Behavioral Neuroscience 9th Edition S.

Mark Breedlove
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 Behavioral Neuroscience
Ninth Edition

BRAIN, n. An apparatus with which we think that we think.

MIND, n. A mysterious form of matter secreted by the brain.

Its chief activity consists in the endeavor to ascertain its own
nature, the futility of the attempt being due to the fact that it
has nothing but itself to know itself with.

Ambrose Bierce, 1911 (The Collected Works of Ambrose Bierce:

Volume VII, The Devil’s Dictionary, p. 41, 217)
 Behavioral
Neuroscience
NINTH EDITION

Courtesy of Dr. Sarah Moghadam, VA Palo

Alto Health Care System, Palo Alto, CA and
Dr. Ahmad Salehi, Dept. of Psychiatry &
Behavioral Sciences, Stanford Medical School

S. MARC BREEDLOVE
Michigan State University
NEIL V. WATSON
Simon Fraser University

SINAUER ASSOCIATES

NEW YORK OXFORD

OXFORD UNIVERSITY PRESS

About the Cover:
Hippocampal (brain) region. Courtesy of Dr.
Sarah Moghadam, VA Palo Alto Health Care
System, Palo Alto, CA and Dr. Ahmad Salehi,
Dept. of Psychiatry & Behavioral Sciences,
Stanford Medical School.
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Library of Congress Cataloging-in-Publication Data
Names: Breedlove, S. Marc, author. | Watson, Neil V. (Neil Verne), 1962−author.
Title: Behavioral neuroscience / S. Marc Breedlove, Michigan State
University, Neil V. Watson, Simon Fraser University.
Description: Ninth edition. | Sunderland, Massachusetts : Sinauer Associates,
Inc. Publishers, [2020] | Includes bibliographical references and index.
Identifiers: LCCN 2019019280 | ISBN 9781605359076 (hardback)
Subjects: LCSH: Psychobiology.
Classification: LCC QP360 .B727 2020 | DDC 612.8--dc23
LC record available at https://lccn.loc.gov/2019019280
987654321
Printed in the United States of America
For Stacey, Collin, and John For Scott and Sherry
S.M.B. N.V.W.
Brief Contents
Chapter 1 Introduction Scope and Outlook 1

PART I Biological Foundations of Behavior 23

Chapter 2 Functional Neuroanatomy
The Cells and Structures of the Nervous System 25
Chapter 3 Neurophysiology The Generation, Transmission, and Integration
of Neural Signals 63
Chapter 4 The Chemistry of Behavior
Neurotransmitters and Neuropharmacology 97
Chapter 5 Hormones and the Brain 137

PART II Evolution and Development of the Nervous System 165

Chapter 6 Evolution of the Brain and Behavior 167
Chapter 7 Life-Span Development of the Brain and Behavior 199

PART III Perception and Action 231

Chapter 8 General Principles of Sensory Processing, Touch, and Pain 233
Chapter 9 Hearing, Balance, Taste, and Smell 269
Chapter 10 Vision From Eye to Brain 309
Chapter 11 Motor Control and Plasticity 347

PART IV Regulation and Behavior 381

Chapter 12 Sex Evolutionary, Hormonal, and Neural Bases 383
Chapter 13 Homeostasis Active Regulation of the Internal Environment 417
Chapter 14 Biological Rhythms, Sleep, and Dreaming 449

PART V Emotions and Mental Disorders 485

Chapter 15 Emotions, Aggression, and Stress 487
Chapter 16 Psychopathology Biological Basis of Behavioral Disorders 521

PART VI Cognitive Neuroscience 555

Chapter 17 Learning and Memory 557
Chapter 18 Attention and Higher Cognition 595
Chapter 19 Language and Lateralization 631
Contents
1 Introduction 1
Scope and Outlook
1.3 Behavioral Neuroscientists Use
Several Levels of Analysis 10
1.4 The History of Research on the Brain
Machine or Human? 1 and Behavior Begins in Antiquity 14
1.1 The Brain Is Full of Surprises 2 BOX 1.2 Bigger Better? The Case of the Brain
BOX 1.1 We Are All Alike, and We Are and Intelligence 18
All Different 6 The Cutting Edge ■ Behavioral Neuroscience
Is Advancing at a Tremendous Rate 20
1.2 Three Approaches Relate Brain
and Behavior 7 Visual Summary 22

PART I Biological Foundations of Behavior 23

2 Functional Neuroanatomy 25
The Cells and Structures of the
Nervous System
3 Neurophysiology 63
The Generation, Transmission, and
Integration of Neural Signals
A Stimulating Experience 25 The Laughing Brain 63
2.1 S
 pecialized Cells Make Up the 3.1 E
 lectrical Signals Are the Vocabulary
Nervous System 26 of the Nervous System 64
BOX 2.1 Visualizing the Cells of the Brain 32 BOX 3.1 Voltage Clamping and Patch
Clamping 72
2.2 T
 he Nervous System Consists of
Central and Peripheral Divisions 36 BOX 3.2 Changing the Channel 76
BOX 2.2 Three Customary Orientations for 3.2 S
 ynapses Transmit Information from
Viewing the Brain and Body 43 One Neuron to Another 77
2.3 T
 he Brain Shows Regional 3.3 A
 ction Potentials Cause the Release
Specialization of Functions 46 of Transmitter Molecules into the
Synaptic Cleft 81
2.4 S
 pecialized Support Systems Protect
and Nourish the Brain 50 3.4 G
 ross Electrical Activity of the Brain
Is Readily Detected 90
2.5 B
 rain-Imaging Techniques Reveal the
Structure and Function of the Living The Cutting Edge ■ Optogenetics: Using Light
Human Brain 54 to Probe Brain-Behavior Relationships 93
BOX 2.3 Isolating Specific Brain Activity 57 Visual Summary 95
The Cutting Edge ■ Two Heads Are Better
Than One 59
Vi su a l S umm a r y 61
4 The Chemistry of Behavior 97 4.8 S
 ubstance Abuse and Addiction Are
Neurotransmitters and Worldwide Social Problems 127
Neuropharmacology The Cutting Edge ■ Uncovering the Insula 131
The Birth of a Pharmaceutical Problem Visual Summary 135
Child 97
4.1 Synaptic Transmission Involves a
Complex Electrochemical Process 98 5 
Hormones and the Brain 137

4.2 M
 any Neurotransmitters Have Been Crafting a Personality through
Identified 100 Hormones 137
4.3 N
 eurotransmitter Systems Form a 5.1 H
 ormones Have Many Actions
Complex Array in the Brain 101 in the Body 138
BOX 4.1 Pathways for Neurotransmitter 5.2 H
 ormones Have a Variety of Cellular
Synthesis 103 Actions 143
4.4 T
 he Effects of a Drug Depend on Its BOX 5.1 Techniques of Modern Behavioral
Site of Action and Dose 106 Endocrinology 146

4.5 D
 rugs Affect Each Stage of
Neural Conduction and Synaptic
Transmission 112
4.6 S
 ome Neuroactive Drugs Ease the
Symptoms of Injury or Psychiatric
Illness 116
4.7 S
 ome Neuroactive Drugs Are Used to
Alter Conscious Experiences
5.3 E
 ach Endocrine Gland Secretes
Specific Hormones 149
The Cutting Edge ■ Bones Secrete
Hormones to Regulate Appetite 158
5.4 H
 ormones Regulate Social Behaviors
and Vice Versa 158
Visual Summary 164
119

PART II Evolution and Development of

the Nervous System
165

6  volution of the Brain and

E 6.4 T
 he Evolution of Vertebrate Brains
Behavior 167 Reflects Changes in Behavior 181
6.5 M
 any Factors Led to the Rapid
We Are Not So Different, Are We? 167 Evolution of a Large Cortex in
6.1 H
 ow Did the Enormous Variety of Primates 186
Species Arise on Earth? 168 BOX 6.3 Evolutionary Psychology 190
6.2 W
 hy Should We Study Other 6.6 E
 volution Continues Today 193
Species? 174
The Cutting Edge ■ Are Humans Still
BOX 6.1 Why Should We Study Particular Evolving? 194
Species? 175
Visual Summary 197
BOX 6.2 To Each Its Own Sensory World 177
6.3 A
 ll Vertebrate Brains Share the Same
Basic Structures 178

VIII CO NT E NTS
7  ife-Span Development of the
L 7.3 L
 ifelong Synapse Rearrangement
Brain and Behavior 199 Is Guided by Experience 215

Overcoming Blindness 199 7.4 E

7.1 Neural Development Is Guided
by the Interaction of Genetic and
Environmental Factors 200
7.2 D
 evelopment of the Nervous System
Can Be Divided into Six Distinct
Stages 204
BOX 7.1 Transgenic and Knockout Mice 209
BOX 7.2 Degeneration and Regeneration of
Nervous Tissue 211
 xperience Can Alter Gene
Expression to Affect Brain
Development 222
The Cutting Edge ■ Harnessing Glia to
Reverse an Inherited Brain Disorder 224
7.5 T
 he Brain Continues to Change
as We Grow Older 226
Visual Summary 23 0

PART III Perception and Action 231

8 
General Principles of Sensory
Processing, Touch, and Pain
233
9  earing, Balance, Taste, and
H
Smell 269
No Ear for Music 269
What’s Hot? What’s Not? 233
HEARING 270
SENSORY PROCESSING 234
9.1 Pressure Waves in the Air Are
8.1 S
 ensory Receptor Organs Detect Perceived as Sound 270
Energy or Substances 234 BOX 9.1 The Basics of Sound 270
8.2 S
 ensory Information Processing Is 9.2 A
 uditory Signals Run from Cochlea
Selective and Analytical 238 to Cortex 277
BOX 8.1 Synesthesia 245
9.3 P
 itch Information Is Encoded in Two
TOUCH: MANY SENSATIONS BLENDED Complementary Ways 279
TOGETHER 246 9.4 B
 rainstem Auditory Systems
8.3 S
 kin Is a Complex Organ That Are Specialized for Localizing
Contains a Variety of Sensory Sounds 281
Receptors 246 9.5 T
 he Auditory Cortex Processes
PAIN: AN UNPLEASANT BUT ADAPTIVE Complex Sounds 283
EXPERIENCE 252 9.6 H
 earing Loss Is a Major Disorder of
8.4 H
 uman Pain Can Be Measured 252 the Nervous System 286
The Cutting Edge ■ Evolving an Indifference VESTIBULAR PERCEPTION 290
to Toxins 256
9.7 A
 n Inner Ear System Senses Gravity
8.5 P
 ain Can Be Difficult to Control 261 and Acceleration 290
Vi su a l S um ma r y 266 THE CHEMICAL SENSES: TASTE
AND SMELL 293

CONTENTS IX
 9.8 C
 hemicals in Foods Are Perceived 10.7 Visual Neuroscience Can Be
as Tastes 293 Applied to Alleviate Some Visual
The Cutting Edge ■ More Than a Matter Deficiencies 340
of Taste 298 The Cutting Edge ■ Seeing the Light 342
9.9 C
 hemicals in the Air Elicit Odor Visual Summary 344
Sensations 299
Visu a l S um m a r y 30 6
11  otor Control and
M
Plasticity 347

10 Vision 309
From Eye to Brain
What You See Is What You Get
11.1 The Behavioral View Considers
347

When Seeing Isn’t Seeing 309 Reflexes versus Plans 348

10.1 The Retina Transduces Light into 11.2 N
 euroscience Reveals Hierarchical
Neuronal Activity 310 Systems 350
10.2 Properties
 of the Retina Shape 11.3 T
 he Spinal Cord Is a Crucial Link in
Many Aspects of Our Vision 314 Controlling Body Movement 357
10.3 N
 eural Signals Travel from the 11.4 P
 athways from the Brain
Retina to Several Brain Regions 318 Control Different Aspects of
Movements 361
BOX 10.1 Eyes with Lenses Have Evolved in
Several Phyla 321 BOX 11.1 Cortical Neurons Can Guide a
Robotic Arm 365
10.4 N
 eurons at Different Levels of the
Visual System Have Very Different 11.5 E
 xtrapyramidal Systems Also
Receptive Fields 322 Modulate Motor Commands 369
10.5 C
 olor Vision Depends on Special The Cutting Edge ■ Cerebellar Glia Play a
Channels from the Retinal Cones Role in Fine Motor Coordination 372
through Cortical Area V4 332 11.6 B
 rain Disorders Can Disrupt
BOX 10.2 Most Mammalian Species Have Movement 373
Some Color Vision 334 BOX 11.2 Prion-Like Neurodegeneration May
10.6 T
 he Many Cortical Visual Areas Be at Work in Parkinson’s 375
Are Organized into Two Major Visual Summary 378
Streams 337

PART IV Regulation and Behavior 381

12 Sex 383
Evolutionary, Hormonal, and Neural
Bases
12.1 R
 eproductive Behavior Can Be
Divided into Four Stages 384

Genitals and Gender: What Makes Us 12.2 T

 he Neural Circuitry of the
Male and Female? 383 Brain Regulates Reproductive
Behavior 388
SEXUAL BEHAVIOR 384

X CO NT E NTS
The Cutting Edge ■ Sexual Experience 13.4 N
 utrient Regulation Helps Prepare
Solidifies Neural Circuits for Mating 390 for Future Needs 429
12.3 T
 he Hallmark of Human Sexual 13.5 A
 Hypothalamic Appetite Controller
Behavior Is Diversity 392 Integrates Multiple Hunger
Signals 433
12.4 F
 or Many Vertebrates, Parental Care
Determines Offspring Survival 396 The Cutting Edge ■ Friends with
Benefits 440
SEXUAL DIFFERENTIATION 397
13.6 O
 besity and Eating Disorders Are
12.5 S
 ex Determination and Sexual Difficult to Treat 441
Differentiation Occur Early in
BOX 13.2 Body Fat Stores Are Tightly
Development 397 Regulated, Even after Surgical Removal
12.6 G
 onadal Hormones Direct Sexual of Fat 442
Differentiation of the Brain and Visual Summary 447
Behavior 402

14
BOX 12.1 The Paradoxical Sexual
Differentiation of the Spotted Hyena 406
 iological Rhythms, Sleep,
B
and Dreaming 449
12.7 D
 o Fetal Hormones Masculinize
Human Behaviors in When Sleep Gets Out of Control 449
Adulthood? 410
BIOLOGICAL RHYTHMS 450
Vi su a l S umm a r y 41 5
14.1 Many Animals Show Daily Rhythms in
Activity 450

