5.1 Techniques of Modern Behavioral
Endocrinology 134
Neurons and Synapses Combine to Make
Circuits 82 Each Endocrine Gland Secretes Specific
Hormones 137
Gross Electrical Activity of the Human Brain 84
BOX 5.2 Stress and Growth: Psychosocial
The Cutting Edge Optogenetics: Dwarfism 143
Using Light to Probe Brain–Behavior
Hormones Affect Behavior in Many Different
Relationships 87
Ways 149
Visual Summary 88
Hormonal and Neural Systems Interact to Produce
Integrated Responses 150
The Cutting Edge Hormones Made By the
Brain, for the Brain 152
Visual Summary 154
© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured
or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd VIII 3/1/13 1:51 PM

 CONTENTS IX

PART

II Evolution and Development of the Nervous System 155

Evolution of the Brain Life-Span Development

6 and Behavior 157
7 of the Brain and
Behavior 185
We Are Not So Different, Are We? 157
How Did the Enormous Variety of Species Arise Overcoming Blindness 185
on Earth? 158 Growth and Development of the Brain Are Orderly
Why Should We Study Other Species? 162 Processes 185

BOX 6.1 Why Should We Study Particular
Species? 163
BOX 6.2 To Each Its Own Sensory World
All Vertebrate Brains Share the Same Basic
Structures 167
The Evolution of Vertebrate Brains Reflects
Changes in Behavior 169
Many Factors Led to the Rapid Evolution of a
Large Cortex in Primates 174
BOX 6.3 Evolutionary Psychology
Evolution Continues Today 179
The Cutting Edge Are Humans
Still Evolving? 181
Visual Summary 183
Development of the Nervous System Can Be
Divided into Six Distinct Stages 187
165 BOX 7.1 Degeneration and Regeneration of
Nervous Tissue 191
BOX 7.2 The Frog Retinotectal System
Demonstrates Intrinsic and Extrinsic Factors
in Neural Development 200
Developmental Disorders of the Brain Impair
Behavior 202
Genes Interact with Experience to Guide Brain
177 Development 204
BOX 7.3 Transgenic and Knockout Mice 205
Experience Is an Important Influence on Brain
Development 209
The Brain Continues to Change as We Grow
Older 212
Two Timescales Are Needed to Describe Brain
Development 216
The Cutting Edge Genetically Reversing
an Inherited Brain Disorder 217
Visual Summary 219

PART

III Biological Foundations of Behavior 221

General Principles of What Type of Stimulus Was That? 225

8 Sensory Processing,
Touch, and Pain 223
Sensory Processing Begins in Receptor Cells
Sensory Information Processing Is Selective and
226

Analytical 228
What’s Hot? What’s Not? 223
BOX 8.1 Synesthesia 235
Sensory Processing 223
Touch: Many Sensations Blended
Sensory Receptor Organs Detect Energy or Together 235
Substances 224

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd IX 3/1/13 1:51 PM

 X CONTENTS

Skin Is a Complex Organ That Contains a Variety of The Human Tongue Discriminates Five Basic
Sensory Receptors 235 Tastes 276
The Dorsal Column System Carries Somatosensory Chemicals in the Air Elicit Odor Sensations 281
Information from the Skin to the Brain 238
The Cutting Edge More Than a Matter of
Pain: An Unpleasant but Adaptive Taste 287
Experience 241
Visual Summary 288
Human Pain Can Be Measured 242
Pain Can Be Difficult to Control 247
Vision: From Eye to
The Cutting Edge Sticks and
Stones… 251
10 Brain 291
When Seeing Isn’t Seeing 291
Visual Summary 253
The Visual System Extends from the Eye
to the Brain 291
Hearing, Vestibular BOX 10.1 The Basics of Light 294
9 Perception, Taste, and
Smell 255 Neural Signals Travel from the Retina to Several
Brain Regions 299
No Ear for Music 255 BOX 10.2 Eyes with Lenses Have Evolved in
Several Phyla 302
Hearing 255
Neurons at Different Levels of the Visual System
BOX 9.1 The Basics of Sound 256 Have Very Different Receptive Fields 303
Each Part of the Ear Performs a Specific Area V1 Is Organized in Columns 312
Function in Hearing 257
Color Vision Depends on Special Channels from the
Auditory System Pathways Run from the Retinal Cones through Cortical Area V4 314
Brainstem to the Cortex 262
BOX 10.3 Most Mammalian Species Have Some
Pitch Information Is Encoded in Two Color Vision 316
Complementary Ways 264
Perception of Visual Motion Is Analyzed by a
Brainstem Auditory Systems Are Specialized Special System That Includes Cortical Area
for Localizing Sounds 266 V5 319
The Auditory Cortex Performs Complex Tasks The Many Cortical Visual Areas Are Organized into
in the Perception of Sound 268 Two Major Streams 320
Hearing Loss Is a Major Disorder of the Nervous Visual Neuroscience Can Be Applied to Alleviate
System 270 Some Visual Deficiencies 322
Vestibular Perception 273 The Cutting Edge Seeing the Light 323
The Receptor Mechanisms for the Vestibular Visual Summary 325
System Are in the Inner Ear 273
Nerve Fibers from the Vestibular Portion of the
Motor Control and
11
Vestibulocochlear Nerve (VIII) Synapse in the
Brainstem 275 Plasticity 327
Some Forms of Vestibular Excitation Produce What You See Is What You Get 327
Motion Sickness 276
The Behavioral View 327
The Chemical Senses: Taste and
Smell 276 The Control Systems View 329

