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Discovering Behavioral Neuroscience
AN INTRODUCTION TO BIOLOGICAL PSYCHOLOGY
Fourth Edition
Laura A. Freberg
California Polytechnic State University,
San Luis Obispo
Australia ● Brazil ● Mexico ● Singapore ● United Kingdom ● United States
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An Introduction to Biological Psychology,
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iii
TO MY FAMILY
Roger, Kristin, Karen, and Karla
About the Author
Laura A. Freberg is Professor of Psychology at California Polytechnic State University,

San Luis Obispo, where she teaches courses in Introductory Psychology, Biological
Psychology, and Sensation and Perception. With John Cacioppo of the University
of Chicago, Laura is the co-author of three editions of Discovering Psychology: The
Science of Mind for Cengage Learning. She is also lead author on a new online Research
Methods textbook.
Laura completed her undergraduate and graduate studies at UCLA, where her
thinking about psychology was shaped by Eric Holman, John Garcia, O. Ivar Lovaas,
Larry Butcher, Jackson Beatty, John Libeskind, Donald Novin, Frank Krasne, and
F. Nowell Jones. She was privileged to study neuroanatomy with Arnold Scheibel, and
she investigated the effects of psychoactive drugs on learning and memory under the
direction of Murray Jarvik and Ronald Siegel in the UCLA Neuropsychiatric Institute.
As a capstone to her education, Laura completed her dissertation with Robert Rescorla,
then at Yale University.
Laura’s teaching career began when she taught her first college course at Pasadena
City College at the age of 23 while still a graduate student at UCLA. Beginning in
2011, to better understand the needs of the online education community, she also be-
gan teaching for Argosy University Online, including courses in Social Psychology,
Sensation/Perception, Cognitive Psychology, Statistics, Research Methods, and Writing
in Psychology. She has also redesigned her Cal Poly introductory course according
to QOLT standards to be administered completely online. She has received Faculty
Member of the Year recognition from Cal Poly Disabilities Resource Center three
times (1991, 1994, and 2009) for her work with students with disabilities. She enjoys
using technology and social media in the classroom and is a Google Glass Explorer.
Laura enjoys collaborating with daughters Kristin Saling (Systems Engineering—U.S.
Military Academy at West Point) and Karen Freberg (Communications—University
of Louisville) on a variety of research projects in crisis management and public rela-
tions as well as in psychology. She serves as the Bylaws and Archives Committee Chair
for the Society for Social Neuroscience and was recently elected President-Elect of the
Western Psychological Association (WPA).
In her spare time, Laura enjoys family time with her husband, Roger, their young-
est daughter Karla, who has autism spectrum disorder, and an active menagerie in-
cluding an Australian shepherd, two cats, and three parakeets. She usually writes while
consuming vast quantities of Gevalia coffee and listening to the Rolling Stones (which
might be apparent in the book’s writing style), and she has been known to enjoy college
football, Harley Davidsons, episodes of Game of Thrones that do not feature weddings,
and Sherlock. Her ringtone is from Nintendo’s Legend of Zelda.
iv
Brief Contents v
Brief Contents
Preface xix
1 What Is Behavioral Neuroscience? 1
2 Functional Neuroanatomy and the Evolution of the

Nervous System 27
3 Neurophysiology: The Structure and Functions

of the Cells of the Nervous System 67
4 Psychopharmacology 107
5 Genetics and the Development of the Human Brain 145
6 Vision 183
7 Nonvisual Sensation and Perception 225
8 Movement 265
9 Homeostasis, Motivation, and Reward 301
10 Sexual Behavior 337
11 Sleep and Waking 375
12 Learning and Memory 415
13 Cognitive Neuroscience 455
14 Emotion, Aggression, and Stress 489
15 Neuropsychology 523
16 Psychopathology 555
References R-1
Name Index I-1
Subject Index/Glossary I-12
vi About the Cover
About the Cover
If you are familiar with the first three editions of this textbook, you know that we like
to pick colorful visuals that portray the biology behind the behavior. For this fourth
edition, we selected an image of the Organ of Corti in the inner ear produced by Sonja
Pyott of the Univeristy of Groningen using a technique known as confocal micros-
copy. Confocal microscopy produces superior resolution compared to conventional,
wide-field microscopes. This stunning image was awarded fourth place in the annu-
al Olympus BioScapes International Digital Imaging Competition in 2007. The green
structures are the hair cells of the cochlea (see Chapter 7), and the nuclei of the inner
hair cells appear in blue. Neurons making contact with the hair cells appear red.
Photo by Dr. Sonja Pyott, University of North Carolina
vi
Contents
Preface xix
1 What Is Behavioral Neuroscience? 1

Neuroscience as an Interdisciplinary Field 2
Historical Highlights in Neuroscience 5
Ancient Milestones in Understanding the Nervous System 5
The Dawn of Scientific Reasoning 5
Modern Neuroscience Begins 6
Interim Summary 1.1 7
Behavioral Neuroscience Research Methods 9
Microscopic Methods 9
Imaging 11
Recording 14
Brain Stimulation 17
Lesion 19
Biochemical Methods 20
Genetic Methods 20
Research Ethics in Behavioral Neuroscience 23
Human Participant Guidelines 23
Animal Subjects Guidelines 24
Chapter Review • Thought Questions • Key Terms 26
CONNECTING TO RESEARCH: Thinking about Your Food as Healthy or

Indulgent Affects Your Physical Reactions to Eating 3
BEHAVIORAL NEUROSCIENCE GOES TO WORK: What Can I Do with a Degree
in Neuroscience? 4
THINKING ETHICALLY: Can We Read Minds with Brain Imaging? 14
BUILDING BETTER HEALTH: When Is It Appropriate to Use Placebos? 25
2 Functional Neuroanatomy and the Evolution

of the Nervous System 27
Anatomical Directions and Planes of Section 28
Protecting and Supplying the Nervous System 30
Meninges 30
Cerebrospinal Fluid 31
vii
viii Contents
The Brain’s Blood Supply 34

The Central Nervous System 36
The Spinal Cord 37
Embryological Divisions of the Brain 38
The Hindbrain 39
The Midbrain 41
The Forebrain 42
The Peripheral Nervous System 55
The Cranial Nerves 55
The Spinal Nerves 55
The Autonomic Nervous System 57
The Endocrine System 61
The Evolution of the Human Nervous System 61
Natural Selection and Evolution 61
Evolution of the Nervous System 62
Evolution of the Human Brain 63
CONNECTING TO RESEARCH: Linking the Brain and the Immune System 31

BEHAVIORAL NEUROSCIENCE GOES TO WORK: Treating Hydrocephalus 34
BUILDING BETTER HEALTH: Using Epidural Stimulation to Improve Standing and
Walking In Patients with Spinal Damage 39
THINKING ETHICALLY: Can We Localize Intelligence in the Brain? 53
3 Neurophysiology: The Structure and Functions of the

Cells of the Nervous System 67
Glia and Neurons 68
Glia 68
The Structure of Neurons 73
Structural Variations in Neurons 80
Functional Variations in Neurons 82
Generating Action Potentials 83
The Ionic Composition of the Intracellular and Extracellular Fluids 83
The Movement of Ions 85
The Resting Potential 86
The Action Potential 86
Propagating Action Potentials 90
The Synapse 94
Gap Junctions 95
Chemical Synapses 96
Axo-axonic Synapses 103
Contents ix
CONNECTING TO RESEARCH: Astrocytes, HIV, and the Blood–Brain Barrier 70

BUILDING BETTER HEALTH: Microglia and Autism Spectrum Disorder 73
THINKING ETHICALLY: Lethal Injection 87
BEHAVIORAL NEUROSCIENCE GOES TO WORK: What Are We Reading in an ECG or EEG? 92
4 Psychopharmacology 107
Neurotransmitters, Neuromodulators, and Neurohormones 108
Identifying Neurochemicals 109
Types of Neurochemicals 109
Mechanisms of Neuropharmacology 123
Agonists and Antagonists 123
Production of Neurochemicals 123
Neurochemical Storage 123
Neurochemical Release 124
Receptor Effects 125
Reuptake and Enzymatic Degradation 126
Basic Principles of Drug Effects 127
Administration of Drugs 128
Individual Differences in Responses to Drugs 128
Placebo Effects 129
Tolerance and Withdrawal 129
Addiction 130
Effects of Selected Psychoactive Drugs 133
Stimulants 133
Opioids 137
Cannabis 138
LSD 139
Alcohol 141
CONNECTING TO RESEARCH: Otto Loewi and “Vagus Stuff” 110