13 Homeostasis 417
Active Regulation of the Internal
Environment
14.2 T
 he Hypothalamus Houses a
Circadian Clock 451
SLEEPING AND WAKING 457
Harsh Reality TV 417
14.3 H
 uman Sleep Exhibits Different
13.1 H
 omeostasis Maintains a Consistent Stages 457
Internal Environment: The Example
of Thermoregulation 418 14.4 W
 hy and How Did Sleep
BOX 13.1 Physiological and Behavioral Evolve? 463
Thermoregulation Are Integrated 422 BOX 14.1 Sleep Deprivation Can Be Fatal 464
FLUID REGULATION 422 14.5 A
 t Least Four Interacting Neural
Systems Underlie Sleep 471
13.2 W
 ater Shuttles between Two Body
Compartments 422 The Cutting Edge ■ Can Individual Neurons
Be “Sleepy”? 476
13.3 T
 wo Internal Cues Trigger
Thirst 425 14.6 S
 leep Disorders Can Be Serious,
Even Life-Threatening 478
FOOD AND ENERGY Visual Summary 482
REGULATION 429

CONTENTS XI
 PART V Emotions and Mental Disorders 485

15  motions, Aggression,
E
and Stress 487 16 Psychopathology 521
Biological Basis of Behavioral
Disorders
The Hazards of Fearlessness 487
“The Voice” 521
15.1 B
 road Theories of Emotion
Emphasize Bodily Responses 488 16.1 S
 chizophrenia Is the Major
Neurobiological Challenge in
BOX 15.1 Lie Detector? 491 Psychiatry 523
15.2 D
 id a Core Set of Emotions Evolve BOX 16.1 Long-Term Effects of
in Humans and Other Animals? 492 Antipsychotic Drugs 532

15.3 S
 pecialized Neural Mechanisms 16.2 D
 epression Is the Most Prevalent
Mediate the Experience and Mood Disorder 537
Expression of Emotions 496 The Cutting Edge ■ Can Our Genes Tell Us
The Cutting Edge ■ Synaptic Changes Which Drugs to Use? 541
during Fear Conditioning 502
BOX 16.2 The Season to Be Depressed? 545
15.4 N
 eural Circuitry, Hormones, and
Synaptic Transmitters Mediate 16.3 E
 xtreme Mood Cycles Define
Violence and Aggression 506 Bipolar Disorder 545

15.5 S
 tress Activates Many Bodily 16.4 T
 here Are Several Types of Anxiety
Responses 510 Disorders 547
Visu a l S um m a r y 51 9 BOX 16.3 Tics, Twitches, and Snorts:
The Unusual Character of Tourette's
Syndrome 552
Visual Summary 553

PART VI Cognitive Neuroscience 555

17 
Learning and Memory 557 BOX 17.1 Emotions and Memory

NEURAL MECHANISMS OF MEMORY

575

Trapped in the Eternal Now 557 STORAGE 576

FUNCTIONAL PERSPECTIVES ON 17.4 M
 emory Storage Requires Physical
LEARNING AND MEMORY 558 Changes in the Brain 576

17.1 There Are Several Kinds of Learning 17.5 S

 ynaptic Plasticity Can Be Measured
and Memory 558 in Simple Hippocampal Circuits 582
The Cutting Edge ■ Artificial Activation
17.2 D
 ifferent Forms of Nondeclarative
of an Engram 587
Memory Involve Different Brain
Regions 565 17.6 In the Adult Brain, Newly Born
Neurons May Aid Learning 589
17.3 S
 uccessive Processes Capture, Store,
and Retrieve Information in the Visual Summary 593
Brain 569

XII CO NT E NTS
18  ttention and Higher
A 19.3 L
 eft-Hemisphere Damage Can
Cognition 595 Cause Aphasia 641

One Thing at a Time 595 19.4 C

 ompeting Models Describe
the Left-Hemisphere Language
ATTENTION 596 System 645
18.1 A
 ttention Selects Stimuli for VERBAL BEHAVIOR: SPEECH AND
Processing 596 READING 651
BOX 18.1 Reaction Time Responses, 19.5 L
 anguage Has Both Learned and
from Input to Output 600 Unlearned Components 652
18.2 Targets of Attention: Attention BOX 19.2 Williams Syndrome Offers Clues
Alters the Functioning of Many about Language 654
Brain Regions 603 BOX 19.3 Vocal Behavior in Birds and
18.3 Sources of Attention: A Network Other Species 657
of Brain Sites Creates and Directs 19.6 R
 eading Skills Are Difficult
Attention 609 to Acquire and Frequently
CONSCIOUSNESS AND EXECUTIVE Impaired 658
FUNCTION 616 RECOVERY OF FUNCTION AFTER
18.4 Consciousness Is a Mysterious BRAIN DAMAGE 662
Product of the Brain 616 19.7 S
 tabilization and Reorganization
BOX 18.2 Phineas Gage 623 Are Crucial for Recovery of
Function 662
The Cutting Edge ■ Building a Better
Mind Reader 627 BOX 19.4 The Amazing Resilience of a
Child’s Brain 663
Vi su a l S umm a r y 6 29
The Cutting Edge ■ Contact Sports Can
Be Costly 665

19  anguage and
L
Lateralization 631
Visual Summary 667

Silencing the Inner Voice 631

Appendix A–1
BRAIN ASYMMETRY AND
LATERALIZATION OF Glossary G–1
FUNCTION 632
References R–1
19.1 T
 he Left and Right Hemispheres
Are Different 632 Author Index AI–1
BOX 19.1 The Wada Test 638
Subject Index SI–1
19.2 R
 ight-Hemisphere Damage Impairs
Specific Types of Cognition 639

CONTENTS XIII

Courtesy of Dr. Sarah Moghadam, VA Palo Alto Health
Care System, Palo Alto, CA and Dr. Ahmad Salehi, Dept. of
Psychiatry & Behavioral Sciences, Stanford Medical School
XIV C HAP T E R 
Preface
Twenty-four years ago, a new kind of textbook was published for University courses
that were often called “Brain and Behavior.” As the field evolved, the book’s title
metamorphosed from Biological Psychology to Behavioral Neuroscience, but the same
drive to provide a definitive and comprehensive survey of the neurosciences lies at
the heart of all our efforts. We strive to keep the book up-to-date while keeping a
conversational tone to make this wealth of information not just accessible, but fas-
cinating. The biggest change in this new edition is the development of Learning
Objectives for each segment of the book, with the idea that telegraphing what’s to
come will focus readers’ attention and facilitate learning. As you finish each section
of text, it would be a good idea to go back and read the associated Learning Objec-
tives to see whether in fact you incorporated the material. If not, a quick review of
that text may be in order.
As always, there have been plenty of new findings to add to this edition. In fact,
the problem we face is which of the many, many new findings to leave out—those
that are not quite essential for a survey of the field. We are pretty picky about what
we add, and still it seems like a deluge of new information and ideas. Hundreds
of new papers are cited in this edition. If that sounds like a lot, let us give you a
perspective on how many new papers were omitted. On our newsfeed site (www.
biopsychology.com/news/), 1,299 new links were added in 2018 alone. Those are just
the findings that were important enough to get the attention of mass media report-
ers. As we note in Chapter 1, over 40,000 new articles indexed under “neuroscience”
appeared that year in PubMed. It would take a thick tome just to list the titles of the
papers from 2018!
While being very, very selective in sampling this flood of findings, we have made
substantial changes in every chapter. For example, in Chapter 3 we have a new
figure comparing “kiss and run” synapses with more traditional models of synaptic
transmission. Chapter 5 has new material about a hormone secreted from bone that
acts on the hypothalamus to reduce appetite. We totally reorganized Chapter 7 for a
more streamlined approach and discuss the growing doubts about whether amyloid
deposits cause Alzheimer’s. Chapter 9 needed a new figure comparing transduction
in the five taste receptors. Chapter 13 talks about yet another factor affecting appetite,
glucagon-like peptide 1. Chapter 16 now discusses the logic of pharmacogenomics
to treat depression, while Chapter 18 was thoroughly reorganized and includes more
about executive function. Honestly, we could go on like this for every chapter. Clearly
this is an exciting era in the neurosciences. As Lewis Carrol put it, “We must run as
fast as we can just to stay in place!”
We’ve also kept several very popular features from previous editions: The Cutting
Edge appears in each chapter, where we explore some of the most exciting examples
of recent research, and each chapter ends with a Visual Summary, where you can see
graphic reminders as you review the principle findings that we just presented. These
Visual Summaries really shine online, where with just a click you can review figures,
animations, and quizzes to help integrate the material. We also continue to open
each chapter with a gripping vignette, relating someone’s real-life experiences that
will be better understood as the content of the chapter unfolds, and we again replaced
several of these vignettes as more recent events bring to the surface many of the
important issues in behavioral neuroscience. Likewise we’ve retained the marginal
glossary that makes it easy to find the definitions that unlock the material, as well
as two features to let you burrow in on a particular subject: the online supplements
called A Step Further cited throughout the text, and the Recommended Reading
at the close of each chapter.
You might think that approaching the quarter-century mark we’d be jaded about
improving and revising our presentations, but we still love it, perhaps because the
dynamic and exciting pace of neuroscience research shows no sign of abating soon.
As always, we welcome all feedback, praise or criticism, cuts or additions, from our
readers. You can email us directly at behavneuro@gmail.com.

Acknowledgments
We continue to feel so lucky to work with the inestimable team at Sinauer Associates,
now a part of Oxford University Press, whose deep skills and generous guidance
transform our hundreds of files, thousands of email attachments and sometimes
scrambled emails into yet another beautiful book. Again, we feel so grateful to benefit
from the experience and exquisite taste of others. In particular, the book could not
exist without the contributions of Senior Acquisition Editor Syd Carroll, Production
Editor Alison Hornbeck, Production Manager Joan Gemme, Book Designer and
Production Specialist Annette Rapier, and Media and Supplements Editor Zan Carter
and her crew. We also fondly bid adieu to the recently retired Chris Small, Production
Manager for all our previous editions. We hope you’re enjoying yourself, Chris, but
how could you abandon us!? A cadre of commandos delved deep in the archives to
deal with copyrights and permissions, so we salute you Michele Beckta, Mark Siddall,
and Tracy Marton. We’d also like to thank our copy editor Lou Doucette, and our
longtime art studio, Dragonfly Media, who bring amazing skill and commitment to
make us look good.
We must also thank the founder of Sinauer Associates, Andy Sinauer, for his
unwavering support over the years, with a touch of sadness upon his retirement.
We are so proud to be a part of Andy’s tremendous legacy, begun all those years
ago with From Neuron to Brain, creating gorgeous books that make even the most
complex topics accessible and enjoyable.
By this point in the evolution of the book, we have benefited from the wisdom
and advice of hundreds of colleagues who have generously served as reviewers of
past editions. Although we don’t have the space to list them all, we want to ac-
knowledge that in many ways the book you are holding is the product of a whole
community of neuroscientists. In this, the Ninth Edition, the following colleagues
have provided invaluable critique and commentary:

Susan Bachus, University of Maryland, Baltimore County

Susan Barron, University of Kentucky
Christopher Beeman, Central Washington University
Jin Bo, Eastern Michigan University
David Brodbeck, Algoma University
Elizabeth Caldwell, University of New Hampshire
James Cherry, Boston University
Michael Cohen, Loyola University Chicago
Paul J. Currie, Reed College
Patrick Cushen, Murray State University
Deana Davalos, Colorado State University
Darragh P. Devine, University of Florida
Christopher W. Drapeau, Valparaiso University
Kelli A. Duncan, Vassar College
Raymond H. Dye, Jr., Loyola University Chicago
Taffeta Elliott, New Mexico Institute of Mining and Technology
Alison A. Fedio, Argosy University, Northern Virginia
Sara B. Festini, University of Tampa

P R EFACE XV
 Cynthia Michelle Finley, College of Marin
Jonathan Franz, SUNY Empire State College
Koren Ganas, University of Illinois
Sophie George, Dixie State University
Aaron Godlaski, Centre College
Brian J. Hock, Austin Peay State University
Jennifer Ingemi, Northeastern University
Mary Ellen Kelly, Haverford College
Susan Kennedy, Denison University
Michael Kerchner, Washington College
Sarita Lagalwar, Skidmore College
Stephen Lippi, George Mason University
Mario L. Mata, California State University, Los Angeles
Alexandra Roach, University of South Carolina, Aiken
Russell Romeo, Barnard College of Columbia University
Timothy Roth, Franklin and Marshall College
Emma Sarro, Dominican College
Peter A. Serrano, Hunter College, City University of New York
Fredric Shaffer, Truman State University
KatieAnn Skogsberg, Centre College
Lucy J. Troup, University of the West of Scotland
Adriana Uruena-Agnes, St. Petersburg College
Jennifer Wilhelm, College of Charleston
Jan R. Wessel, University of Iowa
Susan Zup, University of Massachusetts Boston

Finally, we thank all those tireless colleagues trying to understand the neural basis of
behavior, with techniques that would have seemed like sorcery only a few years ago,
and who share their hard-won findings with us all.