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd X 3/1/13 1:51 PM

 CONTENTS XI

The Neuroscience View 330 Disorders of Muscle, Spinal Cord, or Brain Can
Disrupt Movement 347
Movements Are Controlled at Several Nervous
System Levels 337 The Cutting Edge Cerebellar Glia Play a
BOX 11.1 Cortical Neurons Can Guide a Robotic Role in Fine Motor Coordination 354
Arm 341 Visual Summary 356
Extrapyramidal Systems Also Modulate Motor
Commands 345

PART

IV Regulation and Behavior 359

Sex: Evolutionary, Homeostasis: Active

12 Hormonal, and Neural
Bases 361 13 Regulation of the Internal
Environment 393
Genitals and Gender: What Makes Us Male A Love-Hate Relationship with Food 393
and Female? 361
Homeostasis Maintains a Consistent
Sexual Behavior 361 Internal Environment: The Example of
Thermoregulation 394
Reproductive Behavior Can Be Divided into
Four Stages 362 BOX 13.1 Integrated Physiological and Behavioral
Thermoregulation Helps Young Animals to
The Neural Circuitry of the Brain Regulates Survive 399
Reproductive Behavior 365
Water Moves between Two Major Body
Pheromones Guide Reproductive Behavior in Compartments 400
Many Species 367
Two Internal Cues Trigger Thirst 401
The Hallmark of Human Sexual Behavior Is
Diversity 369 Food and Energy Regulation 405
For Many Vertebrates, Parental Care Determines Nutrient Regulation Helps Prepare for
Offspring Survival 372 Future Needs 405

Sexual Differentiation 373 Insulin Is Crucial for the Regulation of Body

Metabolism 408
Sex Determination and Sexual Differentiation Occur
Early in Development 373 The Hypothalamus Coordinates Multiple Systems
That Control Hunger 409
How Should We Define Gender—by Genes,
Gonads, Genitals, or the Brain? 378 Obesity Is Difficult to Treat 415

Gonadal Hormones Direct Sexual Differentiation of BOX 13.2 Body Fat Stores Are Tightly Regulated,
Even after Surgical Removal of Fat 416
the Brain and Behavior 378
Eating Disorders Are Life-Threatening 418
BOX 12.1 The Paradoxical Sexual Differentiation of
the Spotted Hyena 381 The Cutting Edge A Rumbling in the
Do Fetal Hormones Masculinize Human Behaviors Belly 419
in Adulthood? 386 Visual Summary 421
The Cutting Edge Sex on the Brain 390
Visual Summary 391

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd XI 3/1/13 1:51 PM

 XII CONTENTS

Biological Rhythms, Sleep, Our Sleep Patterns Change across the Life
14 and Dreaming 423 Span 435
Manipulating Sleep Reveals an Underlying
When Sleep Gets Out of Control 423 Structure 437
Biological Rhythms 423 BOX 14.1 Sleep Deprivation Can Be Fatal 438
Many Animals Show Daily Rhythms in Activity 423 What Are the Biological Functions of Sleep? 439
The Hypothalamus Houses a Circadian Clock 425 At Least Four Interacting Neural Systems Underlie
Sleep 443
Some Biological Rhythms Are Longer or Shorter
than a Day 429 Sleep Disorders Can Be Serious, Even
Life-Threatening 448
Sleep and Waking 430
The Cutting Edge Can Individual Neurons
Human Sleep Exhibits Different Stages 430
Be “Sleepy”? 451
Different Species Provide Clues about the
Visual Summary 453
Evolution of Sleep 434