BEHAVIORAL NEUROSCIENCE GOES TO WORK: Substance Abuse Counselors 132
BUILDING BETTER HEALTH: Does Legalization Change Adolescent Attitudes Toward
and Use of Marijuana? 140
THINKING ETHICALLY: Using Addictive Drugs Affects Future Generations 142
5 Genetics and the Development of the Human Brain 145

The Genetic Bases of Behavior 146
From Genome to Trait 146
Sources of Genetic Variability 148
Heritability 151
Epigenetics 153
x Contents
Building a Brain 158

Prenatal Development 158
Effects of Experience on Development 167
Disorders of Nervous System Development 172
The Brain Across the Lifespan 176
Brain Changes During Adolescence and Adulthood 176
Adult Neurogenesis 178
Healthy Brain Aging 178
CONNECTING TO RESEARCH: Epigenetics, Gene Expression, and Stress 155

BEHAVIORAL NEUROSCIENCE GOES TO WORK: Genetics Counseling 156
THINKING ETHICALLY: When are Adolescents Responsible for Their Actions? 179
BUILDING BETTER HEALTH: Nutritional Cognitive Neuroscience and Healthy
Aging 179
6 Vision 183
From Sensation to Perception 184
The Visual Stimulus: Light 185
The Advantages of Light as a Stimulus 185
The Electromagnetic Spectrum 185
Light Interacts with Objects 186
The Structure and Functions of the Visual System 188
Protecting the Eye 188
The Functional Anatomy of the Eye 188
The Layered Organization of the Retina 192
The Photoreceptors 192
Processing by Retinal Interneurons 196
Optic Nerve Connections 201
The Striate Cortex 204
Visual Analysis beyond the Striate Cortex 207
Visual Perception 211
Hierarchies 211
Spatial Frequencies 212
The Perception of Depth 213
Coding Color 214
The Lifespan Development of the Visual System 218
Disorders of the Visual System 219
Amblyopia 219
Cataracts 220
Visual Acuity Problems 220
Blindness 220
Visual Agnosias 221
Contents xi
CONNECTING TO RESEARCH: Hubel and Wiesel Map the Visual Cortex 206
BEHAVIORAL NEUROSCIENCE GOES TO WORK: 3-D Animation 214
THINKING ETHICALLY: Are There Sex Differences in Color Preferences? 216
BUILDING BETTER HEALTH: Does Eating Carrots Really Help Your Vision? 222
7 Nonvisual Sensation and Perception 225

Audition 226
Sound as a Stimulus 226
The Structure and Function of the Auditory System 229
Auditory Perception 235
Hearing Disorders 237
The Body Senses 241
The Vestibular System 241
Touch 243
Pain 251
The Chemical Senses 257
Olfaction 257
Gustation 259
Synaesthesia 262
BUILDING BETTER HEALTH: Earbuds and Hearing Loss 238

THINKING ETHICALLY: Cochlear Prosthetics and Deaf Culture 239
CONNECTING TO RESEARCH: Phantom Limbs, Mirrors, and Longer Noses 250
BEHAVIORAL NEUROSCIENCE GOES TO WORK: What Is a Perfumer? 262
8 Movement 265
Muscles 266
Types of Muscles 266
Muscle Anatomy and Contraction 267
The Effects of Exercise on Muscle 270
The Effects of Aging on Muscles 270
Neural Control of Muscles 272
Alpha Motor Neurons 272
The Motor Unit 272
The Control of Muscle Contractions 273
The Control of Spinal Motor Neurons 274
Reflex Control of Movement 278
Reciprocal Inhibition at Joints 278
The Flexor Reflex 279
Spinal Reflexes Related to Walking 280
Reflexes over the Lifespan 280
xii Contents
Motor Systems of the Brain 281

Spinal Motor Pathways 281
The Cerebellum 283
The Basal Ganglia 284
The Motor Cortex 285
Disorders of Movement 291
Toxins 291
Myasthenia Gravis 292
Muscular Dystrophy 292
Polio 293
Accidental Spinal Cord Injury (SCI) 293
Amyotrophic Lateral Sclerosis (ALS; Lou Gehrig’s Disease) 294
Parkinson’s Disease 295
Huntington’s Disease 297
THINKING ETHICALLY: Gene Doping for Strength 271

CONNECTING TO RESEARCH: Mirror Neurons 289
BUILDING BETTER HEALTH: When Vaccination Is Not Enough 293
BEHAVIORAL NEUROSCIENCE GOES TO WORK: Physical Therapy 294
9 Homeostasis, Motivation, and Reward 301

Homeostasis and Motivation 302
Regulating Body Temperature 303
Adaptations Maintain Temperature 303
Endothermic Responses to Heat and Cold 303
Deviations in Human Core Temperature 305
Brain Mechanisms for Temperature Regulation 307
Thirst: Regulating the Body’s Fluid Levels 309
Intracellular and Extracellular Fluid Compartments 309
Osmosis Causes Water to Move 310
The Kidneys 310
The Sensation of Thirst 311
Hunger: Regulating the Body’s Supply of Nutrients 318
The Process of Digestion 318
The Pancreatic Hormones 319
The Initiation of Eating 320
Satiety 325
Healthy and Disordered Eating 326
Defining Healthy Weight 326
Obesity 327
Disordered Eating 328
Pleasure and Reward 332
Multiple Facets of Reward 333
Contents xiii
Reward Pathways 333

The Neurochemistry of Reward 334
Cortical Processing of Reward 335
BUILDING BETTER HEALTH: Understanding the Benefits of Fever 306

CONNECTING TO RESEARCH: Swallowing Balloons and Growling Stomachs 321
BEHAVIORAL NEUROSCIENCE GOES TO WORK: Dieticians and Nutritionists 327
THINKING ETHICALLY: How Dangerous Is It to Be Overweight or Obese? 329
10 Sexual Behavior 337

Sexual Development 338
The Genetics of Sex 338
Three Stages of Prenatal Development 342
Development at Mini-puberty 346
Development at Puberty 347
Sex Differences in Hormones, Brain Structure, and Behavior 351
The Organizing Role of Sex Hormones 351
The Organizing Role of Sex Chromosome Genes 353
Sexual Dimorphism in the Brain 354
Sex Differences in Behavior and Cognition 356
Sexual Orientation 359
Hormones and Sexual Orientation 360
Brain Structure and Sexual Orientation 361
Genes and Sexual Orientation 362
Sexual Orientation and Cognition 362
Biological Influences on Adult Sexual Behavior 364
The Regulation of Sex Hormones 364
Mood, Menstruation, and Childbirth 365
Hormones and Adult Sexual Behavior 366
Attraction, Romantic Love, Sexual Desire, and Parenting 367
Elements of Physical Attractiveness 368
Romantic Love and Sexual Desire 369
Reproduction and Parenting 371
Sexual Dysfunction and Its Treatment 372
BEHAVIORAL NEUROSCIENCE GOES TO WORK: How Do Therapists Treat Gender

Dysphoria? 348
CONNECTING TO RESEARCH: Simon LeVay and INAH-3 362
THINKING ETHICALLY: Biology’s Role in Explaining Gender and Sexual Orientation 363
BUILDING BETTER HEALTH: Treating Patients with Antidepressant-Induced Sexual
Dysfunction 373
xiv Contents
11 Sleep and Waking 375

Biorhythms 376
Individual Variations in Sleep Patterns 376
Shift Work, Jet Lag, and Daylight Saving Time 378
The Body’s Internal Clocks Manage Circadian Rhythms 379
Major Depressive Disorder with Seasonal Pattern 385
Neural Correlates of Waking and Sleep 388
Electroencephalogram Recordings of Waking and Sleep 388
Brain Networks Control Waking and Sleep 392
Biochemical Correlates of Waking and Sleep 397
The Functions of Sleep 399
Changes in Sleep over the Lifetime 400
Possible Advantages of Sleep 401
Special Benefits of REM Sleep 403
The Possible Functions of Dreaming 404
Sleep–Wake Disorders 407
Insomnia 407
Narcolepsy 408
Breathing-Related Sleep Disorders 409
Sudden Infant Death Syndrome (SIDS) 410
Sleep Talking and Sleep Walking 411
REM Sleep Behavior Disorder 411
Restless Legs Syndrome (RLS) 411
CONNECTING TO RESEARCH: A Composite Scale of Morningness 377