S. Marc Breedlove Neil V. Watson

XVI PREFACE
Media and
Supplements
to accompany
Behavioral Neuroscience,
Ninth Edition
Courtesy of Dr. Sarah Moghadam, VA Palo Alto Health
Care System, Palo Alto, CA and Dr. Ahmad Salehi, Dept. of
Psychiatry & Behavioral Sciences, Stanford Medical School

For the Student

Companion Website (bn9e.com)
The Behavioral Neuroscience Companion Website contains a range of study and review
resources to help students master the material presented in each chapter of the text-
book. Access to the site is included with each new copy of the textbook (see inside
front cover). The site includes the following resources:
• Chapter Outlines that outline each chapter and link to relevant Study Questions
• Brain Explorer that offers an interactive way to explore the brain anatomy discussed
in each chapter
• Activities that help the student review key structures and processes
• Animations and Videos that illustrate many of the complex, dynamic concepts
and processes of behavioral neuroscience
• Media Clips that highlight interesting topics in the chapters (NEW for this edition)
• “A Step Further” essays that offer expanded coverage of selected topics
• Visual Summaries that link to all the Activities, Animations, and Videos,
forming a complete review of each chapter
• Study Questions that help the student master the full range of material in each chapter
• Flashcards that review and reinforce the many new terms introduced in each chapter
• Complete Glossary that provides quick access to definitions of all the important
terminology in the textbook

BioPsychology NewsLink (bn9e.com/news)

This invaluable online resource helps students make connections between the science
of behavioral neuroscience and their daily lives and keeps them apprised of the lat-
est developments in the field. The site includes links to thousands of news stories, all
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ences for course and lecture development, and a list of the chapter’s key terms.
• Videos: A robust collection of video segments from the BBC and other sources
bring to life many important concepts discussed in the textbook. These videos
can be used as excellent lecture-starters and/or discussion topics.
• Animations: These detailed animations from the Companion Website help
enliven lectures and illustrate dynamic processes.
• Animation Quizzes: These quizzes test the student’s understanding of the topic
(NEW for this edition).
• Chapter Quizzes: Quiz questions for each chapter in two formats: Available
in Blackboard, Canvas, D2L platform, or as MS Word files.
• Multiple choice tests student comprehension of the material covered in
each chapter.
• Essays challenge students to synthesize and apply what they have learned.
• Test Bank: The Test Bank consists of a broad range of questions covering key facts
and concepts in each chapter. Multiple choice, essay, and paragraph development
questions are included. Questions are ranked according to Bloom’s Taxonomy and
referenced to specific textbook sections. NEW for this edition, questions are also
aligned to the textbook Learning Objectives. (Available in Blackboard, Canvas,
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ME DIA AND SUP P LEMENTS  XIX

 Psychiatry & Behavioral Sciences, Stanford Medical School
Care System, Palo Alto, CA and Dr. Ahmad Salehi, Dept. of
Introduction

Courtesy of Dr. Sarah Moghadam, VA Palo Alto Health

Scope and Outlook
Machine or Human?
In the near future depicted in the HBO series Westworld, people visit a theme park set
in the Old West, with steam locomotives, saloons, and brothels, populated with an-
droids, called “hosts,” to entertain humans. The mechanical hosts provide their guests
with anything, from casual banter to gunfights, harmless flirting to kinky sex, the only

1
restriction being that the robots are never to harm the humans. The android hosts are
so lifelike in appearance and behavior that visitors may have a hard time distinguishing
whether someone is a fellow guest or a robot. To make the androids’ simulation of hu-
mans complete, they are given backstories, false memories of a life before their appear-
ance for each new batch of guests. Importantly, none of the androids know that they
are mechanical beings rather than humans. It’s probably not much of a spoiler to say
that several plot lines in the series hinge on androids slowly discovering their true nature,
moving from shock and shame that they are mere machines, to openly rebelling from
the notion that they are to be used, and abused, as mere playthings for the humans.
We aren’t told too much about how the android “brains” in Westworld work, be-
cause, of course, such technology remains far outside our grasp, so the writers,
reduced to mere speculation, remain rather vague. But apparently the knowledge and
personality for any particular android lies in a “control unit,” a golf-ball-size device that
can be extracted from the head of one host and implanted into the head of another,
interchangeable body. Presumably, if we had enough knowledge and surgical skill, we
could remove your brain from your head and connect it up to the head of some other
body. Would you still be you? Even if we put your brain into a body of the opposite sex?
Come to think of it, are you entirely sure there is a brain in your head, and not one of
those control units?

Our aim in this book is to help you learn what is known so far about how brains work,
and about how much more we have yet to learn. We will explore the many ways in
which the structures and actions of the brain produce mind and behavior. But that is
only half of our task. We are also interested in the ways in which behavior and experi-
ence modify the structures and actions of the brain. One of the most important lessons
we want to convey is that interactions between brain and behavior are reciprocal. The
brain controls behavior and, in turn, behavior and experience alter the brain.
We hope to give an interesting account of the main ideas and research in be-
havioral neuroscience, which is of great popular as well as scientific interest. Most
important, we try to communicate our own interest and excitement about the mys-
teries of mind and body.

Go to Brain Explorer
bn9e.com/be1

neuron Also called nerve cell. The basic
unit of the nervous system.
2 C HA PT E R 1
1.1 The Brain Is Full of Surprises
Learning Objectives
After reading this section, you should be able to:
1.1.1 
Name the main type of cells found in the brain, and name the connections
between them.
1.1.2 
List the names of some of the many fields of study related to behavioral
neuroscience.
1.1.3 
Describe five different perspectives taken in understanding the biology
of behavior.
I used to think that the brain was the most wonderful organ in my body. Then I
realized who was telling me this.
—Emo Philips
(American comedian)
Of course we should always consider the source when evaluating an idea, but even
so, the brain indeed seems like a pretty wonderful organ. For one thing, brains pro-
duced the entire extent of human knowledge, everything we understand about the
universe, however limited that may be. Brains also produced every written descrip-
tion of that hard-won knowledge (including this book you hold in your hands), as
well as every work of visual art, from doodles to the sweeping frescos on the ceiling
of the Sistine Chapel.
Most of us have a hard time grasping the idea of a billion of anything, but your
head contains an estimated 86 billion nerve cells, or neurons (from the Greek word
for “nerve” or “cord”) (Herculano-Houzel, 2012). Each neuron contacts many other
cells at points called synapses, so there are trillions of those between your ears. A
specialized extension of neurons, called an axon, is microscopically slender, yet it may
be several feet long. We’ll learn that axons produce electrical impulses that travel
hundreds of miles per hour. FIGURE 1.1 offers a list of just a few of the things we will
© Dwayne Godwin, 2011
1.1 Your Brain by the Numbers The cerebral cortex is the outermost portion of the brain.
learn about the human brain in the course of this book. All this hardware isn’t just for neuroscience The study of the
show—it allows you to take in all the information in that figure in less than a minute. nervous system.
behavioral neuroscience Also called
What is behavioral neuroscience? biological psychology. The study of the
No treaty or trade union agreement defines the boundaries of behavioral neuroscience. neural bases of behavior and mental
The first people to study the relationships between brain and behavior regarded them- processes.
selves as philosophers, and their findings contributed to the births of biology and psy-
chology. Those disciplines merged in the twentieth century to form biological psychology,
the field that relates behavior to bodily processes. With the modern explosion of neuro-
science, the study of the brain, this research has evolved to the point that behavioral
neuroscience offers a more accurate description. Whichever name is used, the main
goal of this field is to understand the neuroscience underlying behavior and experience.
Behavioral neuroscience is a field that includes many players who come from quite
different backgrounds: psychologists, biologists, physiologists, engineers, neurolo-
gists, psychiatrists, and many others. Thus, there are many career opportunities, in
both universities and private industry, for people with interests in this field (Hitt,
2007). FIGURE 1.2 maps the relations of behavioral neuroscience to these many oth-
er disciplines. Clearly, the behavioral neuroscience umbrella opens very wide.

Cognitive
science
Computer
Anthropology
science

Cognitive
psychology
Evolutionary Sociobiology Artificial Psychiatry
biology intelligence
Cognitive
Behavioral neuroscience Behavioral
ecology/ethology Social Neural medicine
neuroscience modeling
Comparative/ Health
Paleontology evolutionary psychology Neurology
Paleoneuro- psychology Clinical
Cognitive neuro-
anatomy Comparative
neuro- psychology
neuroanatomy BEHAVIORAL psychology
Neural NEUROSCIENCE Neuro-
Neuro- imaging physiology Electro-
anatomy physiology
Anatomy Developmental Psycho- Physiology
psychobiology pharmacology
Behavior Psychoneuro-
Developmental genetics immunology Pharmacology
neurobiology Behavioral
endocrinology
Developmental Genetics/ Neuro- Biochemistry
biology epigenetics immunology
Neuro-
endocrinology

Molecular Immunology
biology
Endocrinology

1.2 What’s in a Name? In this graphical representation of the relationships among behavioral
neuroscience and other scientific disciplines, fields toward the center of the map are closest to
behavioral neuroscience in their history, outlook, aims, and/or methods.

In trodu ctio n 3
 Five viewpoints explore the biology of behavior
In our effort to understand the neuroscience bases of behavior, we use several dif-
ferent perspectives. Because each one yields information that complements the
others, the combination of perspectives is especially powerful. We will discuss five
major perspectives:
1. Describing behavior
2. Observing the development of behavior and its biological characteristics
over the life-span
3. Studying the biological mechanisms of behavior
4. Studying applications of behavioral neuroscience—for example, its application
to dysfunctions of human behavior
5. Studying the evolution of behavior

These perspectives are discussed in the sections that follow, and TABLE 1.1 illustrates
how each perspective can be applied to three kinds of behavior.

Behavior can be described according to different criteria

Until we describe what we want to study, we cannot accomplish much. Depending
on our goals, we may describe behavior in terms of detailed acts or processes, or in
terms of results or functions. An analytical description of arm movements might
record the successive positions of the limb or the contraction of different muscles.
A functional behavioral description, by contrast, would state whether the limb was
being used in walking or running, texting or sexting. To be useful for scientific
study, a description must be precise and reveal the essential features of the behavior,
using accurately defined terms and units.

TABLE 1.1 Five Research Perspectives Applied to Three Kinds of Behavior

Research perspective Sexual behavior
DESCRIPTION
Structural What are the main patterns
of reproductive behavior
and sex differences in
behavior?
Functional How do specialized patterns
of behavior contribute
to mating and to care of
young?
ONTOGENY How do reproductive
(development) and secondary sex
characteristics develop
over the life-span?
MECHANISMS What neural circuits and
hormones are involved in
reproductive behavior?
APPLICATIONS Low doses of testosterone
restore libido in some
postmenopausal women.
EVOLUTION How does mating depend
on hormones in different
species?
Learning and memory
In what main ways does
behavior change as a
consequence of experience—
for example, conditioning?
How do certain behaviors lead
to rewards or avoidance of
punishment?
How do learning and memory
change as we grow older?
What anatomical and chemical
changes in the brain hold
memories?
Gene therapy and behavioral
therapy improve memory in
some senile patients.
How do different species
compare in kinds and speed
of learning?
Language and
communication
How are the sounds of speech
patterned?
What behavior is involved in
making statements or asking
questions?
What changes in the brain
when a child learns to
speak?
What brain regions are
particularly involved in
language?
Speech therapy, in conjunction
with amphetamine treatment,
speeds language recovery
following stroke.
How did the human speech
apparatus evolve?

4 C HA PT E R 1
The body and behavior develop over the life-span conserved In the context of evolution,
referring to a trait that is passed on from
Ontogeny is the process by which an individual changes in the course of its life-
a common ancestor to two or more
time—that is, grows up and grows old. Observing the way in which a particular descendant species.
behavior changes during ontogeny may give us clues to its functions and mech-
ontogeny The process by which an
anisms. For example, we know that learning ability in monkeys increases over individual changes in the course of its
the first years of life. Therefore, we can speculate that prolonged maturation of lifetime—that is, grows up and grows old.
brain circuits is required for complex learning tasks. In rodents, the ability to form
long-term memories lags somewhat behind the maturation of learning ability. So,
young rodents learn well but forget more quickly than older ones, suggesting that
learning and memory involve different processes. Studying the development of
reproductive capacity and of differences in behavior between the sexes, along with
changes in body structures and processes, throws light on body mechanisms un-
derlying sexual behaviors.

Biological mechanisms underlie all behavior

To learn about the mechanisms of an individual’s behavior, we study how his or her
present body works. To understand the underlying mechanisms of behavior, we must
regard the organism (with all due respect) as a “machine,” made up of billions of
neurons. We must ask, How is this thing constructed to be able to do all that? These
are sometimes described as proximate questions—questions about the physical inter-
actions that control a particular behavior. How cells in your eye respond differently
to light of different wavelengths is a proximate question. On the other hand, why
color vision, once it arose, benefited our ancestors is an evolutionary question.
Our major aim in behavioral neuroscience is to examine body mechanisms that
make particular behaviors possible. In the case of learning and memory, for example,
we would like to know the sequence of electrical and biochemical processes that oc-
cur when we learn something and retrieve it from memory. What parts of the nervous
system are involved in that process? In the case of reproductive behavior, we also want
to understand the neuronal and hormonal processes that underlie mating behaviors.

Research can be translated to address human problems

Like other sciences, behavioral neuroscience is also dedicated to improving the
human condition. Numerous human diseases involve malfunctioning of the brain.
Many of these are already being alleviated as a result of research in the neuro-
sciences, and the prospects for continuing advances are good. Attempts to apply
knowledge also benefit basic research. For example, the study of memory disorders
in humans has pushed investigators to extend our knowledge of the brain regions
involved in different kinds of memory (see Chapter 17).

We compare species to learn how the brain and behavior

have evolved
Nature is conservative. Once particular features of the body or behavior evolve, they
may be maintained for millions of years and may be seen in animals that otherwise
appear very different. For example, the electrical messages used by nerve cells (see
Chapter 3) are essentially the same in a jellyfish, a cockroach, and a human being.
Some of the chemical compounds that transmit messages through the bloodstream
(hormones) are also the same in diverse animals (see Chapter 5). Species share
these conserved characteristics because the features first arose in a shared ancestor
(BOX 1.1 on the next page). But mere similarity of a feature between species does
not guarantee that the feature came from a common ancestral species. Similar solu-
tions to a problem may have evolved independently in different classes of animals.
Charles Darwin’s theory of evolution through natural selection is central to
all modern biology. From this perspective emerge two rather different emphases:
(1) the continuity of behavior and biological processes among species that reflects
shared ancestry and (2) the species-specific differences in behavior and biology that
have evolved as adaptations to different environments.

In tro ductio n 5
 B OX
1.1 We Are All Alike, and We Are All Different
Each person has some characteristics shared by…
all
animals…
All animals
use DNA to
store genetic
information.

all
vertebrates…

All vertebrates
have a backbone
and spinal cord.

all
mammals…
Whether knowledge gained about
All mammals a process in another species applies
suckle their to humans depends on whether we
young.
are like that species in regard to that
process. The fundamental research on
all the mechanisms of inheritance in the
primates…
bacterium Escherichia coli proved so
All primates have a
hand with an opposable widely applicable that some molecular
thumb and a relatively biologists proclaimed, “What is true
large, complex brain.
of E. coli is true of the elephant.” To a
remarkable extent, that statement is
all true, but there are also some important
humans differences in the genetic mechanisms
(people)…
All humans use symbolic of E. coli and mammals.
language to communicate With respect to each biological
with each other.
property, researchers must deter-
mine how animals are identical and
How do similarities and differ- how they are different. When we seek
some
people… ences among people and animals animal models for studying human
fit into behavioral neuroscience? behavior or biological processes, we
Some people like Each person is in some ways like must ask the following question: Does
to eat beets (no one
knows why). all other people, in some ways like the proposed animal model really
some other people, and in some have some things in common with the
ways like no other person. As the process at work in humans (Seok et
figure shows, we can extend this al., 2013)? We will see many cases in
No two people, even observation to the much broader which it does.
identical twins, are alike
no other in each and every way, as range of animal life. Each person is Even within the same species,
person. individual experiences in some ways like all other animals however, individuals differ from one
leave their unique stamp (e.g., needing to ingest complex another: cat from cat, blue jay from
on every brain.
organic nutrients), in some ways like blue jay, and person from person.
all other vertebrates (e.g., having a Behavioral neuroscience seeks to
spinal column), in some ways like understand individual differences
all other mammals (e.g., nursing our as well as similarities. Therefore, the
young), and in some ways like all way in which each person is able to
other primates (e.g., having a hand process information and store the
with an opposable thumb and a memories of these experiences is
relatively large, complex brain). another part of our story.
Behavioral Neuroscience 9E
Breedlove
Sinauer Associates

Breedlove9e_Box01.01.ai Date 07-15-19

6 C HA P T E R 1
 1.2 Three Approaches Relate Brain and Behavior somatic intervention An approach
to finding relations between body variables
Learning Objectives and behavioral variables that involves
manipulating body structure or function and
After reading this section, you should be able to:
looking for resultant changes in behavior.
1.2.1 
Differentiate between the independent and dependent variables in
scientific experiments. independent variable The factor that
is manipulated by an experimenter.
1.2.2 
Name the type of research in which a part of the brain is manipulated to
observe effects on behavior, and offer examples. dependent variable The factor that
1.2.3 
Name the type of research in which behavior or experience is manipulated an experimenter measures to monitor a
to observe effects on the brain, and offer examples. change in response to manipulation of an
1.2.4 
Describe correlational research about the brain and behavior, and independent variable.
offer examples.
1.2.5 
Explain why the brain must be capable of changing its structure, and name
the term to describe that changeability.