PART

V Emotions and Mental Disorders 455

Emotions, Aggression, Psychopathology: Biological

15 and Stress 457
16 Basis of Behavioral
Disorders 491
Trouble in Paradise 457
What Are Emotions? 458 “My Lobotomy” 491
Broad Theories of Emotion Emphasize Bodily The Toll of Psychiatric Disorders Is Huge 492
Responses 458 Schizophrenia Is the Major Neurobiological
BOX 15.1 Lie Detector? 461 Challenge in Psychiatry 492
How Many Emotions Do We Experience? 462 BOX 16.1 Long-Term Effects of Antipsychotic
Drugs 502
Emotions from the Evolutionary Viewpoint 465
Mood Disorders Are a Major Psychiatric
Do Distinct Brain Circuits Mediate Emotions? 468 Category 506
Neural Circuitry, Hormones, and Synaptic BOX 16.2 The Season to Be Depressed 511
Transmitters Mediate Violence and
There Are Several Types of Anxiety Disorders 513
Aggression 476
BOX 16.3 Tics, Twitches, and Snorts: The Unusual
Stress Activates Many Bodily Responses 479
Character of Tourette’s Syndrome 516
Stress and Emotions Affect the Immune Neurosurgery Has Been Used to Treat Psychiatric
System 482 Disorders 517
The Cutting Edge Synaptic Changes Abnormal Prion Proteins Destroy the Brain 518
during Fear Conditioning 487
The Cutting Edge Are Abnormal Eye
Visual Summary 489 Movements an Endophenotype for People
at Risk for Schizophrenia? 519
Visual Summary 521

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd XII 3/1/13 1:51 PM

 CONTENTS XIII

PART

VI Cognitive Neuroscience 523

Learning and Attention and Higher

17 Memory 525 18 Cognition 561
Trapped in the Eternal Now 525 One Thing at a Time 561
Functional Perspectives on Memory 525 Attention 561
There Are Several Kinds of Memory and Attention Selects Stimuli for Processing 561
Learning 526
Attention May Be Endogenous or Exogenous 565
Memory Has Temporal Stages: Short, Intermediate,
BOX 18.1 Reaction-Time Responses, from Input to
and Long 530 Output 566
Successive Processes Capture, Store, and Retrieve Electrophysiological Techniques Trace Rapid
Information in the Brain 532 Changes of Brain Activity 570
BOX 17.1 Emotions and Memory 534 Many Brain Regions Are Involved in Processes
Different Brain Regions Process Different Aspects of Attention 574
of Memory 536 Two Cortical Networks Collaborate to Govern
Neural Mechanisms of Memory Attention 578
Storage 542 Disorders Provide Clues about the Organization
Memory Storage Requires Neuronal of Attention 581
Remodeling 542 Consciousness 583
Invertebrate Nervous Systems Show Consciousness Is the Most Mysterious Property
Plasticity 545 of the Nervous System 583
Synaptic Plasticity Can Be Measured in Simple The Frontal Lobes Govern Our Most Complex
Hippocampal Circuits 547 Behavior 588
Some Simple Learning in Mammals Relies on BOX 18.2 Phineas Gage 590
Circuits in the Cerebellum 551
The Cutting Edge Putting the You in
In the Adult Brain, Newly Born Neurons May Aid YouTube 593
Learning 554
Visual Summary 595
Learning and Memory Change as We Age 555
The Cutting Edge Artificial Activation of an
Engram 557
Visual Summary 559

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd XIII 3/1/13 1:51 PM

 XIV CONTENTS

Language and Hemispheric

19 Asymmetry 597
Putting a Name to a Face 597
Language Has Both Learned and Innate Components 597
Language Disorders Result from Region-Specific Brain Damage 605
BOX 19.1 Williams Syndrome Offers Clues about Language 600
BOX 19.2 The Wada Test 605
Competing Models Describe Left-Hemisphere Language
Specializations 608
Reading Skills Are Difficult to Acquire and Frequently Impaired 611
Brain Stimulation Provides Information about the Organization of
Language in the Brain 614
Functional Neuroimaging Tracks Activity in the Brain’s Language
Zones 616
The Left Brain Is Different from the Right Brain 618
Deficits in Spatial Perception Follow Right-Hemisphere Damage 624
Following Some Injuries, the Brain Can Recover Function 626
BOX 19.3 The Amazing Resilience of a Child’s Brain 627
BOX 19.4 Contact Sports Can Be Costly 628
The Cutting Edge Studying Connectivity in the
Living Brain 630
Visual Summary 632

Appendix A–1
Glossary G–1
Illustration Credits IC–1
References R–1
Author Index AI–1
Subject Index SI–1

© Sinauer Associates, Inc. This material cannot be copied, reproduced, manufactured

or disseminated in any form without express written permission from the publisher.

00_FM_Breedlove.indd XIV 3/1/13 1:51 PM