THINKING ETHICALLY: Artificial Lighting and Circadian Rhythms 386
BUILDING BETTER HEALTH: Do Smartphone Sleep Apps Work? 389
BEHAVIORAL NEUROSCIENCE GOES TO WORK: Sleep Medicine 407
12 Learning and Memory 415

Categorizing Learning and Memory 416
Types of Learning 416
Types of Memory 418
Mechanisms of Synaptic Plasticity 421
Learning in Simple Organisms 421
Long-Term Potentiation (LTP) 425
Working Memory 432
Memory Consolidation 433
Reactivation and Reconsolidation 433
Neural Systems Supporting Learning and Memory 435
Early Efforts to Locate Memory Functions 435
Contents xv
Systems Supporting Classical Conditioning 436

The Temporal Lobe and Episodic Memory 441
Semantic Memory Networks 444
Systems Supporting Working Memory 445
Systems Supporting Procedural Memory 446
The Effects of Stress and Healthy Aging on Learning and Memory 448
Stress Effects on Memory 448
The Effects of Healthy Aging on Memory 451
BEHAVIORAL NEUROSCIENCE GOES TO WORK: What Is Neuro Education? 421

CONNECTING TO RESEARCH: Karl Lashley’s Search for the Engram 436
THINKING ETHICALLY: Should We Erase Traumatic Memories? 451
BUILDING BETTER HEALTH: Can We Avoid Age-Related Memory Deficits? 452
13 Cognitive Neuroscience 455

Understanding Cognitive Neuroscience 456
Hemispheric Asymmetry and Its Behavioral Correlates 456
Learning About Asymmetry 456
The Evolution of Lateralization 459
The Development of Lateralization 460
Implications of Asymmetry for Behavior 461
Language 468
The Origins of Language 468
Communication in Nonhuman Animals 469
Multilingualism 471
American Sign Language (ASL) 472
Communication Disorders and Brain Mechanisms for Language 473
Paul Broca and Patient Tan 473
Aphasia 473
Disorders of Reading and Writing 478
Stuttering 479
Intelligence 481
Assessing Intelligence 481
Intelligence and Genetics 482
Structural and Functional Correlates of Intelligence 482
The Neuroscience of Decision Making 485
CONNECTING TO RESEARCH: Savants and Laterality 466

BEHAVIORAL NEUROSCIENCE GOES TO WORK: Speech and Language Pathology 480
THINKING ETHICALLY: Performance-Enhancing Drugs for the Mind? 484
BUILDING BETTER HEALTH: Enriched Environments, Infectious Load, and IQ 485
xvi Contents
14 Emotion, Aggression, and Stress 489

Emotion 490
The Evolution and Adaptive Benefits of Emotion 490
Theories of Emotion 491
The Expression and Recognition of Emotion 495
Biological Correlates of Emotion 501
Emotion Regulation 506
Social Cognition 507
Aggression and Violence 508
Genetics, Environment, Epigenetics, and Aggression 509
Brain Structures and Aggression 510
Biochemistry and Aggression 511
Stress 513
Hans Selye and the General Adaptation Syndrome (GAS) 514
Responses to Stress 515
Stress and Epigenetics 517
Stress, the Immune System, and Health 518
CONNECTING TO RESEARCH: Facial Expressions Predict Assault 498

THINKING ETHICALLY: Using Neuroscience to Assess “Dangerousness” 510
BEHAVIORAL NEUROSCIENCE GOES TO WORK: Pet Therapy for Stress 519
BUILDING BETTER HEALTH: Stress Management 520
15 Neuropsychology 523
What Is Neuropsychology? 524
Who Are the Neuropsychologists? 524
Neuropsychological Assessment 524
Neurocognitive Disorders 525
Alzheimer’s Disease 526
Vascular Disease (Stroke) 528
Traumatic Brain Injury (TBI) 532
Substance/Medication-Induced Neurocognitive Disorder 534
HIV-Associated Neurocognitive Disorder (HAND) 534
Prion Diseases 535
Neurocognitive Disorders Due to Other Medical Conditions 540
Brain Tumors 540
Infections 543
Epilepsy 545
Multiple Sclerosis 548
Migraine 549
Contents xvii
Recovery and Treatment in Neurocognitive Disorders 550

Plasticity and Recovery 550
Cognitive Reserve 551
Rehabilitation for Neurocognitive Disorders 552
BEHAVIORAL NEUROSCIENCE GOES TO WORK: Preparing to Be a Clinical

Neuropsychologist 525
THINKING ETHICALLY: Apoe ɛ4 in Younger Adults 529
BUILDING BETTER HEALTH: Recognizing the Signs of a Stroke 531
CONNECTING TO RESEARCH: Stanley Prusiner and the Prion 537
16 Psychopathology 555
What Does It Mean to Have a Mental Disorder? 556
Autism Spectrum Disorder (ASD) 558
Causes of ASD 559
Brain Structure and Function in ASD 560
Treatment of ASD 562
Attention Deficit Hyperactivity Disorder (ADHD) 563
Causes of ADHD 564
Brain Structure and Function in ADHD 564
Treatment of ADHD 565
Schizophrenia 566
Genetic Contributions to Schizophrenia 567
Environmental Influences on Schizophrenia 568
Brain Structure and Function in Schizophrenia 569
The Biochemistry of Schizophrenia 571
Treating Schizophrenia 572
Bipolar Disorder 574
Genetics and Bipolar Disorder 575
Brain Structure and Function in Bipolar Disorder 575
Biochemistry and Treatment of Bipolar Disorder 576
Major Depressive Disorder (MDD) 577
Genetic Contributions to MDD 577
Environmental Influences on MDD 578
Brain Structure and Function in MDD 578
Biochemistry of MDD 579
Treatment of MDD 580
Anxiety Disorders 582
Obsessive-Compulsive Disorder (OCD) 583
Posttraumatic Stress Disorder (PTSD) 585
Brain Structure and Activity in PTSD 586
Biochemistry and Treatment of PTSD 587
xviii Contents
Antisocial Personality Disorder (ASPD) 587

Genetics and ASPD 587
Brain Structure and Function in ASPD 588
Treatment of ASPD 589
CONNECTING TO RESEARCH: Genetic Overlap in Five Disorders 557