Behavioral neuroscientists use three approaches to understand the relationship be-

tween brain and behavior: somatic intervention, behavioral intervention, and cor-
relation. In the most common approach, somatic intervention (FIGURE 1.3A), we
alter a structure or function of the brain or body to see how this alteration changes
behavior. Here, somatic intervention is the independent variable, and the behav-
ioral effect is the dependent variable; that is, the resulting behavior depends on
how the brain has been altered. For example, in response to mild electrical stimula-
1.3 Three Main Approaches to
tion of one part of her brain, not only did one patient laugh, but she found whatever
Studying the Neuroscience of Behavior
she happened to be looking at amusing (Fried et al., 1998). (A) In somatic intervention, investigators
In later chapters we describe many kinds of somatic intervention with both hu- change the body structure or chemistry of
mans and other animals, as in the following examples: an animal in some way and observe and
• A hormone is administered to some animals but not to others; various behaviors measure any resulting behavioral effects.
(B) Conversely, in behavioral intervention,
of the two groups are later compared.
researchers change an animal’s behavior
• A part of the brain is stimulated electrically, or by use of light to stimulate only a or its environment and try to ascertain
particular class of neurons, and behavioral effects are observed. whether the change results in physiological
or anatomical changes. (C) Measurements
• A connection between two parts of the nervous system is cut, and changes in
of both kinds of variables allow researchers
behavior are measured. to arrive at correlations between somatic
changes and behavioral changes. (D) Each
approach enriches and informs the others.
(A) Manipulating the body may affect behavior (B) Experience affects the body (including the brain)
Somatic interventions Behaviors affected Somatic effects Behavioral interventions
Strength of mating Changes in hormone Put male in presence
Administer a hormone
behavior levels of female

Stimulate brain region Movement toward Changes in electrical Present a visual

electrically goal object activity of brain stimulus

Cut connections between Recognition of Anatomical changes

parts of nervous system stimulus Give training
in nerve cells

(C) Body and behavioral measures covary (D) Behavioral neuroscience seeks to understand all
Somatic variables Behavioral variables these relationships

Brain size Correlations Learning scores

Somatic
intervention
Strength of mating
Hormone levels Correlations
behavior Somatic variables Correlations Behavioral variables

Enlarged cerebral Schizophrenic Behavioral

Correlations intervention
ventricles symptoms

Introductio n 7
behavioral intervention An approach
to finding relations between body variables
and behavioral variables that involves
intervening in the behavior of an organism
and looking for resultant changes in body
structure or function.
correlation The covariation of
two measures.
neuroplasticity Also called neural
plasticity. The ability of the nervous
system to change in response to
experience or the environment.
Go to Media Clip 1.1
Neuroplasticity
bn9e.com/mc1.1
8 C HA P T E R 1
The approach opposite to somatic intervention is psychological or behavioral
intervention (FIGURE 1.3B). In this approach, the scientist intervenes in the be-
havior or experience of an organism and looks for resulting changes in body struc-
ture or function. Here, behavior is the independent variable, and change in the
body is the dependent variable. Among the examples that we will consider in later
chapters are the following:
• Putting two adults of opposite sex together may lead to increased secretion of
certain hormones.
• Exposing a person or animal to a visual stimulus provokes changes in electrical
activity and blood flow in parts of the brain.
• Training of animals in a maze is accompanied by electrical, biochemical, and
anatomical changes in parts of their brains.
The third approach to brain-behavior relations, correlation (FIGURE 1.3C), con-
sists of finding the extent to which a given body measure varies with a given behav-
ioral measure. Later we will examine the following questions, among others:
• Are people with large brains more intelligent than people with smaller brains
(a topic we’ll take up later in this chapter)?
• Are individual differences in sexual behavior correlated with levels of certain
hormones in the individuals?
• Is the severity of schizophrenia correlated with the magnitude of changes in
brain structure?
Such correlations should not be taken as proof of causal relationship. For one thing,
even if a causal relation exists, the correlation does not reveal its direction—that is,
which variable is independent and which is dependent. For another, two factors might
be correlated only because a third, unknown factor affects the two factors measured.
If you and your study partner get similar scores on an exam, that’s not because your
performance caused her to get the score she did, or vice versa. What a correlation does
suggest is that the two variables are linked in some way—directly or indirectly. Such
a correlation often stimulates investigators to formulate hypotheses and to test them
by somatic or behavioral intervention. Only by moving on to such intervention ap-
proaches can we establish whether one variable is causing changes in the other.
Combining these three approaches yields the circle diagram of FIGURE 1.3D, incor-
porating the basic approaches to studying relationships between bodily processes and
behavior. It also emphasizes the theme that the relations between brain and behavior
are reciprocal: each affects the other in an ongoing cycle of bodily and behavioral in-
teractions. We will see examples of this reciprocal relationship throughout the book.
Neuroplasticity: behavior can change the brain
The idea that there is a reciprocal relationship between brain and behavior has embed-
ded within it a concept that is, for most people, startling. When we say that behavior and
experience affect the brain, we mean that they, literally, physically alter the brain. The
brain of a child growing up in a French-speaking household assembles itself into a con-
figuration different from that of the brain of a child who hears only English. That’s why
the first child, as an adult, understands French effortlessly while the second does not. In
this case we cannot tell you what the structural differences are exactly, but we do know
one part of the brain that is being altered by these different experiences (see Chapter 19).
Numerous examples, almost all in animal subjects, show that experience can
affect the number or size of neurons, or the number or size of connections between
neurons. This ability of the brain, both in development and in adulthood, to be
changed by the environment and by experience is called neuroplasticity (or neural
plasticity, or simply plasticity).
Today when we hear the word plastic, we think of the class of materials found in
so many modern products. But originally, plastic meant “flexible, malleable” (from the
Greek plassein, “to mold or form”), and the modern materials were named plastics be-
cause they can be molded into nearly any shape. In 1890, William James (1842–1910)
described plasticity as the possession of a structure weak enough to Only in this brain region
yield to an influence but strong enough not to yield all at once. 12
was growth stunted by the
In the ensuing years, research has shown that the brain is even lack of opportunity to play.
more plastic, more yielding, than James suspected. For example, parts 10

Volume (mm3 × 10–1)

of neurons known as dendritic spines (see Chapter 2) appear to be in Social
8
constant motion, changing shape in the course of seconds. We will Isolated
see many examples in which experience alters the structure and/or 6
function of the brain: In Chapter 5, you’ll read that hearing a baby cry
4
causes the mother’s brain to secrete a hormone. In Chapter 7, we’ll see
that visual experience in kittens directs the formation of connections 2
in the brain. In Chapter 12, we’ll discuss how a mother rat’s grooming
of her pups affects the survival of spinal cord neurons. And Chapter 0
Posterodorsal Anterodorsal Anteroventral Posteroventral
17 talks about how a sea slug learning a task changes the connections
Quadrants of the medial amygdala
between two particular neurons.
1.4 The Role of Play in Brain
Behavioral neuroscience and social psychology are related Development A brain region involved in
The plasticity of the human brain has a remarkable consequence: other individuals processing odors (the posterodorsal por-
can affect the physical structure of your brain! Indeed, the whole point of coming to a tion of the medial amygdala) was smaller
in male rats housed individually than in
lecture hall is to have the instructor use words and figures to alter your brain so that
males housed together and allowed to
you can retrieve that information in the future (in other words, teach you something). play. Other nearby regions were identical
Many of these alterations in your brain last only until you take an exam, but every once in the two groups. (After B. M. Cooke et
in a while the instructor may tell you something that you’ll remember for the rest of al., 2000. Behav Brain Res 117: 107–113.)
your life. Most aspects of our social behavior are learned—from the language we speak
to the clothes we wear and the kinds of food we eat—so the mechanisms of learning
and memory (see Chapter 17) are important for understanding social behavior.
For an example from an animal model, consider the fact that rats spend a lot
of time investigating the smells around them, including those coming from other
rats. Cooke et al. (2000) took young male rats, just weaned from their Behavioral
mother, and
Neuroscience 9E
raised them in two different ways: either alone in separate cages, orBreedlove
with other
males in group cages so they could engage in play (including a lot ofSinauer Associates
sniffing of
each other’s butts). Examination of these animals as adults found only one brain
Breedlove9e_1.04.ai Date 07-15-19
difference between the groups: a region of the brain known to process odors was
smaller in the isolated males than in the males raised with playmates (FIGURE
1.4). Was it the lack of play (N. S. Gordon et al., 2003), the lack of odors to investi-
gate, or the stress of isolation that made the region smaller? Whatever the mecha-
nism, social experience affects this brain structure. In Chapter 17 we’ll see more
examples of social experience altering the brain.
Here’s an example of how social influences can affect human brain function.
When people were asked to put a hand into moderately hot water (47°C), part of
the brain became active, presumably because of the discomfort involved (Rain-
ville et al., 1997). But people who were led
to believe the water would be very hot had
a more activated brain than did those led to
believe the discomfort would be minimal

From P. Rainville, 1998. Presented at INABIS '98, 5th Internet

(FIGURE 1.5), even though the water was

University, Canada, Dec 7–16th. Available at http://www.

the same temperature for everyone. The so- mcmaster.ca/inabis98/woody/rainville0419/index.html

World Congress on Biomedical Sciences at McMaster

cially induced psychological expectation af-
fected the magnitude of the brain response,

1.5 Pictures of Pain People told to expect

only mild discomfort from putting a hand into
47°C water (left) showed less activation in a par-
ticular brain region (the anterior cingulate cortex)
than did people expecting more discomfort (right)
from water of the very same temperature. Areas
of high activation are indicated by orange, red,
and white.

In trodu ctio n 9

reductionism The scientific strategy of
breaking a system down into increasingly
smaller parts in order to understand it.
levels of analysis The scope of
experimental approaches. A scientist may
try to understand behavior by monitoring
molecules, nerve cells, brain regions, or
social environments, or some combination
of these levels of analysis.
10 C HAP T E R 1
even though the physical stimulus was exactly the same. (By the way, the people
with the more activated brains also reported feeling more pain.)
In most cases, biological and social factors continually interact and affect each
other in an ongoing series of events as behavior unfolds. For example, the level of the
hormone testosterone in circulation can affect dominance behavior and aggression
(see Chapter 15). The dominance may be exhibited in a great variety of social settings,
ranging from playing chess to physical aggression. In humans and other primates,
the level of testosterone correlates positively with the degree of dominance and with
the amount of aggression exhibited. Winning a contest, whether a game of chess or a
boxing match, raises the level of testosterone; losing a contest lowers the level. Thus,
at any moment the level of testosterone is determined, in part, by recent dominant-
submissive social experience, and the level of testosterone determines, in part, the de-
gree of dominance and aggression in the future. Of course, social and cultural factors
also help determine the frequency of aggression; cross-cultural differences in rates of
aggression exist that cannot be correlated with hormone levels, and ways of express-
ing aggression and dominance are influenced by sociocultural factors.
Perhaps nothing distinguishes neuroscience from the other sciences more clearly
than this fascination with neuroplasticity and the role of experience. Neuroscientists
have a pervasive interest in how experience physically alters the brain and therefore
affects future behavior. We will touch on this theme in every chapter of this book.
1.3 Behavioral Neuroscientists Use Several Levels
of Analysis
Learning Objectives
After reading this section, you should be able to:
1.3.1 
Name and describe the scientific approach of explaining mechanisms at
simpler and simpler levels.
1.3.2 
Give a survey of important ongoing questions about the relationship
between the brain and behavior.
1.3.3 Offer estimates of the extent of neurological and psychiatric disorders.
1.3.4 
Explain the importance of research with animals for neuroscience, and
discuss the ethics of such research.
Scientific explanations of systems or structures or functions usually involve break-
ing them down into smaller parts, as a way of understanding them. This approach is
known as reductionism. In principle, it is possible to reduce each explanatory series
down to the molecular or atomic level, though for practical reasons this extent of
reductionism is rare. For example, most chemists deal with large, complex molecules
and the laws that govern them; seldom do they seek explanations in terms of sub-
atomic quarks and bosons.
Understanding behavior often requires several levels of biological analysis. The units
of each level of analysis are simpler in structure and organization than those of the level
above. The levels of analysis range from social interactions to the brain, continuing to
successively less complex units until we arrive at single nerve cells and their even sim-
pler, molecular constituents.
Naturally, in all fields different problems are carried to different levels of analy-
sis, and fruitful work is often being done simultaneously by different workers at
several levels (FIGURE 1.6). Thus, in their research on visual perception, cognitive
neuroscientists advance analytical descriptions of behavior. They try to determine
how the eyes move while looking at a visual pattern, or how the contrast among
parts of the pattern determines its visibility. Meanwhile, other behavioral neurosci-
entists study the differences in visual abilities among species and try to determine
the adaptive significance of these differences. For example, how is the presence
(or absence) of color vision related to the life of a species? At the same time, other
investigators trace out brain structures and networks involved in different kinds of
visual discrimination. Still other scientists try to ascertain the electrical and chemi-
cal events that occur in the brain during vision.
 Social level: Neural systems level:
Individuals behaving Eyes and visual brain regions
in social interaction Organ level:
Brain, spinal cord, Brain region level:
peripheral nerves, Visual cortex
and eyes

Circuit level:
Local neural circuit

Cellular level:
Single neuron

Molecular level
Synaptic level

Membrane receptors

1.6 Levels of Analysis in Behavioral Neuroscience The scope of behavioral neuro-

science ranges from the level of the individual interacting with others, to the level of the mol-
ecule. Depending on the question at hand, investigators use different techniques to focus
on these many levels, but always with an eye toward how their findings apply to behavior.

We will encounter many diverse brain and behavior topics

Here areNeuroscience
Behavioral some examples
9E of research topics considered in this book:
Breedlove
• How does the brain grow, maintain, and repair itself over the life-span (see
Sinauer Associates
Chapter 7), and how are these capacities related to the growth and development
Breedlove9e_1.06.ai
of the mind and behaviorDatefrom07-15-19
the womb to the tomb?
• How does the nervous system capture, process, and represent information about
the environment? For example, sometimes brain damage causes a person to lose
the ability to identify other people’s faces (see Chapter 18); what does that tell us
about how the brain recognizes faces?
• How does sexual orientation develop? Some brain regions are different
in straight versus gay men (see Figure 12.26); what, if anything, do those
differences tell us about the development of human sexual orientation?
• Some people suffer damage to the brain and afterward seem alarmingly unafraid
in dangerous situations and unable to judge the fearfulness of other people; what
parts of the brain are damaged to cause such changes (see Figure 15.16)?
• How does the brain manage to change during learning (see Chapter 17), and
how are memories retrieved?
• What brain sites and activities underlie feelings and emotional expression? Are
particular parts of the brain active in romantic love, for example (FIGURE 1.7A)?
• Why are different brain regions active during different language tasks
(FIGURE 1.7B)?

Introdu ctio n 11
 (A) (B)

From A. Bartels and S. Zeki, 2000.

Hearing words Seeing words

Neuroreport 11: 3829–3834

Courtesy of Marcus Raichle

Reading words Generating words

1.7 “Tell Me Where Is Fancy Bred, Or in the Heart Or in the Head?” (A) The parts
of the brain highlighted here become especially active when a person thinks about his or
her romantic partner. (B) Different brain regions are activated when people perform four
different language tasks. The techniques used to generate such images are described in
Chapter 2.

The relationship between the brain and behavior is, on the one hand, very mys-
terious because it is difficult to understand how a physical device, the brain, could
be responsible for our subjective experiences of fear, love, and awe. Yet despite
this mystery, we all use our brains every day. Perhaps it is the “everyday miracle”
Behavioral Neuroscience 9E aspect of the topic that has generated so much folk wisdom about the brain. Think
Breedlove
Sinauer Associates of it as “neuromythology.”
Sometimes these popular ideas about the brain are in line with our current
Breedlove9e_1.07.ai Date 07-15-19
knowledge, but in many cases we know they are false. For example, the notion
that we normally use only 10% of our brain is commonplace—a survey of teachers
found that nearly half of them agreed with this notion (Howard-Jones, 2014)—but
it is patent nonsense. Brain scans make it clear that the entire brain is activated by
even fairly mundane tasks. Indeed, although the areas of activation shown in Fig-
ure 1.7 appear rather small and discrete, we will show in Box 2.3 that experimenters
must work very hard to create images that separate activation related to a particular
task from the background of widespread, ongoing brain activity.
We offer a list of other commonly held beliefs about the brain and behavior on
the website in A STEP FURTHER 1.1 : NEUROMYTHOLOGY: FACTS OR FABLES?
Throughout the book we offer such opportunities for you to explore a given topic in
more detail on the website, bn9e.com.

Behavioral neuroscience contributes to our understanding of

human disorders
One of the great promises of neuroscience is that it can help us understand brain
disorders and devise treatment strategies. Like any other complex mechanism, the
brain is subject to a variety of malfunctions and breakdowns. People afflicted by
disorders of the brain are not an exotic few—a European survey estimated that at
least 38% of the population would suffer from a mental disorder at some point in
a typical year (Wittchen et al., 2011). At least one person in five around the world
currently has neurological and/or psychiatric disorders that vary in severity from
mild challenges to complete disability. FIGURE 1.8A shows the estimated numbers
of U.S. residents afflicted by some of the main neurological disorders. FIGURE

12 C HA P T E R 1
(A) Prevelence of neurological disorders (B) Prevelance of psychiatric disorders 1.8 The Toll of Brain Disorders Estimated
numbers of people in the United States with
Other neurological disorders (A) and number of
dementias people worldwide with psychiatric disorders
2,200,000 Depression (B). (Part A after C. L. Gooch et al., 2017. Ann
Alzheimer’s 268,000,000
5,300,000 Neurol 81: 479–484; B after H. Ritchie and M.
Epilepsy
2,800,000 Alcohol use Roser, 2019. "Mental Health". Published online
109,000,000 at OurWorldInData.org. Retrieved from: https://
ourworldindata.org/mental-health. Underlying
Drug use
Anxiety data available from http://ghdx.healthdata.org/
Stroke 73,000,000
disorders gbd-results-tool.)
6,800,000 289,000,000

Traumatic
brain injury
1,700,000

Spinal cord MS Parkinson’s Eating Schizophrenia Bipolar

injury 600,000 disease disorders 19,500,000 disorder
340,000 1,000,000 16,000,000 47,000,000

1.8B gives estimates of the numbers of adults worldwide with certain psychiatric
disorders. The percentage of U.S. adults suffering from mental illness may be in-
creasing (Twenge, 2015; Twenge et al., 2010).
The toll of these disorders is enormous, in terms of both individual suffering
and social costs (Demyttenaere et al., 2004). The National Advisory Mental Health
Council has estimated that direct and indirect costs of behavioral and brain dis-
orders amount to $400 billion a year in the United States alone. For example, the
cost for treatment of dementia (severely disordered thinking) exceeds the costs
of treating cancer and heart disease combined. The World Health Organization
(2004) estimates that over 15% of all disease burden, in terms of lost productivity,
is due to mental disorders. The high cost in suffering and expense has compelled
researchers to try to understand the mechanisms involved in these disorders and
to try to alleviate or even prevent them.
In this quest, the distinction between clinical and laboratory approaches begins 1.9 Identical Twins but Nonidentical
Brains and Behavior In these images
to fade away. For example, when clinicians encounter a pair of twins, one of whom of the brains of identical twins, the fluid-
has schizophrenia while the other seems healthy, the discovery of structural dif- filled cerebral ventricles are prominent
ferences in their brains (FIGURE 1.9) immediately raises questions for laboratory as dark “butterfly” shapes. The brain of
scientists: Did the structural differences arise before the symptoms of schizophre- the twin with schizophrenia (A) has the
nia, or the other way around? Were the brain differences present at birth, or did enlarged cerebral ventricles that some
they arise during puberty? Does medication that reduces symptoms affect brain researchers believe are characteristic of
structure? When genes associated with schizophrenia in people are introduced this disorder. The other twin does not
have schizophrenia; his brain (B) clearly
into mice, will that change the mouse brains (see Figure 16.7)? This has smaller ventricles.
last question is just one instance of when working with animals is
essential, an issue we address next. (A) Twin with (B) Unaffected twin
schizophrenia
Animal research makes vital contributions
Because we will draw on animal research throughout this book, we
want to comment on some of the ethical issues of experimentation on
animals. Human beings’ involvement and concern with other species
predates recorded history. Early humans had to study animal behavior
MRIs courtesy of E. Fuller Torrey

and physiology in order to escape some species and hunt others. To

study biological bases of behavior inevitably requires research on ani-
and Daniel Weinberger

mals of other species as well as on human beings. Psychology students

usually underestimate the contributions of animal research because
the most widely used introductory psychology textbooks often pres-
ent major findings from animal research as if they were obtained with
human participants (Domjan and Purdy, 1995).

Behavioral Neuroscience 9e
Fig. 01.08
Dragonfly Media Group Intro ductio n 13
07/25/19

© Santa Cruz Sentinel/ZUMAPRESS.com
1.10 Car Firebombed by Animal
Rights Activists The extremists target-
ed the cars and homes of two scientists
who work with animals at the University
of California in Santa Cruz in 2008. The
next year, the car of a researcher at
UCLA was torched.
14 C HAP T E R 1
Because of the importance of carefully reg-
ulated animal research for both human and
animal health and well-being, the National
Research Council (NRC Committee on Ani-
mals as Monitors of Environmental Hazards,
1991) undertook a study on the many uses
of animals in research. The study noted that
93% of the mammals used in research are
laboratory-reared rodents. It also reported that
most Americans believe that animal research
should continue. Of course, researchers have
an obligation to minimize the discomfort of
their animal subjects, and ironically enough,
animal research has provided us with the
drugs and techniques to make most research
painless for the animal subjects (Sunstein and
Nussbaum, 2004).
Nevertheless, a very active minority of peo-
ple believe that research with animals, even if
it does lead to lasting benefits, is unethical. For
example, in his 1975 book Animal Liberation, Peter Singer asserts that research with
animals can be justified only if it actually produces benefits. The trick, of course,
is how to predict which experiment will lead to a breakthrough. Thus Singer re-
fuses to say that animal experimentation is never justified (Neale, 2006). In the
meantime, animal rights groups have vandalized labs, burned down buildings,
and exploded bombs in laboratories (Conn and Parker, 2008). In 2008, animal
rights extremists set off firebombs at the homes of two scientists in Santa Cruz,
California. One scientist’s family, including two young children, had to flee their
home through a second-story window (FIGURE 1.10) (Paddock and La Ganga,
2008). These personal attacks on individuals appear to be intended to intimidate
and frighten scientists (D. Grimm, 2014), and they have already hounded at least
one researcher out of the field (Nature Neuroscience, 2015).
Perhaps in a future where robots can be made that look and act like humans,
methods will be available to clearly see all the processes at work in a living, work-
ing human brain. In the meantime, there’s no substitute for research with animal
subjects. Every chapter in this book is teeming with information that was gathered
from humane experiments with animals.
1.4 The History of Research on the Brain and Behavior
Begins in Antiquity
Learning Objectives
After reading this section, you should be able to:
1.4.1  race the historical point at which the brain was recognized as the control
T
unit for behavior.
1.4.2  iscuss the importance of the Renaissance in better understanding human
D
anatomy.
1.4.3  xplain Descartes’s contribution to early neuroscience and his now-
E
discredited ideas of dualism.
1.4.4  race the history of phrenology and the relationship to modern thinking
T
about brain and behavior.
1.4.5  iscuss the difficult question of consciousness and the explosion of
D
neuroscience research.
Only recently have scientists recognized the central role of the brain in controlling
behavior. When Egyptian pharaoh Tutankhamen was mummified (about 1300 bce),
five important organs were preserved in his tomb: liver, lungs, stomach, intestines,
and heart. All these organs were considered necessary to ensure the pharaoh’s con-
tinued existence in the afterlife. The brain, however, was picked out through the
nostrils (FIGURE 1.11) and thrown away. Although the Egyptian version of the after-
life entailed considerable struggle, the brain was not considered an asset.
Neither the Hebrew Bible (written from the twelfth to the second century bce)
nor the New Testament ever mentions the brain. However, the Bible mentions the
heart hundreds of times and makes several references each to the liver, the stom-
ach, and the bowels as the seats of passion, courage, and pity, respectively. “Teach
us … that we may gain a heart of wisdom” (Psalms 90:12).
The heart is also where Aristotle (about 350 bce), the most prominent scientist
of ancient Greece, located mental capacities. We still reflect this ancient notion
when we call people kindhearted, openhearted, fainthearted, hardhearted, or heartless
and when we speak of learning by heart. Aristotle considered the brain to be only
a cooling unit to lower the temperature of the hot blood from the heart.
Also about 350 bce, the Greek physician Herophilus (the “Father of Anatomy”)
advanced our knowledge of the nervous system by dissecting bodies of both peo-
ple and animals. He traced nerves from muscles and skin into the spinal cord and
noted that each region of the body is connected to separate nerves.
A second-century Greco-Roman physician, Galen, treated the injuries of gladi-
ators. His reports of behavioral changes caused by injuries to the heads of gladia-
tors drew attention to the brain as the controller of behavior. Galen advanced the

Photograph by Neil Watson

idea that animal spirits—a mysterious fluid—passed along nerves to all regions of
the body. But Galen’s ideas about the anatomy of the human brain were very inac-
curate because dissecting humans was illegal in Rome at that time.

Renaissance scientists began to understand brain anatomy

The eminent Renaissance painter and scientist Leonardo da Vinci (1452–1519) stud-
ied the workings of the human body and laid the foundations of anatomical drawing.
He especially pioneered in providing views from different angles and cross-sectional 1.11 Brain Removal Kit Ancient Egyp-
tians had little regard for the brain. During
representations. His artistic renditions of the body included portraits of the nerves in mummification, they would use tools like
the arm and the fluid-filled ventricles in the brain (FIGURE 1.12). these to reach through the nostrils to pick
Renaissance anatomists emphasized the shape and appearance of the external out brain pieces and throw them away.
surfaces of the brain because these were the parts that were easiest to see when the
skull was removed. It was immediately apparent to anyone who looked that the brain

(A) Early drawing (B) Later drawing based on observation

Her Majesty Queen Elizabeth II, copyright reserved

Reproduced with gracious permission of

1.12 Leonardo da Vinci’s Changing View of the (B) Later he made a drawing based on direct observation:
Brain (A) In an early representation, Leonardo simply after making a cast of the ventricles of an ox brain by
copied old schematic drawings that represented the fluid- pouring melted wax into the brain and letting it set, he cut
filled cerebral ventricles as a linear series of chambers. away the tissue to reveal the true shape of the ventricles.