BUILDING BETTER HEALTH: The Gut Microbiota and Mental Disorders 557
BEHAVIORAL NEUROSCIENCE GOES TO WORK: Applied Behavior Analysis 563
THINKING ETHICALLY: Are Psychopaths Responsible for their Behavior? 589
Reference R-1
Name Index I-1
Subject Index/Glossary I-12
Preface
“[I]n teaching, you must simply work your pupil into such a state of interest in
what you are going to teach him that every other object of attention is banished
from his mind; then reveal it to him so impressively that he will remember the
occasion to his dying day; and finally fill him with devouring curiosity to know
what the steps in connection with the subject are.”
—William James (1899, p. 10)
James’s goals for the classroom instructor might seem lofty to some, but many of us
who teach neuroscience have enjoyed the peak experience of seeing students “turn on”
to the material in just the way James describes.
This is an exciting time to be a neuroscientist. Every day, science newsfeeds an-
nounce some new and dramatic breakthroughs in our knowledge about the nervous
system and the human mind. Important questions raised in the past now have de-
finitive answers. In 1890, James commented that “blood very likely may rush to each
region of the cortex according as it is most active, but of this we know nothing” (vol. 1,
p. 99). With today’s technology, it is safe to say we now know much more than “noth-
ing” about this phenomenon James described.
Much has changed in the field of neuroscience since our first edition of this text-
book in 2006. More than half of the four-year universities in the United States now
offer bachelor’s degrees in neuroscience, and most offer at least a minor in the disci-
pline. Neuroscience reflects a general academic trend of the twenty-first century, in
which the walls separating specializations are giving way to new, transdisciplinary
research teams, courses, and educational programs. In recognition of these changes,
we modified the title of the third edition of this textbook from Discovering Biological
Psychology to Discovering Behavioral Neuroscience: An Introduction to Biological
Psychology. This change was welcomed by our faculty and student audiences. A great-
er emphasis on the neurosciences in general is also achieved by renaming some chap-
ter titles. Psychology still provides a foundation for the study of behavioral neuro-
science, as without the ability to ask the right questions about behavior and mental
processes, all of the technology on the planet wouldn’t do us much good. Our current
behavioral neuroscience students, however, are just as likely to be preparing for ca-
reers in the health professions, biomedical engineering, or even scientific journalism
as they are in psychology.
A major reflection of the transdisciplinary approach exemplified by the neurosci-
ences is the inclusion of psychology and behavioral neuroscience content in the revised
edition of the Medical College Admission Test (MCAT) beginning in 2015. One hundred
years ago, the leading killers of humans were infectious diseases. Today’s top killers—
heart disease, diabetes, and cancer—have far stronger relationships with behavior, not
only in their causes but also in their treatments. A simple five-minute conversation
with a health professional about the need to quit smoking is sufficient to lead to absten-
tion for one year by 2 percent of patients (Law & Tang, 1995). This might not sound like
much, but given the 20 percent or so of American adults who smoke and the billions
of dollars their health care and lost productivity represent, the stakes are high. Imagine
what could be accomplished by health care providers who have a deep understanding
of learning, motivation, and social influences on behavior. In response to these and
similar trends, the current edition of the textbook explores relevant applications to
students pursuing fields of study other than psychology whenever these are relevant.
xix
xx Preface
This fourth edition continues and expands on the goals of the previous three:
▶ To provide challenging, current content in a student-friendly, accessible form.
▶ To stimulate critical thinking about neuroscience by presenting controversial and
cutting-edge material.
▶ To promote active student engagement and excitement about the neurosciences.
▶ To integrate across chapters rather than treating them as stand-alone modules; to
encourage students to see the connections among the topics. For example, connec-
tions are made between glutamate as a chemical messenger, its role in learning, the
effects of psychoactive drugs on glutamate, its role in psychosis, and its importance
to the causes and treatments of stroke and epilepsy.
Pedagogical Features
We realize that a course in behavioral neuroscience can be challenging for many students,
particularly those who are underprepared for science courses. To make the process of mas-
tering behavioral neuroscience concepts easier, we have included the following features:
▶ Accessible Writing Style Many textbooks are classified by “level,” but it is my
opinion that the most complex topics can be mastered by students across a wide
range of preparation if the writing style is clear. Students and instructors from the
community college through the top R1 universities have kindly complimented me
on the accessible writing style used in this textbook. The textbook is also widely
adopted in non-English-speaking countries, which suggests that the writing style
is manageable for those for whom English is not a first language.
▶ Clear, Large, Carefully Labeled Illustrations Our medical-quality anatomical il-
lustrations help students visualize the structures and processes discussed in each
part of the textbook. Behavioral neuroscience is similar to geography in its highly
visual nature, and both fields require more visual aids than most other courses. The
illustrations in the textbook are augmented by a set of online animations that help
the student grasp processes over time, such as the propagation of action potentials
down the length of an axon.
▶ Learning Objectives and Chapter Outlines Each chapter begins with a concise set
of learning objectives designed to tap into higher levels of Bloom’s taxonomy, as well
as an outline of the chapter’s content. These features assist students in planning their
learning and in becoming familiar with main terms and concepts to be covered.
▶ Margin Glossaries We regularly provide margin definitions for many difficult
terms. Unlike many textbooks, we do not restrict margin definitions to key terms
only. In the electronic forms of the book, these take the form of pop-up definitions,
with which students are familiar from their online searching experience.
▶ Key Terms We provide a concise list of key terms to help students focus their
learning. Behavioral neuroscience can often seem more like a foreign language
course than a science course, and students benefit from guidance regarding which
terms should be prioritized.
▶ Interim Summaries Each chapter features two to three interim summaries where
students can catch their breath and check their mastery of the material before pro-
ceeding. These summaries feature summary points keyed to the learning objectives
listed at the beginning of each chapter as well as review questions. Most interim
summaries also include a helpful table that pulls together key concepts from the
previous section in one convenient place.
▶ Chapter Integration To emphasize how the material fits together and to promote
elaborative rehearsal, we make references to other chapters relevant to the topic
at hand.
Preface xxi
▶ Chapter Review At the end of each chapter, the student will find some thought
questions that can also serve as essay or discussion prompts. The Chapter Reviews
also include the list of key terms.
▶ Practice Tests In the electronic version of the textbook (see the description of
MindTap), practice tests will be available for each main heading with a compre-
hensive practice test at the end of each chapter.
Additional Features
Students have told me that the narrative of the textbook is “packed” and that skimming
paragraphs is usually a recipe for disaster, as each sentence “counts.” In defense, I re-
spond that we have so much to say and so little room to say it in that there is little space
for “fluff.” At the same time, psychological science shows that spaced learning is supe-
rior to massed learning, so it is a good idea to provide regular breaks in the narrative to
allow students to catch their breath and digest what they have read. We like to think of
these breaks as cool stepping-stones in the flow of lava.
One type of break that we used in the previous editions and continue here in the
fourth is the use of interim summaries that include section summary points and review
questions. Most also feature tables that pull together chunks of material in a way that
makes it easy to learn. Any complex field like the neurosciences entails a bit of simple,
rote memorization to form a foundation for later analysis and critical thinking. The
more quickly we can bring students up to speed on the basics, the faster we can move
on to higher levels of discussion. Chapter summaries include thought questions de-
signed to push students to think more actively and deeply about what they have read.
In addition to the interim and chapter summaries, each chapter includes four types
of features. We recognize that “boxing” material often encourages students to overlook
content unless expressly instructed to read the boxes, but we trust instructors to use
these in ways consistent with their personal style. Obviously, we hope that the content is
sufficiently engaging that students will read the material regardless of “what’s on the test.”
▶ Thinking Ethically features introduce controversial, contemporary questions that
require the students to use the information in the chapter in critical ways. Our stu-
dents will graduate to become community leaders, and they need to be able to think
ethically about future cultural choices related to the neurosciences. For example,
this feature in Chapter 1 follows a discussion of brain imaging technology with
questions about the potential use of brain imaging as a “lie detection” technology.
▶ Connecting to Research features highlight either classic or very contemporary
single studies in behavioral neuroscience. This provides students with a “soft”
segue into the scholarly literature, which might otherwise seem somewhat in-
timidating. The feature emphasizes the type of critical thinking and creativity
required to advance science. For example, this feature in Chapter 2 describes the
recent paradigm-shifting discovery about the links between the brain and the
immune system.
▶ Behavioral Neuroscience Goes to Work features expose students to some of the
many real-world career paths that relate to behavioral neuroscience. In my expe-
rience, many students are unaware of a number of these options. They love the
material but have no idea how they can meld this passion with their need to find
employment. In this feature in Chapter 5, we describe the role of the genetics coun-
selor, whose insights will be increasingly important as the public obtains more in-
formation about personal genotypes. As a bridge between biological sciences and
the counseling professions, this career has become increasingly popular with my
students. At least a dozen who are now enrolled or who have completed genetics
counseling master’s degrees attribute their career choice to my “selling” this con-
cept in class.
xxii Preface
▶ Building Better Health features provide an additional opportunity for students to

think critically about behavioral neuroscience in the context of real-world health
problems. How does your mindset regarding food (healthy versus indulgent) affect
your satiety? Why are microglia important for our understanding of autism spectrum
disorder? Will legalization of marijuana influence attitudes and prevalence of use?
New Content for the Fourth Edition

This new edition contains hundreds of new citations to reflect the advances in the field
that have occurred since the previous edition went to press. Textbook authors are often
challenged by colleagues with questions about “why do you need a new edition?” In
behavioral neuroscience, this question is easy to answer: we have so much new, exciting
research to share.
Illustrations have also been updated to reflect the new content. Because space is
so precious, illustrations are viewed as “teachable moments” that expand on or further
explain the narrative rather than redundant, “pretty” placeholders. We are especially
proud of our medical-quality anatomical illustrations, which have been the source of
much positive feedback through the previous editions.
Space does not permit me to provide an exhaustive list of the updates, but here are
some of the chapter-by-chapter highlights:
Chapter 1 What Is Behavioral Neuroscience?