In troductio n 15

Behavioral Neuroscience 9E
Breedlove
Sinauer Associates
Breedlove9e_1.13.ai
Date 07-15-19
dualism Here, the notion promoted by
René Descartes that the mind is subject
only to spiritual interactions while the body
is subject only to material interactions.
phrenology The outmoded belief that
bumps on the skull reflect enlargements
of brain regions responsible for certain
behavioral faculties.
16 C HAP T E R 1
1.13 An Early Account of Reflexes In this depiction of an explanation by Des-
cartes, when a person’s toe touches fire, the heat causes nervous activity to flow up
the nerve to the brain (blue arrows). From there the nervous activity is “reflected” back
down to the leg muscles (red arrows), which contract, pulling the foot away from the
fire; the idea of activity being reflected back is what gave rise to the word reflex. In
Descartes’s time, the difference between sensory and motor nerves had not yet been
discovered, nor was it known that nerve fibers normally conduct in only one direction.
Nevertheless, Descartes promoted thinking about bodily processes in scientific terms,
and this focus led to steadily more accurate knowledge and concepts.
has an extraordinarily complex shape. To Renaissance artists like Michelangelo
(1475–1564), this marvelous structure was God’s greatest gift to humankind. So,
in Michelangelo’s painting on the ceiling of the Sistine Chapel, God seems to
ride the form of the human brain when bestowing life to Adam (Meshberger,
1990), while in another scene God’s throat resembles the base of the brain (Suk
and Tamargo, 2010).
In 1633, René Descartes (1596–1650) wrote an influential book (De Homine
[On Man]) in which he tried to explain how the behavior of animals, and to
some extent the behavior of humans, could be like the workings of a machine.
Descartes proposed the concept of spinal reflexes and a neural pathway for
them (FIGURE 1.13). Attempting to relate the mind to the body, Descartes sug-
gested that the two come into contact in the pineal gland, located within the brain.
He suggested the pineal gland for this role because (1) whereas most brain struc-
tures are double, located symmetrically in the two hemispheres, the pineal gland
is single, like consciousness, and (2) he believed, erroneously, that the pineal gland
exists only in humans and not in animals.
As Descartes was preparing to publish his book, he learned that the Catholic
Church had forced Galileo to renounce his teaching that Earth revolves around the
sun, threatening to execute him if he did not recant. Fearful that his own specula-
tions about mind and body could also incur the wrath of the church, Descartes
withheld his book from publication. It did not appear in print until 1662, after his
death. Descartes believed that if people were nothing more than intricate machines,
they could have about as much free will as a pocket watch, with no opportunity
to make the moral choices that were so important to the church. He asserted that
humans, at least, had a nonmaterial soul as well as a material body. This notion
of dualism spread widely and left other philosophers with the task of determin-
ing how a nonmaterial soul could exert influence over a material body and brain.
Mainstream neuroscientists reject dualism and insist that all the workings of the
mind can also, in theory, be understood as purely physical processes in the material
world, specifically in the brain.
The concept of localization of function arose in the
nineteenth century
By the end of the 1600s, the English physician Thomas Willis (1621–1675), with his
detailed descriptions of the structure of the human brain and his systematic study
of brain disorders, convinced educated people in the Western world that the brain is
the organ that coordinates and controls behavior (Zimmer, 2004). A popular notion
of the nineteenth century, called phrenology, elaborated on this idea by asserting
that the cerebral cortex consisted of separate functional areas and that each area was
responsible for a behavioral faculty such as love of family, perception of color, or curi-
osity. Investigators assigned functions to brain regions anecdotally, by observing the
behavior of individuals and inferring, from the shape of the skull, which underlying
regions of the brain were more or less developed (FIGURE 1.14A).
Opponents rejected the entire concept of localization of brain function, insisting
that the brain, like the mind, functions as a whole. Today we know that the whole
brain is indeed active when we are doing almost any task. When we are performing
particular tasks, however (as we saw earlier in this chapter), certain brain regions
become even more activated. Different tasks activate different brain regions. Modern
(A) (B)
Voluntary eye Face Motor execution
Visual spatial movements Hand
attention Foot
Anticipation Motor
Analytic preparation Face Somatosensory cortex
and figural Hand
reasoning Foot
Visual spatial
Spatial working attention
memory Analytic
reasoning
Mathematical
approximations Mathematical
approximations
Anticipation
of pain Visual spatial
attention
Object
working Motion perception
© The Print Collector/Alamy Stock Photo

memory Speed perception

Exact Primary visual
mathematical cortex
calculations
Color perception
Olfaction
Pleasant Face recognition
touch Olfaction Pain Auditory cortex

Speech Anticipation Semantic priming Spoken language

production of pain of visual words comprehension

1.14 Old and New Phrenology (A) In the early nineteenth century, certain “faculties,”
such as skill at mathematics or a tendency toward aggression, were believed to be directly
associated with particular brain regions. Phrenologists used diagrams like this one to measure
bumps on the skull, which they took as an indication of how fully developed each brain region
was in an individual, and hence how fully that person should display particular qualities.
(B) Today, technology enables us to roughly gauge how active different parts of the brain
are when a person is performing various tasks (see Chapter 2). But virtually the entire brain
is active during any task, so the localization of function that such studies provide is really a
measure of where peak activity occurs, rather than a suggestion of a single region involved in
a particular task. (B after M. J. Nichols and W. T. Newsome, 1999. Nature 402: C35–C38.)
Behavioral Neuroscience 9E
Breedlove
brain maps
Sinauer of these places
Associates where peaks of activation occur (FIGURE 1.14B) bear a
passing resemblance to their phrenological predecessors, differing only in the spe-
Breedlove9e_1.14.ai Date
cific locations of functions. But 07-15-19
unlike the phrenologists, we confirm these modern
maps by other methods, such as examining what happens after brain damage.
Even as far back as the 1860s, the French surgeon Paul Broca (1824–1880) argued
that language ability was not a property of the entire brain but rather was localized
in a restricted brain region. Broca presented a postmortem analysis of a patient
who had been unable to talk for several years. The only portion of the patient’s
brain that appeared damaged was a small region within the frontal portions of
the brain on the left side—a region now known as Broca’s area (labeled “Speech
production” in Figure 1.14B). The study of additional patients further convinced
Broca that language expression is mediated by this specific brain region rather than
reflecting activities of the entire brain.
In 1890, William James’s book The Principles of Psychology signaled the begin-
nings of a modern approach to behavioral neuroscience. The strength of the ideas
described in this book is evident from the continuing frequent citation of the work. In
James’s work, psychological ideas such as consciousness and other aspects of human
experience came to be seen as properties of the nervous system. A true behavioral
neuroscience began to emerge from this approach.
These nineteenth-century observations form the background for a continuing
theme of research in behavioral neuroscience—notably, the search for distinguishing
differences among brain regions on the basis of their structure, and the effort to re-
late different kinds of behavior to different brain regions (M. Kemp, 2001). An addi-
tional theme emerging from these studies is the relation of brain size to ability across
species (see Figure 6.9), and even across various people (BOX 1.2 on the next page).

Intro ductio n 17
 B OX
1.2 Bigger Better? The Case of the Brain and Intelligence
Does a bigger brain indicate greater
intelligence? Brain size does seem
to explain many species differences
in complex behavior, and the hu-
man brain has expanded remarkably
over the past few million years (see
Chapter 6). But do variations in brain
size within our species correlate with
intelligence? This question has been
the subject of lively controversy for at
least two centuries. Sir Francis Galton
(1822–1911), who invented the correla-
tion coefficient, stated that the great-
est disappointment in his life was his
failure to find a significant relationship
between head size and intelligence.
But Galton had to use head size, when
he really wanted to measure brain
size. In addition, he had to rely on
teachers’ estimates of their students’
intelligence, and every student knows
that teachers can be quite wrong.
Other investigators in the nineteenth
century measured the volumes of
skulls of various groups and estimated
intelligence on the basis of people’s
occupations or other doubtful criteria
(S. J. Gould, 1981).
(A)
Courtesy of Nancy Andreasen
(A) Examples of measuring specific features of brain size from scans.
18 C HA P T E R 1
Behavioral Neuroscience 9E
The development and standardiza- est thickening of the outer layer of the
tion of intelligence quotient (IQ) tests in brain, especially in the front (P. Shaw
the twentieth century provided invalu- et al., 2006). Other brain-imaging
able help for one side of the question, studies report correlations between IQ
and these scores indeed correlate, scores and the extent of connectivity
with ranges from +0.08 to +0.22, between brain regions of about +0.50
with estimates of brain size based on (Figure B) (Haász et al., 2013; Malpas
known head size (Van Valen, 1974). et al., 2016).
Newer, noninvasive techniques Thus, on the basis of modern tech-
(discussed in detail in Chapter 2) to niques, the long-standing controversy
visualize the brains of healthy people appears to have been settled in favor
now make it possible to directly mea- of a significant correlation between
sure brain size in living humans. One brain size and intelligence (as shown
study found a significant correlation in Figure B). Note, however, that the
coefficient of about +0.26 between modest size of the correlations, while
brain size and IQ (Posthuma et al., statistically significant, indicates that
2002). In another study, brain scans only about 10–20% of variability in
were used for measuring the sizes of IQ is accounted for by brain size. So,
different brain regions (Figure A). After there is plenty of room for other factors
correction for body size, the correla- to contribute to overall IQ. There is
tion between brain size and IQ scores abundant evidence that early experi-
was +0.38 (Andreasen et al., 1993). IQ ence can affect IQ (Chapter 17). In
seems to correlate better with the vol- addition, many people dispute whether
ume of the front of the brain than that IQ tests really measure a general prop-
of the back (Colom et al., 2013). When erty of intelligence (Stanovich, 2009).
the brains of children were measured (Figure B after C. B. Malpas et al.,
at age 6 and again at 11, those with 2016. J Clin Neurosci 24: 128–134.)
the highest IQs displayed the great-
(B)
Connectivity (fractional anisotropy)
0.56
0.54
0.50
0.46
0.42
80 90 100 110 120 130 140
IQ score
(B) Correlation between IQ and brain connectivity.
Modern behavioral neuroscience arose in the twentieth century
The end of the nineteenth century brought many important developments for be-
havioral neuroscience. German psychologist Hermann Ebbinghaus showed in 1885
how to measure learning and memory in humans. In 1898, American psychologist
Edward L. Thorndike demonstrated how to measure learning and memory in ani-
mals. Early in the twentieth century, Russian physiologist Ivan P. Pavlov announced
research in his laboratory on conditioning in animals.
American psychologist Shepard I. Franz (1902) sought the site of learning and
memory in the brain by removing different brain regions in animal subjects. This
work started a search for the traces of experience in the brain—a quest that Karl S.
Lashley (1890–1958) referred to as the “search for the engram.”
Behavioral neuroscience bears the strong imprint of Canadian psychologist Don-
ald O. Hebb (1904–1985), a student of Lashley (P. M. Milner, 1993). In his book The
Organization of Behavior (1949), Hebb showed, in principle, how complex cognitive
behavior could be accomplished by networks of active neurons. He suggested how
brain cell connections that are initially more or less random could become orga-
nized, by sensory input and stimulation, into strongly interconnected groups that
he called cell assemblies. His hypothesis about how neurons strengthen their con-
nections through use gave rise to the concept of the Hebbian synapse, a topic much
studied by current neuroscientists (see Chapters 7 and 17).

Consciousness is a thorny problem

Almost anyone using this book has at some time wondered about consciousness: the
personal, private awareness of our emotions, intentions, thoughts, and movements
and of the sensations that impinge upon us. How is it possible that you are aware of
the words on this page, the room you are occupying, the goals you have in life?
In his review of theories of consciousness, Adam Zeman (2002) notes that al-
most all scientists agree on some aspects of consciousness:
• Consciousness matters; it permits us to do certain important things, like
planning and mentally “simulating” what might happen in the future.
• Consciousness is bound up somehow with the activity of the brain. consciousness The state of awareness
• We are not aware of all of our brain’s activities. Some brain activity, and therefore of one’s own existence and experience.
some of our behavior, is unconscious.
• The deepest parts of our brain are important for arousal.
• The topmost parts of the brain are responsible for whatever we experience from
moment to moment.
In the chapters to come, we will see many examples of experiments that demon-
strate these properties of consciousness. However consciousness is brought about,
any satisfying understanding would be able, for example, to explain why a certain
pattern of activity in your brain causes you to experience the sensation of blue when
looking at the sky (FIGURE 1.15), or the smell of cinnamon when entering a bakery.
A good theory would let us predict that after messing about with your brain, chang-
ing particular connections or activating particular neurons, you would experience
yellow when seeing the sky.
Unfortunately, we are nowhere near understanding consciousness this clearly. We
describe some intriguing (and disturbing) experiments explicitly directed at human
consciousness in Chapter 18. In the rest of the book, we rarely use the words conscious
or consciousness. Normally we cannot say anything about the particulars of what hu-
mans or animal subjects are experiencing, but only whether their behavior suggests
© iStock.com/Roman Sigaev

that the brain detected a signal or event. Thus, we are in no position to know whether
complicated machines like androids are, or might one day be, conscious.

1.15 How Blue IS the Sky? We would all agree that this sky is
the color everyone calls “blue.” But in Chapter 18 we will ask whether
everyone who sees that sky has the same experience of the color.

Intro ductio n 19
 1017 1.16 The Human Brain Project The goal of massive
2023 projects in both the United States and Europe is to
1016 Complete human have a digital re-creation of the neurons and connec-
brain (100 billion
Petabyte

1015 tions found in a human brain. As this projection from

neurons)
2011
the European project (bluebrain.epfl.ch) shows, this
Computer memory (bytes)

1014 100 columns of would require a computer that is vastly faster than any
cortex (1 million made to date, as well as a truly staggering amount of
1013
computer memory. (After M. M. Waldrop, 2012. Nature
Terabyte

neurons)
1012 2014 482: 456–458.)
Complete rat
1011 brain (100
million neurons)
1010 2005 Both the United States and Europe have begun projects
Gigabyte

Single 2008
109 neuron Single half-millimeter hoping to map an entire human brain, a truly formidable task
model column of cerebral (Waldrop, 2012), which we discuss in Chapter 2. It will re-
108 cortex (10,000 neurons)
quire a computer with vastly more memory and faster pro-
107 cessing than any yet devised (FIGURE 1.16). Some people
Megabyte

106 even doubt whether our “merely human” brains will ever be
109 1010 1011 1012 1013 1014 1015 1016 1017 1018 able to understand something as complicated as conscious-
Gigaflop Teraflop Petaflop Exaflop ness (Yong, 2019). Nevertheless, any gains we make in un-
Computing speed (flops) derstanding how the brain works, which is the subject of
this book, will bring us closer to that goal.

The Cutting Edge

Behavioral Neuroscience Is Advancing
at a Tremendous Rate
It is difficult to convey the rate at which we are learning new things about the brain.
Each year, over 25,000 neuroscientists meet at the annual meeting of the Society for
Neuroscience (sfn.org). On the website (bn9e.com), the NewsLink tab directs you to
Behavioral Neuroscience 9E news stories about behavioral neuroscience for a general audience, with more than
Breedlove 20 articles added every week, over a thousand per year. The predominant index for
Sinauer Associates biomedical research, PubMed (pubmed.gov), classified over 40,000 articles under
Breedlove9e_1.16.ai Date 07-15-19 neuroscience in 2018 alone (FIGURE 1.17). Somehow, we didn’t quite get around to
reading them all. This incredible pace of research is driven, in part, by talented young
scientists who are more excited by neuroscience than competing fields. Excitement

45,000

40,000
Incidence of terms in biomedical journals

35,000
Neuroscience
30,000

The number of scientific

25,000 articles indexed by these
terms in PubMed.gov
1.17 Neuroscience on the Rise 20,000 keeps growing.
In the major index of biomedical journals,
PubMed (pubmed.gov), occurrence of
15,000
the term neuroscience was rather rare in Neural
the first 20 years covered by the index.
However, use of the term has increased 10,000
steadily since 1990 and risen very
sharply in the past decade. It seems safe 5,000
to say that no one has read, or ever will
read, the 40,451 such articles published 0
in 2018 alone. (www.pubmed.gov.Ac- 1950 1960 1970 1980 1990 2000 2010 2020
cessed July 2019.) Year

20 C HA P T E R 1
about understanding the brain is also the reason that undergraduate majors in neuro-
science are now being offered in dozens of colleges and universities around the United
States and the world.
Given this explosion of information in behavioral neuroscience, it is difficult for
us to keep this book up to date. We’ve done our best in every chapter to convey
those exciting new concepts that seem to be holding up to the scrutiny of the field,
citing over 500 new articles to keep the text current. Each chapter includes a special
feature: The Cutting Edge. Here we present exciting new findings about the area
under discussion. These are the types of findings that have scientists in the field
buzzing among themselves. In addition to highlighting new and exciting ideas, we
use The Cutting Edge to describe experimental approaches in more detail, to give
you a better feel for the process of scientific reasoning and hypothesis testing. We
really enjoy writing these breaking news stories about behavioral neuroscience, and
we hope that they excite you too. ■

Recommended Reading
Blackmore, S., and Troscianko, E. T. (2018). Consciousness: An Introduction (3rd ed.).
London: Routledge.
Carter, R. (2009). The Human Brain Book. London: Dorling Kindersley.
Dehaene, S. (2014). Consciousness and the Brain: Deciphering How the Brain Codes Our
Thoughts. New York: Penguin Books.
Godfrey-Smith, P. (2017). Other Minds: The Octopus, the Sea, and the Deep Origins of
Consciousness. New York: Farrar, Straus and Giroux.
Kaku, M. (2015). The Future of the Mind: The Scientific Quest to Understand, Enhance, and
Empower the Mind. New York: Random House.
Koch, C. (2012). Consciousness: Confessions of a Romantic Reductionist. Cambridge, MA:
MIT Press.
Wickens, A. P. (2014). A History of the Brain: From Stone Age Surgery to Modern Neuroscience.
New York: Psychology Press.
Zimmer, C. (2004). The Soul Made Flesh: The Discovery of the Brain—and How It Changed the
World. New York: Basic Books.

In tro ductio n 21
 1 Visual Summary bn9e.com/vs1
You should be able to relate each summary to the adjacent illustration, including structures and processes.
If you go to the website for this text (bn9e.com), you can follow links to figures, animations, and activities
that will help you consolidate the material.

Cognitive
science
2 Behavioral neurosci-
1 Behavioral neuroscience entists balance three
Computer
Anthropology
science

Cognitive

is a branch of neuroscience general research per-

psychology
Evolutionary Sociobiology Artificial Psychiatry
biology
Cognitive
intelligence
Somatic
that focuses on the biologi-
Behavioral neuroscience
intervention
spectives—correlation,
Behavioral
ecology/ethology Social Neural medicine
neuroscience modeling
Comparative/ Health

cal bases of behavior. It is

evolutionary psychology

somatic intervention,
Paleontology Neurology
Paleoneuro- psychology Clinical
Cognitive neuro-
anatomy Comparative
neuro- psychology
neuroanatomy BEHAVIORAL psychology
Somatic variables Correlations Behavioral variables
closely related to many and behavioral inter-
Neural NEUROSCIENCE Neuro-
Neuro- imaging physiology Electro-
anatomy physiology
Anatomy Developmental Psycho- Physiology

other neuroscience disci- vention—in designing

psychobiology pharmacology
Behavior Psychoneuro-
Developmental
neurobiology
genetics immunology
Behavioral
Pharmacology Behavioral
plines. Review Figure 1.2 intervention
their research. Review
endocrinology
Developmental Genetics/ Neuro- Biochemistry
biology epigenetics immunology
Neuro-

Figure 1.3
endocrinology

Molecular Immunology
biology
Endocrinology

Behavioral Neuroscience 9E
3 Research in behavioral Breedlove Behavioral Neuroscience 9E
Sinauer Associates Dragonfly Media Group Breedlove Other
neuroscience is conduct- dementias
Sinauer2,200,000
Associates Dragonfly
Depression
Media Group
Breed9e_VS_01.01.ai
Alzheimer’s
ed at levels of analysis Date 07-18-19
5,300,000
268,000,000
Epilepsy
Alcohol use 4 The prevalence of neu-
Breed9e_VS_01.02.ai
2,800,000
ranging from molecular 109,000,000

Drug use
Date 07-18-19
rological and psychiatric
events to the functioning Stroke
6,800,000
Anxiety
disorders
73,000,000
disorders exacts a very
289,000,000
of the entire brain and high emotional and
complex social situa- Traumatic
brain injury
1,700,000 economic toll. Review
tions. Review Figure 1.6 Spinal cord MS Parkinson’s Eating Schizophrenia Bipolar Figure 1.8
injury 600,000 disease disorders 19,500,000 disorder
340,000 1,000,000 16,000,000 47,000,000

Behavioral Neuroscience 9E Behavioral Neuroscience 9E

Breedlove Breedlove
Sinauer Associates Dragonfly Media Group
Sinauer Associates Dragonfly Media Group
5 Although genes can have Breed9e_VS_01.03.ai Date 07-18-19 6
Although humans have
a major impact on brain Breed9e_VS_01.04.ai Date 07-26-19
wondered about the control
function, it is clear that of behavior for thousands of
experience physically years, only comparatively
alters the brain and that recently has a mechanistic
genetically identical people view of the brain taken
will not necessarily have hold. Review Figure 1.13
the same brain disorders.
Review Figure 1.9

7 The concept of localization of
function, which originated in
phrenology—despite obvious
flaws with the phrenologists’
methodology—was an impor-
tant milestone for behavioral
neuroscience. Today we know
that the part of the brain that
shows a peak of activity varies
in a predictable way depend-
ing on what task we’re doing.
Review Figure 1.14
Behavioral Neuroscience 9E
Breedlove
Sinauer Associates Dragonfly Media GroupSinauer Associates Dragonfly Media Group
Breed9e_VS_01.05.ai
Behavioral Neuroscience 9E
Breedlove
Sinauer Associates Dragonfly Media Group
Behavioral Neuroscience 9E
Breedlove
Localization of cognitive func- 8
Date Breed9e_VS_01.06.ai
07-18-19 Date 07-18-19
tions remains a major focus of
behavioral neuroscience. With
modern imaging technology
and a more carefully validated
understanding of cognitive
abilities, a detailed view of the
organization of the brain is
emerging. Review Figure 1.14
Behavioral Neuroscience 9E
Breedlove
Sinauer Associates Dragonfly Media Group
Breed9e_VS_01.08.ai Date 07-18-19

Breed9e_VS_01.07.ai Date 07-18-19

 Biological
Foundations
of Behavior

Courtesy of Dr. Ryo Egawa

PART I
CHAPTER 2
Functional Neuroanatomy:
The Cells and the
Structures of the Nervous
System

CHAPTER 3
Neurophysiology:
The Generation,
Transmission, and
Integration of
Neural Signals

CHAPTER 4
The Chemistry
of Behavior:
Neurotransmitters
and Neuropharmacology

CHAPTER 5
Hormones and
the Brain
Another random document with
no related content on Scribd:
mistake. He has not troubled himself to seek in his royal features for
something by which he might be distinguished from the people about
him. Winged genii, king and viziers, all have the same eye, the same
nose and the same mouth. One would say that for each group of
bas-reliefs the original designer only drew one head, which was
repeated by tracing or some other process as often as there might
be heads in the composition, and that it was afterwards carved and
modelled in the alabaster by the chisel of the journeyman.
No, in spite of all that has been said, the Assyrians made no
portraits. They did not even attempt to mark in any precise fashion,
those physical characteristics by which they themselves were so
sharply divided from many of the races by whom they were
surrounded. Among the numerous peoples that figure in the sieges
and battles that cover the palace walls, although some, like the
Chaldæans, the Jews, and the Syrians, were near relations of their
own, others belonged either to the Aryan or Turanian family; but any
one who will examine the reliefs as we have done, will see that all
the prisoners of war and other vanquished enemies have the same
features as their conquerors.[162] The only exception to which we
can point is in the case of certain bas-reliefs of Assurbanipal in which
the episodes of an expedition into Susiana are retraced. There we
can perceive in some of the figures—by no means in all—an
endeavour on the part of the sculptor to mark the difference of race
otherwise than by details of costume and head-dress. Here and
there we find a head that suggests a negro;[163] but his
characteristics are never as clearly marked as in Egypt. This may be
merely the result of caprice on the part of some individual artist who
has amused himself by reproducing with the edge of the chisel some
head which had struck his fancy; but even here we only find one
profile several times repeated. The modelling is far from searching,
but wherever the work is in fair condition and the scale not too small
the character we have described may be easily distinguished. The
only differences over which the Assyrian sculptors naturally troubled
themselves were those of costume and equipment; thus we find
them recording that the people subdued in one of the expeditions of
Sennacherib wore a crown or wreath of feathers about their heads
(Fig. 48).[164] So, too, in the relief of a man with apes, the foot-
covering, a kind of buskin with upturned toes (Fig. 64), should be
noticed. But the lines of his profile remain unchanged; and yet there
can be no doubt that the sculptor here meant to represent a man of
negro race, because, as Layard, who dug up the monument, tells us,
traces of black paint might be distinctly perceived upon the faces of
this man and his companions.[165] On a Babylonian stele that we
have already figured (Fig. 43), some have attempted to recognize a
Mongol type, and thence to confirm the hypothesis that would make
a Turanian race the founders of the Chaldæan civilization. This, too,
we think a mistake.[166]
At first sight this curious monument surprises those who are
accustomed to Assyrian art, but the nature of the material has not a
little to do with that. The hardness and darkness of basalt affect the
treatment of the sculptor in quite a different way from a gypseous
stone like alabaster. Add to this that the proportions are quite unlike
those of the Ninevite reliefs. This Marduk-idin-akhi is a work of the
ancient school, which made its figures far shorter than those of such
Assyrian reliefs as have come down to us. Finally the head-dress
should be noticed. In place of being conical it is cylindrical, a form
which overweights the figure and shortens its apparent proportions.
On the whole, any one looking at this stele without bias on one side
or the other, will, we think, acknowledge that the type it presents is
the same as the figures at Nimroud, Khorsabad, and Kouyundjik. It
is, moreover, identical with that we see in monuments even older
than this royal Babylonian stele, such as the fragmentary relief found
by M. de Sarzec at Sirtella (Fig. 67).
 Fig. 67.—Fragment of a Chaldæan bas-relief. Louvre.
Limestone. Height 3¾ inches.
The type which crops up so often in the pages of this history was
fixed, in all its main features, in the earliest attempts at plastic art
made by the Chaldæans. By them it was transmitted to their
scholars, the Assyrians, and during long centuries, until the fall of
Nineveh and Babylon, the painters and sculptors of Mesopotamia,
from the shores of the Persian Gulf to the foot of the mountains of
Armenia, did not cease to reproduce and perpetuate it, I might say to
satiety; they reproduced it with infinite patience, and, so far as we
can see, without once suspecting that the human visage might
sometimes vary its lines and present another aspect.

§ 4. On the Representations of Animals.

In the preceding pages our chief aim has been to determine the
nature and the mode of action of the influences under which the
Assyro-Chaldæan sculptor had to do his work. We have explained
how certain conditions hampered his progress and in some respects
arrested the development of his skill.
The height to which the plastic genius of this people might have
carried their art had their social habits been more favourable to the
study of the nude, may perhaps be better judged from their treatment
of animals than anything else. Some of these, both in relief and in
the round, are far superior to their human figures, and even now
excite the admiration of sculptors.
The cause of this difference is easily seen. When an artist had to
represent an animal, his study of its form was not embarrassed by
any such obstacle as a long and heavy robe. The animal could be
watched in its naked simplicity and all its instinctive and
characteristic movements grasped. The sculptor could follow each
contour of his model; he could take account of the way in which the
limbs were attached to the trunk; he saw the muscles swell beneath
the skin, he saw them tighten with exertion and relax when at rest.
He was not indifferent to such a sight; on the contrary, he eagerly
drank in the instruction it afforded, and of all the works he produced
those in which such knowledge is put into action are by far the most
perfect; they show us better than anything else how great were his
native gifts, and what a fund of sympathy with the beauties of life and
with its inexhaustible variety his nature contained. Whether he model
an animal separately or introduce it into some historic scene, it is
always well rendered both in form and movement.
This is to be most clearly seen in the rich and varied series of
Assyrian reliefs, but the less numerous works of the same kind of
Babylonian origin show the same tendency and at least equal talent.
In copying the principal types of the animal world with fidelity and
vigour, the Assyrian sculptors only followed the example set them by
their south-country masters.
 Fig. 68.—Head of a cow, bronze.
British Museum. Width across the
cheeks 3¾ inches.
A cow’s head in bronze, which was brought from Bagdad by Mr.
Rassam, is broad in treatment and of great truth (Fig. 68); the same
good qualities are to be found in a terra-cotta tablet found by Sir
Henry Rawlinson in the course of his excavations in the Birs-
Nimroud (Fig. 69). It represents a man, semi-nude and beardless
and with a stout stick in his hand, leading a large and powerfully
made dog by a plaited strap. It is a sort of mastiff that might be used
for hunting the wild beasts in the desert and marshes, the wild boar,
hyena, and panther, if not the lion. The characteristics of the species
are so well marked that naturalists have believed themselves able to
recognise it as that of a dog which is still extant, not in Mesopotamia
indeed, but in Central Asia.[167] We may seek in it for the portrait of
one of those Indian hounds kept, in the time of Herodotus, by the
Satrap of Babylon. His pack was so numerous that it took the
revenues of four large villages to support it.[168]
Similar subjects were represented upon other tablets of the same
origin. One of them shows a lion about to devour a bull and disturbed
by a man brandishing a mace. Nothing could be more faithful than
the action of the animal; without letting go his prey he raises a paw,
its claws opened and extended and ready to be buried in the side of
the rash person who interrupts his meal.[169]

Fig. 69.—Terra-cotta tablet. British Museum. Height

3⅗ inches.
We may also mention a cylinder which, from its style, M. Ménant
does not hesitate to ascribe to the first Chaldæan monarchy. It
represents two oxen in a field of wheat. The latter, by a convention
that also found favour with the Greeks, is indicated by two of those
huge ears that so greatly astonished Herodotus.[170] Was it on a
similar principle that the Chaldæan engraver gave his oxen but one
horn apiece? In spite of this singularity and the peculiar difficulties
offered by work in intaglio on a very hard material, the forms are well
understood, and the artist has not been content to give them merely
in outline. At the croup and under the belly an effort has been made
to model the figure and to mark its thickness.
Judging from their style and inscriptions, several more of these
engraved stones may be ascribed to the oldest Chaldæan schools of
art, but we are satisfied with again reminding our readers that it was
in Lower Mesopotamia that everything had its beginning. We shall
take our remaining examples from the richer deposits of Assyria.