▶ Expanded discussion of microscopic methods, including flourescent microscopy
▶ Added description of optical imaging
▶ Updated and expanded discussion of optogenetics
▶ Added section on genetic screens
Chapter 2 Functional Neuroanatomy and the Evolution

of the Nervous System
▶ Added discussion of work by Louveau et al. (2015) demonstrating connectivity
between the brain and the immune system
▶ Expanded discussion of the cingulate cortex, amygdala, and periaqueductal gray
▶ Refreshed Connecting to Research feature with a detailed explanation of Louveau
et al. (2015) and connections between the immune system and the brain
▶ Refreshed Building Better Health feature with a discussion of epidural stimulation
and standing/walking in patients with spinal damage
Chapter 3 Neurophysiology: The Structure and Function

of the Cells of the Nervous System
▶ Added section on ependymal cells
▶ Added discussion of exosomes
▶ Updated and expanded discussion of microglia
▶ Expanded discussion of gap junctions
▶ Distinguished between wiring transmission and volume transmission
▶ Refreshed Building Better Health feature with a discussion of microglia and autism
spectrum disorder
Preface xxiii
Chapter 4 Psychopharmacology
▶ Updated discussion of neurotransmitter, neuromodulator, and neurohormone
terms
▶ Updated discussion of cocaine and amphetamine methods of action
▶ Expanded discussion of opioids
▶ Added discussion of hallucinogen persisting perception disorder (HPPD)
▶ Refreshed Building Better Health feature with discussion of the implications of
legalization on attitudes toward and the prevalence of use of marijuana
Chapter 5 Genetics and the Development

of the Human Brain
▶ Added discussion of partial dominance and codominance
▶ Added discussion of gene silencing to section on epigenetics
▶ Added discussion of the Zika virus and microencephaly
▶ Expanded and updated discussion of brain development in childhood, adolescence,
and young adulthood
▶ Refreshed Building Better Health feature with a discussion of nutritional cognitive
neuroscience and healthy aging
▶ Added comprehensive summary table of milestones in prenatal nervous system
development
▶ Added comprehensive summary table of major neurodevelopment events from
birth to late adulthood
Chapter 6 Vision
▶ Expanded and clarified discussion of visual receptive fields, including additional
illustration
▶ Updated discussion of vision restoration
▶ Added discussion of “The Dress” to section on color constancy
Chapter 7 Nonvisual Sensation and Perception

▶ Added section on synaesthesia
▶ Updated and expanded discussion of loudness perception
▶ Added discussion of the McGurk effect
▶ Added discussion of the effects of using profanity to the experience of pain
▶ Expanded discussion of the connections between olfaction and dementia
Chapter 8 Movement
▶ Added discussion of M lines and tropomyosin
▶ Expanded discussion of motor units
▶ Added discussion of direct and indirect systems linking the thalamus, basal ganglia,
and cerebral cortex
▶ Updated discussion of amyotrophic lateral sclerosis (ALS)
▶ Added discussion of CRISPR to Thinking Ethically feature on Gene Doping for
Strength
xxiv Preface
Chapter 9 Homeostasis, Motivation, and Reward

▶ Moved topic of reward from Chapter 14 to better reflect its role in motivated
behavior
▶ Added discussion of drive and incentive approaches to motivation
▶ Streamlined discussions of temperature, thirst, and hunger
▶ Updated discussion of obesity
▶ Introduced distinction between liking and wanting to organize discussion of reward
pathways
▶ Expanded and updated discussion of the neurochemistry of reward
Chapter 10 Sexual Behavior

▶ Added discussion of mini-puberty
▶ Updated discussion of Turner syndrome
▶ Added summary section on lessons learned from sex chromosome abnormalities
▶ Added discussion of the organizing roles of sex chromosome genes
▶ Added discussion of brain imaging and gender dysphoria
▶ Updated discussion of hormones and sexual interest
▶ Updated discussion of oxytocin and bonding
▶ Updated discussion of the treatment of sexual dysfunction
▶ Updated Building Better Health feature on antidepressant-induced sexual dysfunction
Chapter 11 Biorhythms
▶ Updated discussion of shift work type circadian disorders
▶ Expanded discussion on mechanisms of light therapy
▶ Updated and expanded discussion of default mode network (DMN) and its rela-
tionship to levels of consciousness
▶ Expanded discussion of eye movements during REM
▶ Updated and expanded discussion of memory consolidation and reconsolidation
during sleep, including elimination and maintenance of synapses during REM
▶ Added section on sleep and emotion regulation
Chapter 12 Learning and Memory

▶ Reorganized chapter material to reflect the continuity between learning and
memory
▶ Expanded and clarified discussion of long-term potentiation (LTP)
▶ Expanded discussion of working memory, including a further discussion of the
role of LTP
▶ Expanded discussion of memory consolidation and reconsolidation, including the
role of transcription factors
▶ Updated discussion of conditioned fear
▶ Updated discussion of Henry Molaison (Patient H.M.) and the relationships be-
tween the hippocampus and cortex in memory formation
▶ Updated the discussion of stress effects on memory
Preface xxv
Chapter 13 Cognitive Neuroscience

▶ Changed chapter title to reflect the organization of topics
▶ Added section on decision making, including a discussion of neuroeconomics
▶ Updated section on the development of lateralization
▶ Updated section on the lateralization of emotion and music abilities
▶ Expanded discussion of intelligence to include genome-wide association studies
▶ Updated Thinking Ethically feature on performance-enhancing drugs for the mind
Chapter 14 Emotion, Reward, Aggression, and Stress

▶ Reordered topics to present major theories earlier in the chapter
▶ Added sections on emotion regulation and social cognition
▶ Updated and expanded discussion of detecting deception
Chapter 15 Neuropsychology
▶ Updated section on Alzheimer’s disease
▶ Updated section on brain tumors
▶ Added information about the Zika virus
▶ Updated section on migraine
Chapter 16 Psychopathology
▶ Added discussion of the NIMH Research Domain Criterion project (RDoC)
▶ Updated section on autism spectrum disorder (ASD), including discussion of pa-
ternal age, default mode network differences, and microglia actions
▶ Clarified and updated research on saccades and schizophrenia
▶ Expanded discussion of the roles of dopamine and glutamate in schizophrenia
▶ Updated discussion of tardive dyskinesia
▶ Updated discussion of causes of bipolar disorder
▶ Introduced the hypothesis that inflammation plays a role in major depressive dis-
order (MDD)
▶ Expanded the discussion of obsessive-compulsive disorder (OCD)
▶ Updated the section on posttraumatic stress disorder (PTSD)
▶ Updated the section on antisocial personality disorder (ASPD)
▶ Refreshed the Building Better Health feature with a discussion of the gut micro-
biota and mental disorders
MindTap
MindTap® Psychology: MindTap® for Discovering Behavioral Neuroscience: An
Introduction to Biological Psychology 4th edition is the digital learning solution that
helps instructors engage and transform today’s students into critical thinkers. Through
paths of dynamic assignments and applications that you can personalize, real-time
course analytics, and an accessible reader, MindTap helps you turn cookie cutter into
cutting edge, apathy into engagement, and memorizers into higher-level thinkers.
xxvi Preface
As an instructor using MindTap you have at your fingertips the right content and
unique set of tools curated specifically for your course all in an interface designed
to improve workflow and save time when planning lessons and course structure. The
control to build and personalize your course is all yours, focusing on the most relevant
material while also lowering costs for your students. Stay connected and informed in
your course through real-time student tracking that provides the opportunity to adjust
the course as needed based on analytics of interactivity in the course.
Instructor Ancillaries
Online Instructor’s Manual: The manual includes learning objectives, key terms,
a detailed chapter outline, a chapter summary, lesson plans, discussion topics, student
activities, media tools, a sample syllabus and an expanded test bank. The learning
objectives are correlated with the discussion topics, student activities, and media tools.
Online PowerPoints: Helping you make your lectures more engaging while
effectively reaching your visually oriented students, these handy Microsoft
PowerPoint® slides outline the chapters of the main text in a classroom-ready
presentation. The PowerPoint® slides are updated to reflect the content and
organization of the new edition of the text.
Cengage Learning Testing, powered by Cognero®: Cengage Learning Testing,
Powered by Cognero®, is a flexible online system that allows you to author, edit,
and manage test bank content. You can create multiple test versions in an instant and
deliver tests from your LMS in your classroom.
Acknowledgments xxvii
Acknowledgments
I view this text as a work in progress. Please take a moment to share your thoughts
and suggestions with me: lfreberg@calpoly.edu. You can also find me on Facebook, on
Twitter as @biopsych, and on my blog: http://www.laurafreberg.com/blog.
I am enormously grateful for the team that Cengage assembled to help make this
book a reality. First, I would like to thank my professional colleagues who reviewed the
many drafts of one or more of the four editions:
John Agnew, University of Colorado at Paul J. Kulkosky, Colorado State

Boulder University Pueblo
James E. Arruda, Mercer University Gloria Lawrence, Wayne State College
Giorgio Ascoli, George Mason University Simon Levay
Ronald Baenninger, Temple University Charles F. Levinthal, Hofstra University
Aileen Bailey, St. Mary’s College of David R. Linden, West Liberty State College
Maryland Michael R. Markham, Florida
Jeffrey S. Bedwell, University of Central International University
Florida Richard Mascolo, El Camino College
Steve Bradshaw, Bryan College Robert Matchock, Penn State University,
Virginia Bridwell, Bellevue College Altoona
Gayle Brosnan-Watters, Slippery Rock Janice E. McPhee, Florida Gulf Coast
University University
John P. Bruno, Ohio State University Jody Meerdink, Nebraska Wesleyan
Allen E. Butt, Indiana State University University
Deborah Carroll, Southern Connecticut Melinda Meszaros, Clarke College
State University Maura Mitrushina, California State
David A. Cater, John Brown University University Northridge
James Chrobak, University of Robert R. Mowrer, Angelo State University
Connecticut, Storrs Mark Nawrot, North Dakota State
Cynthia Cimino, University of South University
Florida Irene Nielsen, Bethany College, Lindsborg
James R. Coleman, University of South Terry Pettijohn, Ohio State University,
Carolina Marion
Sherry Dingman, Marist College David W. Pittman, Wofford College
Aaron Ettenberg, University of California, Joseph H. Porter, Virginia
Santa Barbara Commonwealth University
Bob Ferguson, Buena Vista University Jerome L. Rekart, Rivier College
Thomas M. Fischer, Wayne State University John C. Ruch, Mills College
John P. Galla, Widener University Ronald Ruiz, Riverside City College
Cynthia Gibson, Creighton University Lawrence J. Ryan, Oregon State University
Ben Givens, Ohio State University Carl Samuels, Glandale Community College
Karen Glendenning, Florida State Anthony C. Santucci, Manhattanville
University College
C. Hardy, Columbia College Virginia F. Saunders, San Francisco
James G. Holland, University of Pittsburgh State University
Richard Howe, College of the Canyons Royce Simpson, Spring Hill College
Joyce Jadwin, Ohio University Don Smith, Everett Community College
Robert A. Jensen, Southern Illinois Marcello Spinella, Richard Stockton
University College of New Jersey
Katrina Kardiasmenos, Bowie State Sheralee Tershner, Western New
University England College
Joshua Karelitz, Pennsylvania State C. Robin Timmons, Drew University
University, New Kensington Barbara Vail, Rocky Mountain College
Camille Tessitore King, Stetson University Rachael Volokhov, Kent State University,
Norman E. Kinney, Southeast Missouri Salem
State University Dana Wallace, Jamestown College
xxviii Acknowledgments
Linda L. Walsh, University of Northern Yeuping Zhang, Lewis and Clark College
Iowa Margaret H. White, California State
Frank M. Webbe, Florida Institute of University Fullerton
Technology Xiojuan Xu, Grand Valley State University
Kimberly Wear, High Point University Robert M. Zacharko, Carleton University
Stephen P. Weinert, Cuyamaca College Phillip Zoladz, Ohio Northern University
The many instructors who responded to market research surveys and interviews over
the past several years have also contributed greatly to the text and its ancillaries.
I would like to give a special thank you to several colleagues. Skirmantis Janusonis
of the University of California, Santa Barbara, has provided not only encouragement
but also gentle suggestions for improved accuracy. Skirmantis kindly provided me with
an opportunity to photograph some of his human brains (one appears in Chapter 1),
and he shared an image of his fantastic dissection of the hippocampus for us to use in
Chapter 2. Simon LeVay is a gifted researcher and a truly elegant writer. His careful
review of the chapter on sexual behavior for the first edition is deeply appreciated. I
am indebted to Simon for pointing out many important studies included in the sec-
tions on sexual orientation and gender differences in cognition. Marie Banich at the
University of Colorado at Boulder provided her expert opinions on the art program,
and Gayle Brosnan-Watters at Arizona Christian University contributed many helpful
suggestions as a reviewer and while working with me on the third edition of the Test
Bank. John Cacioppo of the University of Chicago generously shared his considerable
insight into the social aspects of neuroscience. Konstantinos Priftis of the University
of Padova supplied several helpful corrections to the second edition. Robert Matchock
of the University of Pennsylvania, Altoona, has been a friendly voice of encouragement
beginning with the first edition. The late Larry Butcher of UCLA, one of my former
professors, provided valuable assistance in the fine-tuning of my cholinergic pathway
image in Chapter 4. Rick Howe and the students and faculty of the psychology depart-
ment at College of the Canyons in California not only took the time to meet with me to
share their reactions to the first edition but fed me cookies as well.
At Cengage, Erin Schnair, product manager for psychology, has championed my
text through this edition. Linda Man, my content developer for the fourth edition, was
my sounding board and partner every step of the way. Ruth Sakata Corley, senior proj-
ect manager, guided the production of the text and coordinated innumerable aspects of
the art and text program. Photo researcher Geeta Kamath tenaciously pursued the pho-
tos that enliven the text. Vernon Boes, art director, guided our design to new heights in
appeal and usability. One faculty adopter referred to the design as “Steve Jobs-esque,”
which we view as the ultimate compliment. I would also like to thank the many other
capable and creative professionals at Cengage who contributed to this project in ways
large and small over the years of its development.
Finally, I would like to thank my family for their patience and support. Roger, my
husband of 45 years, read every word, helped me choose photos, and even contributed
some of our original sketches when I had trouble articulating what I wanted to show.
My daughters, Kristin, Karen, and Karla, frequently offered their encouragement and
perspectives. My Australian shepherd, Ronnie, waited patiently by my desk until it was
time to clear the mind with a walk through some beautiful San Luis Obispo scenery.
I am very much indebted to all.
Laura A. Freberg
CHAPTER 1
What Is Behavioral Neuroscience?
Learning Objectives chapter OutLine

L01 Classify the subfields of neuroscience, and explain Neuroscience as an Interdisciplinary Field
how behavioral neuroscience fits within the field. Historical Highlights in Neuroscience
Ancient Milestones in Understanding the Nervous
L02 Interpret the significance of the major historical System
highlights in the study of the nervous system. The Dawn of Scientific Reasoning
Modern Neuroscience Begins
L03 Differentiate the brain imaging technologies,
Interim Summary 1.1
including CT, PET, SPECT, MRI, fMRI, and DTI.
Behavioral Neuroscience Research Methods
L04 Assess the use of microscopic, recording, Microscopic Methods
stimulation, optogenetic, lesion, and biochemical Imaging
methods in behavioral neuroscience. Recording
Brain Stimulation
L05 Analyze the relative strengths and weaknesses Lesion
of twin studies, adoption studies, and genetic Biochemical Methods
screens for understanding behavior. Genetic Methods
Interim Summary 1.2
L06 Evaluate the ethical standards used to protect
Research Ethics in Behavioral Neuroscience
human and animal research participants.
Human Participant Guidelines
Animal Subjects Guidelines
Interim Summary 1.3
Chapter Review
CONNECTING TO RESEARCH: Thinking about Your Food

as Healthy or Indulgent Affects Your Physical Reactions
to Eating
BEHAVIORAL NEUROSCIENCE GOES TO WORK:
What Can I Do with a Degree in Neuroscience?
THINKING ETHICALLY: Can We Read Minds with Brain
Imaging?
BUILDING BETTER HEALTH: When Is It Appropriate to
Use Placebos?
2 C HAPT ER 1 What Is Behavioral Neuroscience?
Neuroscience as an Interdisciplinary Field