Fig. 70.—Cylinder of black marble. National Library, Paris.

Among all those animals that attracted the attention of man either
by their size or strength, either by the services they rendered or the
terror they caused, there were none that the chisel of the Assyrian
sculptor did not treat and treat with taste and skill. With their passion
for the chase the kings and nobles of Assyria were sure to love dogs
and to train them with scrupulous care.[171] They did more. They
employed sculptors in making portraits of them. In the palace of
Assurbanipal terra-cotta statuettes of his best dogs have been found
(Fig. 71). They belong to the same race as the Chaldæan mastiff
above mentioned, but their strength, their fire, I might almost say
their ferocity, is better shown in those pictures where they are no
longer in a state of repose, but in movement and action. Look at the
series of slabs representing the departure for the chase. The hounds
are held in the leash by attendants who carry bags on their
shoulders for the smaller game (see Fig. 72). Mark the tightened
cord, the straining bodies, the tension of every muscle in their desire
to get at their quarry! We can almost fancy we hear the deep,
confused bayings with which they prelude the regular music of the
hunt itself when the game is afoot. These animals are represented
with no less truth and vivacity when a kill has taken, or is about to
take, place. As an example of this we may point out a relief from the
same palace in which two of these bloodhounds launch themselves
upon a wild ass whose flight has been arrested by an arrow. The ass
still manages to stagger along, but he will not go far; the hounds are
already upon him and have buried their teeth in his flanks and croup.
[172]

Fig. 71.—Terra-cotta dog. British Museum. Height

2⅖ inches.
Other domestic animals are figured with no less sure a hand; to
each is given the proportions and attitudes that really characterise it.
We shall now study them all in succession; others have done so, and
have found much precious information upon the fauna of Western
Asia and upon the state of Mesopotamian civilization;[173] we shall
content ourselves with mentioning the principal types and those in
which the sculptor has shown most skill.
Fig. 72.—The hounds of Assurbanipal. British Museum. Height 26 inches. Drawn
by Saint-Elme Gautier.

The colossi of the gateways have already given us an opportunity

for showing how art enlisted the powerful limbs and natural majesty
of the bull in its service. Elsewhere the bovine race occupies a less
important part in Assyrian sculpture than in that of Egypt, in whose
tombs scenes of agricultural art are of such constant occurrence. We
find, however, the wild bull,[174] which the kings of Calah hunted in
the neighbouring desert (Fig. 15), and the draught ox, which, after a
lucky raid, the terrors of Asia drive before them with their prisoners
and other booty (Vol. I. Fig. 30).[175]—We may also point to the
heifer’s head in ivory which acts as tail-piece to the third chapter of
our first volume. We sometimes find also sheep and goats of both
sexes (Fig. 54);[176] but of all the animals that have close relations
with man, that which occurs most often on the palace walls is the
horse. They did not use him as a beast of burden; it was the mule
that was used for drawing carts (Vol. I. Fig. 31), for carrying women
and children and merchandise (Vol. I. Figs. 30 and 115). As with the
Arabs of to-day, the horse was reserved for war and hunting. But the
Assyrians were not, like the Egyptians, content to harness him to the
chariot; they rode him as well. Their armies comprised a numerous
and well-provided cavalry; and the Assyrian artist drew the horse a
great deal better than his Egyptian confrère.
The horses we meet with in the Assyrian sculptures are of a
heavier breed than Arabs; they are generally shorter and more
thickly set. Travellers believe the breed to still exist in the horses of
Kurdistan, a country which was bordered by ancient Assyria and
dependent upon it.[177] The head is small, well-formed, and well-
carried (Fig. 73), the shoulders sloping, the neck and limbs well set
on, and the muscles strongly marked. We have already had
occasion to figure horses at full speed (Vol. I. Fig. 5), standing still
(Vol. I. Figs. 67 and 115), and proceeding at a slow pace (Figs. 21
and 31).[178] No observer can avoid being struck by the truth of
attitude, and movement given by the Assyrian sculptor to horses
both driven and mounted. Nowhere is this merit more conspicuous
than in one of those bas-reliefs of Assurbanipal that figure the
episodes of a chase of wild asses (Fig. 74).
 Fig. 73.—Chariot horses; from Layard.
Contrary to their usual habits the herd have allowed themselves
to be surprised. One of those armies of beaters who are yet
employed by eastern sovereigns on such occasions, has driven
them upon the hunters. The latter, preceded by their dogs, throw
themselves upon the herd, which breaks up in all directions. They
pierce those that are within reach with their arrows; those that do not
fall at once are pursued and brought down by the hounds. We
cannot reproduce the whole scene,[179] but we doubt whether there
is any school of animal painters that has produced anything more
true to nature than the action of this poor beast stopping in the
middle of his flight to launch futile kicks at his pursuers.
Fig. 74.—Wild ass. From the hunt of Assurbanipal, in the British Museum. Drawn
by Saint-Elme Gautier.
The ibex and the wild goat figure in the same sculptured pictures.
One marching in front of the herd turns and anxiously sniffs the wind,
while her companion quietly browses by her side; farther off, two kids
trot by the side of their mother. The alarm has not yet been given,
but upon the next slab the artist shows the headlong flight that
follows the discovery of the enemy. Naturally it is the wild and
domestic animals of Mesopotamia and the districts about that are
most commonly figured in these reliefs, but the sculptor also took
advantage of every opportunity and pretext for introducing into his
repertory those rare and curious animals which were only seen in
Nineveh on rare occasions. Thus the camel that we find in so many
pictures is the same as that which now occupies the same region
and marches in its slow caravans;[180] but on the obelisk of
Shalmaneser we find the double-humped Bactrian camel (Fig. 49).
[181] The clumsy tribe of the pachyderms is not only represented by
the wild boars that still have their lairs in the marshes of the lower
Euphrates;[182] the rhinoceros and the Indian elephant also occur on
the obelisk (Vol. I. Fig. 111).[183] The apes shown in our Fig. 64 also
seem to belong to an Indian species.[184]
The sculptor was not always as happily inspired by these exotic
animals as by those of his own country, and in that there is nothing
surprising. He only caught a passing glimpse of them as they defiled,
perhaps, before the people in some triumphal procession. On the
other hand, the fauna of his native land were known to him through
long habit, and yet his reproductions of the elephant and the
dromedary are very good, much better than those of the semi-human
ape. His idea of the rhinoceros is very faulty; the single horn planted
on the nose leaves no doubt as to his meaning, but the lion’s mane
with which the animal’s back is clothed has never belonged to the
rhinoceros. The artist may have worked from a description.
In these pictures birds hold a very secondary place; Assyrian
sculpture was hardly light enough of hand to render their forms and
feathers. For such a task, indeed, painting with its varied handling,
its delicate lines and brilliant colours is required. It was with the
brush that the Egyptians succeeded, in the frescoes of their tombs,
in figuring the principal birds of the Nile Valley with all their elegance
of form and brilliant variety of plumage. In Assyria, among a nation of
soldiers and in an art whose chief inspiration had to do with war, the
only bird we find often reproduced is the eagle, the symbol of victory,
who floats over the chariot of the king, and the vulture who devoured
where they fell the bodies of the enemies of Assyria; and even these
images are rather careless and conventional, which may perhaps be
accounted for by their partially symbolic character and their frequent
repetition.[185] A group of partridges rising and, in those sculptures of
the later Sargonids in which the artists show a love for picturesque
detail, birds hopping in the trees or watching over their nestlings,
have been mentioned as showing technical excellence of the same
kind as the hunting scenes.[186] The ostrich appears on the
elaborate decorations of the royal robes (Fig. 75) and upon the
cylinders (Fig. 76). Perhaps it was considered sacred.
 Fig. 75.—Embroidery on the king’s robe; from Layard.

Fig. 76.—Fight between a man and an ostrich.

Chalcedony. National Library, Paris.
As for fishes, crabs, and shells, these were scattered broadcast
over the watercourses in the reliefs, but they are never studied with
any great care (see Vol. I., Figs. 34 and 157), nor is any attempt
made to distinguish their species. They seem to have been
introduced merely as hints to the spectator, to dispel any doubt he
may entertain as to the meaning of those sinuous lines by which the
sculptor suggested rivers and the sea. Where these indications are
not given we might indeed very easily mistake the artist’s intention
(see Vol. I., Figs. 38 and 71).
 Some of the animals in the Assyrian reliefs are then nothing but
determinative signs, a kind of pictorial gloss. Of these it will suffice to
mention the existence. Their forms are so much generalized that
they offer no matter for study. On the other hand, our best attention
should be given to those figures whose modelling has strongly
interested the artist, who has taken a lively pleasure in reproducing
their various aspects and in making them live again in all the
originality of their powerful and exceptional natures. In this respect
the lion deserves particular notice. He interested the Assyrian
sculptors more profoundly than any other animal and they devoted
extraordinary attention to illustrating his various attitudes and
characteristics. One is inclined to believe that the more skilful among
them chose a lion for treatment when they wished to display all the
talent they possessed and to gain a reputation for complete mastery
of their art.[187]
Here we find the great beast stretched carelessly upon the
ground, full of confidence in his strength and careless of danger
(Plate XI.); there he rises to his feet and advances ready to collect
himself and spring upon any threatening enemy or passing prey
(Plate VIII.). We sometimes find both these motives united, as in a
bas-relief of Assurbanipal, which is unfortunately mutilated (Fig. 77).
Here a lioness is stretched upon the ground, her head upon her
forepaws and her tail outstretched behind her, in a favourite attitude
of very young cats. The lion stands upright before her in a proud,
extended attitude like that of the colossal lion from Nimroud (Plate
VIII.); his head and the hind parts of his body are unfortunately
missing.
 BRONZE LION
FROM KHORSABAD
Louvre

Elsewhere we find the lion cautiously emerging from a stoutly-

built timber cage (Fig. 78). He has been captured in a net or snare
and shut up in this narrow prison until the day of some great hunt.
[188] When that arrives the door is raised at a given signal by a man
perched on the top of the cage and protected by a timber grating. In
spite of this defence the service would hardly be free from danger
but that the lion is too pleased to find himself at liberty to look behind
him.[189]
 Fig. 77.—Lion and lioness in a park. British Museum.

The lion finds himself confronted by the Royal huntsman who

fights, as a rule, from his chariot, where two or three companions,
chosen from his bravest and most skilful servants, are ready to lend
him help if necessary. The British Museum possesses a great
number of sculptured pictures in which every incident of the hunt is
figured up to its inevitable end. We reproduce two figures from the
slabs representing the great hunt of Assurbanipal. The first shows a
huge lion mortally wounded by an arrow which still stands in his
body. It has transfixed some great vessel, and the blood gushes in a
wide torrent from his open mouth. Already the chills of death are
upon him and yet with his back arched, and his feet brought together
and grasping the soil, he collects his energies in a last effort to
prevent himself rolling over helplessly on the sand.
Fig. 78.—Lion coming out of his cage. Height of relief about 22 inches. British
Museum.
Drawn by Saint-Elme Gautier.
 Fig. 79.—Wounded lion. Height of slab about 22 inches. British Museum. Drawn
by Saint-Elme Gautier.
Still more expressive, perhaps, and more pathetic, is the picture
of a lioness struck down by the same hand, but in a different fashion
(Fig. 80). One of three arrows that have reached her has transfixed
the spinal column at the loins. All the hinder part of the body is
paralysed. The hind feet drag helplessly on the ground, while the
poor animal still manages for a moment to support herself on her
fore paws. She still faces the enemy, her half opened jaws are at
once agonised and menacing, and, as we gaze upon her, we can
almost fancy that we hear her last groan issue from her dying lips.
We might multiply these examples if we chose, but the two
fragments we have reproduced will, we hope, send our readers to
the British Museum to see the Hunt of Assurbanipal for themselves.
In any case they are enough to prove that the Assyrian sculptor
studied the lion from nature. He was not without opportunities. He
was, no doubt, allowed to assist at those great hunts of which he
was to be the official chronicler. He there saw the king of beasts
throw himself on the spears of the footmen or fly before the arrows of
the charioteers, and break the converging line of beaters; he saw
him fall under his repeated wounds and struggle in his last
convulsions. Later on he could supplement his recollections, he
could complete and correct his sketches by the examination of the
victims.[190] At the end of the day the “bag” was displayed as it is
now at the end of a modern battue, when the keepers bring
pheasants, hares and rabbits, and lay them in long rows in some
clearing or corner of the covert. In one of the Kouyundjik reliefs we
see the king standing before an altar and doing his homage to the
gods after the emotions and dangers of a hunt that was almost a
battle.[191] He seems to pour the wine of the libation upon four dead
lions, which his attendants have arranged in line upon the ground.
There must also have been tame lions in the palaces and royal
parks. Even now they are often to be met with in that country, under
the tent of the Arab chief or in the house of the bey or pacha.[192]
When captured quite young the lion is easily educated, and,
provided that his appetite is never allowed to go unsatisfied, he may
be an inoffensive and almost a docile companion until he is nearly
full grown. We are ready to believe that the lion and lioness shown in
our Fig. 77 were tame ones. The background of the relief suggests a
park attached to the royal residence, rather than a marsh, jungle or
desert. Vines heavy with fruit and bending flowers rise above the
dozing lioness; we can hardly suppose that wild animals could
intrude into such a garden. It follows, then, that the artist could study
his models as they moved at freedom among the trees of the royal
demesne, basking idly in the sun or stretching themselves when they
rose, or burying their gleaming teeth on the living prey thrown to
them by their keepers.
Thanks to such facilities as these the Ninevite sculptors have
handed down to us more faithful reproductions of the lion than their
more skilful successors of Greece or Rome. For the latter the lion
was little more than a conventional type from which ornamental
motives might be drawn. Sometimes no doubt they obtained very
fine effects from it, but they always considered themselves free to
modify and amplify, according to the requirements of the moment.
Thus they were often led to give him full and rounded forms, which
had a beauty of their own but were hardly true to nature. The