Neuroscience is the scientific study of the brain and nervous system, in health and in
disease (UCLA, 2008). Neuroscientists strive to understand the functions of the brain
and nervous system across a number of levels of analysis, using molecular, cellular,
synaptic, network, computational, and behavioral approaches (see ● Figure 1.1). You
might think of this field as analogous to Google Earth. We can zoom in to see the tiniest
detail and then zoom back out again to see the “big picture.”
Beginning at the most microscopic level, the molecular neuro-
scientist explores the nervous system at the level of the molecules
that serve as its building blocks. We will cover their work in our
ZouZou/Shutterstock.com
chapters on neural cell physiology (Chapter 3), psychopharmacol-

ogy (Chapter 4), and genetics (Chapter 5). Starting with DNA and
RNA and the proteins resulting from gene expression, the molecular
Behavioral neuroscientist attempts to understand the chemicals that build the
system and make neural functioning possible.
Zooming out just a bit from the molecular level of analysis, we
find the cellular neuroscientist hard at work outlining the structure,
physiological properties, and functions of single cells found within
the nervous system. These isolated cells would be of no use unless
they could forge connections, which they do at junctions we call
Network synapses. Synaptic neuroscience examines the strength and flexibil-
ity of neural connections, which underlie complex processes such as
learning and memory.
Beyond the single synapse, we find that interconnected neurons
iStock.com/Henrik5000
form pathways or networks. In contemporary neuroscience, we are

seeing a move away from the idea that “this structure engages in
Synaptic Computational
this function” to ideas that more accurately reflect neural networks
that have been identified. We are more likely to say that “this struc-
ture participates in a network connecting these other structures to
engage in this type of processing.”
Zooming out perhaps to the most global point of view, we find
behavioral neuroscience, also known as biological psychology,
which is the primary focus of this textbook. Behavioral neuroscien-
Cellular tists use all of the previous levels of analysis, from the molecular up
through the network, in their efforts to understand the biological
correlates of behavior. The relationship between biology and behav-
ior is reciprocal—biology can impact our behavior, and behavior,
Courtesy of Laura Freberg
including your cognitions and emotions, impacts biology. The

experiment described in the Connecting to Research feature illus-
trates this reciprocal relationship. The way you think about what
Molecular
you’re eating can influence your physical reaction to the food.
Like the neurosciences in general, behavioral neuroscience
● Figure 1.1 The Neurosciences: Building
looks at the activity of the nervous system in health and in cases
from Molecule to Behavior
of illness, injury, and psychological disorder. Subspecialties within
behavioral neuroscience include cognitive neuroscience, or the
study of the biological correlates of information processing, learning
and memory, decision making, and reasoning. We investigate these topics in depth
in chapters on sensation and perception (Chapters 6 and 7), learning and memory
(Chapter 12), and cognition (Chapter 13). Social neuroscience, covered in Chapter 14,
neuroscience The scientific study explores the interactions between the nervous system and our human social environ-
of the brain and nervous system
in health and in disease. ment and behavior.
behavioral neuroscience/ Computational neuroscience runs parallel to the types of neuroscience described
biological psychology The study so far, but it draws from computer science, electrical engineering, mathematics, and
of the biological foundations of
behavior, emotions, and mental physics to produce models of the nervous system from the molecular up through the
processes. behavioral levels of analysis. The predictions from these computational models can
Neuroscience as an Interdisciplinary Field 3
Connecting to Research
THINKING ABOUT YOUR FOOD AS HEALTHY OR INDULGENT AFFECTS
YOUR PHYSICAL REACTIONS TO EATING
O ne of the key aspects of behavioral neuroscience

that we introduce in this chapter is the reciprocal
nature of the relationship between biology and behavior.
At the same time, the way you think about eating
can affect your body’s responses to food. Simply telling
research participants that their 380-calorie milkshake
Most of us are quite familiar and comfortable with the was either “sensible” or “indulgent” produced big
idea that our biology can influence our behavior, but the differences in the amount of ghrelin released (Crum,
influence of behavior on biology is perhaps less obvious. Corbin, Brownell, & Salovey, 2011) (see ● Figure 1.2).
A clever study by Alia Crum and her colleagues (2011) After consuming the “sensible” shake, ghrelin levels
provides a compelling illustration of the reciprocal influ- didn’t change much, which means that the participants
ences of biology and behavior by exploring factors that still felt hungry and unsatisfied. After consuming the
lead to hunger and feelings of fullness or satiety. “indulgent” shake, ghrelin levels dropped significantly
When you haven’t eaten for a while, you release and the participants felt more satisfied. In other words,
a gut hormone called ghrelin that tells your brain that thinking about the shake as sensible or indulgent (be-
you’re hungry (see Chapter 9). Once you eat some- havior) had a remarkable impact on biology (large or
thing, the release of ghrelin is suppressed, and you no small ghrelin release). There are no guarantees that
longer feel hungry. So your biology (how much ghrelin praising your healthy salad as indulgent will help you
is released) initiates behavior (feeling hungry or feeling stick to your diet, but it can’t hurt to try!
satisfied). So far, so good.
Courtesy of Alia Crum and Gibbs Graphics
980
● Figure 1.2 Biology and Behavior Form
960 Reciprocal Relationships Participants’ ghrelin
940 levels fluctuated differently after drinking an
Ghrelin in pg/ml
identical 380-calorie milkshake depending on

920 whether the participant viewed a “Sensi-Shake”
or “Indulgent” label at Minute 20. By Minute 90,
900
participants viewing the “Indulgent” label had
880 lower ghrelin levels, associated with feeling
satisfied, than participants viewing the “Sensi-
860 Indulgence
Shake” label. Thinking about the milkshake a
Sensi-Shake
certain way (behavior) had a dramatic effect on
840
ghrelin release (biology).
20 60 90
Time (minutes) Source: Gibbs Graphics, Crum, A. J., Corbin,
W. R., Brownell, K. D., & Salovey, P. (2011). Mind
over milkshakes: Mindsets, not just nutrients,
determine ghrelin response. Health Psychology,
30(4), 424–429.
then be tested against living systems, forming a cooperative symbiosis with research-
ers in other areas of neuroscience. One of the practical applications of computational
neuroscience is the use of neural decoding, or using neural activity to estimate what
the brain is doing, in the development of sophisticated prosthetic devices. The devices
can use neural activity to figure out the brain’s intentions to move in certain ways or
perceive certain types of input as touch.
These different levels of analysis complement each other rather than compete
with one another. Because of the diversity of skills needed to pursue each of these
approaches, neuroscience is an essentially interdisciplinary field of study, reaching
across traditional academic departments of biology, chemistry, psychology, medicine,
mathematics, physics, engineering, and computer science.
The need for better understanding of the nervous system has never been greater.
The Society for Neuroscience (2012) reported that neurological illnesses impact one
out of six Americans annually at a cost of more than $500 billion for treatment, which
does not include the cost of disability. Delaying the onset of Alzheimer’s disease on
an average of five years would save the United States $50 billion in annual health care
costs. Connections between biology and behavior are not just relevant to neurological
disease, but they inform our understanding of health in general. Compared to 100
years ago, when most people died from infectious diseases, today’s killers (cancer,
diabetes, heart disease) are tightly linked to behavior. Reflecting recognition of the
role of behavior in illness, the standardized Medical College Admission Test (MCAT)
for medical school applicants now contains a significant number of questions about
psychology and behavioral neuroscience.
Illness is only part of the human equation. We also need to understand how
the nervous system responds in typical ways to promote well-being, including better
relationships, better parenting, better child development, and better thinking and
learning. Through improved understanding of the nervous system and its interac-
tions with behavior, scientists and practitioners will be more thoroughly prepared
to tackle the significant challenges to health and well-being faced by contemporary
world populations.
Behavioral
Neuroscience WHAT CAN I DO WITH A DEGREE IN NEUROSCIENCE?
GOES TO WORK
O ne of the pleasures of teaching courses in behav-

ioral neuroscience occurs when a student sudden-
ly falls in love with the field. “This is for me,” the student
neuroscience graduates work in substance abuse
counseling or in mental health facilities. Neuroscience
is used in some unexpected places as well. A grow-
might say, but the question that usually follows is, “But ing trend in advertising agencies is to use brain imag-
how can I make a living doing this?” The answers to ing and other technologies to gauge public reactions
this question are as diverse as the field. Because neuro- to advertising. Web and application designers use
science is so broad, opportunities can be found down eye-tracking technology to evaluate the user experi-
many different paths. ence (UX), such as whether a person “sees” and pro-
Like many other fields, neuroscience has more cesses the important features of a webpage and has a
opportunities for people with more education. Many good time while doing so.
practicing neuroscientists have medical degrees, The ongoing burst in neuroscience technologies
PhDs, or even both. This does not mean that jobs are is likely to continue to shape the field, and additional
unavailable for students with undergraduate degrees, opportunities are likely to emerge. In the meantime,
however. Students with undergraduate degrees can any student interested in neuroscience would benefit
be employed as research assistants in pharmaceutical from gaining the best possible skills in general science,
firms, universities, and government agencies. Some research methods, mathematics, and statistics.
Historical Highlights in Neuroscience

The history of neuroscience parallels the development of tools for studying the ner-
vous system. Early thinkers made progress in spite of limited scientific methods and
technologies.
Ancient Milestones in Understanding the Nervous System

Our earliest ancestors apparently had at least a rudimentary understanding about
the brain’s essential role in maintaining life. Archaeological evidence of brain sur-
gery suggests that as long as 7,000 years ago, people tried to cure others by drilling
holes in the skull, a process known as trephining or trepanation (see ● Figure 1.3).
Because some skulls have been located that show evidence of healing following the
drilling procedure, we can assume that the patient lived through the procedure and
that this was not a postmortem ritual. What is less clear is the intent of such surgeries.
Possibly, these early surgeons hoped to release demons or relieve feelings of pressure
(Clower & Finger, 2001).
The Edwin Smith Surgical Papyrus represents
the oldest known medical writing in history yet fea-
tures many sophisticated observations (Breasted,
1930). The Egyptian author of the Papyrus clearly
understood that paralysis and lack of sensation in the
body resulted from nervous system damage. Cases
Mary Martin/Science Source/Photo Researchers, Inc

of nervous system damage were usually classified as
“an ailment not to be treated,” indicating the author’s
understanding of the relatively permanent damage
involved.
Building on the knowledge taken from ancient
Egypt, the Greek scholars of the fourth century
BCE proposed that the brain was the organ of sen-
sation. Hippocrates (460–379 BCE) correctly iden-
tified epilepsy as originating in the brain, although
the most obvious outward signs of the disor- ● Figure 1.3 Prehistoric Brain Surgery As far back in history
der were muscular convulsions (see Chapter 15). as 7,000 years ago, people used trepanation (trephining), or the
Galen (130–200 CE), a Greek physician serving the drilling of holes in the skull, perhaps to cure “afflictions” such
Roman Empire, made careful dissections of animals as demonic possession. Regrowth around some of the holes indi-
(and we suspect of the mortally wounded gladiators in cates that at least some of the patients survived the procedure.
his care as well). Galen believed erroneously that the More recently, trephining has resurfaced as a DIY (do it yourself)
ventricles played an important role in transmitting process, possibly as a type of self-injurious behavior.
messages to and from the brain, an error that influ-
enced thinking about the nervous system for another
1,500 years (Aronson, 2007).
The Dawn of Scientific Reasoning

The French philosopher René Descartes (1596–1650) argued in favor of mind–body
dualism. For Descartes and other dualists, the mind is neither physical nor accessible
mind–body dualism A
to study through the natural sciences. In contrast, the modern neurosciences are based philosophical perspective put
on monism rather than dualism. The monism perspective proposes that the mind is forward by René Descartes in
the result of activity in the brain, which can be studied scientifically. Descartes’s ideas which the body is mechanistic,
whereas the mind is separate and
were very influential, and even today some people struggle with the idea that factors nonphysical.
such as personality, memory, and logic simply represent the activity of neurons in the monism A philosophical
brain. Later in the chapter, our discussion of research ethics presents another legacy perspective characteristic of the
neurosciences in which the mind
of Descartes’s ideas. Because many shared his view of animals as mechanical, not sen- is viewed as the product of activity
tient, beings, experiments were carried out on animals that seem barbaric to many in the brain.
● Figure 1.4 Luigi Galvani

Demonstrated a Role
for Electricity in Neural
Communication This engrav-
ing illustrates the basement
laboratory of Luigi Galvani,
where his experiments with
frogs not only helped establish
understanding of the electrical
nature of neural communica-
tion, but also inspired Mary
Shelley to write Frankenstein.
Time Life Pictures/Getty Images

modern thinkers. In 1865, Claude Bernard (1813–1878) wrote that “the science of life
is a superb and dazzingly lighted hall which may be reached only by passing through
a long and ghastly kitchen” (p. 15).
Between 1500 and 1800, scientists made considerable progress in describing the
structure and function of the nervous system (see ● Figure 1.4). The invention of the
light microscope by Anton van Leeuwenhoek in 1674 opened up a whole new level of
analysis. Work by Luigi Galvani and Emil du Bois-Reymond established electricity as
the mode of communication used by the nervous system. British physiologist Charles
Bell (1774–1842) and French physiologist François Magendie (1783–1855) demon-
© Bettmann/CORBIS
strated that information traveled in one direction, not two, within sensory and motor
nerves.
● Figure 1.5 Phrenology

Modern Neuroscience Begins
Bust Franz Josef Gall and As late as the beginning of the twentieth century, many scientists, including Italian
his followers used maps like researcher Camillo Golgi, continued to support the concept of the nervous system as a
this one to identify traits vast, interconnected network of continuous fibers. Others, including the Spanish anat-
located under different omist Santiago Ramón y Cajal, argued that the nervous system was composed of an
parts of the skull. Bumps on
array of separate, independent cells. Cajal’s concept is known as the Neuron Doctrine.
the skull were believed to
Golgi and Cajal shared the Nobel Prize for their work in 1906. Ironically, Cajal used a
indicate that the underlying
trait had been “exercised.”
stain invented by Golgi to prove that Golgi was incorrect.
Although Gall’s system was The road to our current understanding of the nervous system has not been without
an example of very bad sci- its odd turns and dead ends. The notion that specific body functions are controlled
ence, the underlying princi- by certain areas of the brain, called localization of function, began with an idea pro-
ple that functions could be posed by Franz Josef Gall (1758–1828) and elaborated by Johann Gasper Spurzheim
localized in the brain turned (1776–1832). These otherwise respectable scientists proposed a “science” of phrenology
out to be valuable. that maintained that the structure of people’s skulls could be correlated with their indi-
vidual personality characteristics and abilities. A phrenologist could “read” a person’s
character by comparing the bumps on his or her skull to a bust showing the supposed
location of each trait (see ● Figure 1.5). Although misguided, Gall and Spurzheim’s
phrenology The misguided effort
to correlate character traits with work did move us away from the metaphysical, nonlocalized view of the brain that had
bumps in the skull. persisted from the time of Descartes. Instead, Gall and Spurzheim proposed a more
modern view of the brain as the organ of the mind, composed of interconnected, coop-
erative, yet relatively independent functional units.
Further evidence in support of localization of function in the brain began to
accumulate. In the mid-1800s, a French physician named Paul Broca correlated the
damage he observed in patients with their behavior and concluded that language
functions were localized in the brain (see Chapter 13). Incidentally, it was also Broca
who brought the practice of trepanation, described earlier in this chapter, to the
attention of the scientific community (Clower & Finger, 2001). In 1870, Gustav
Theodor Fritsch (1838–1927) and Eduard Hitzig (1838–1907) described how elec-
trically stimulating the cortex of a rabbit and a dog produced movement on the
opposite side of the body. Localization of function in the brain became a generally
accepted concept.
Further evidence of the mind’s physical, nonmystical nature was provided by
physiologists and psychophysicists (see Chapters 6 and 7). Hermann von Helmholtz
(1821–1894) demonstrated that the mind had a physical basis by asking participants
to push a button as soon as they felt a touch. The participants reacted faster when their
thigh was touched than when their toe was touched because the more distant signal
from the toe would take more time to reach the brain.
The founding of modern neuroscience has often been attributed to the British neu-
rologist John Hughlings Jackson (1835–1911). Hughlings Jackson proposed that the
nervous system was organized as a hierarchy, with simpler processing carried out by
lower levels and more sophisticated processing carried out by higher levels, such as the
cerebral cortex. For example, we normally inhibit aggressive behaviors, associated with
activity in lower levels of the brain, by engaging our higher cortical executive func-
tions. When a person consumes alcohol, however, the cortical inhibition might fail,
leaving the lower parts of the brain to initiate a bar fight. We meet Hughlings Jackson
again in Chapter 15, in which his contributions to the understanding of epilepsy will be
discussed.
Progress in the neurosciences over the past 100 years accelerated rapidly as new
methods became available for studying the nervous system. Charles Sherrington not
only coined the term synapse (defined as the point of communication between two
neurons) but also conducted extensive research on reflexes and the motor systems of
the brain (see Chapter 8). Otto Loewi demonstrated chemical signaling at the synapse
(see Chapter 3), using an elegant research design that he claims came to him while
asleep. Sir John Eccles, Bernard Katz, Andrew Huxley, and Alan Hodgkin furthered
our understanding of neural communication. You will meet many more contemporary
neuroscientists as you read the remainder of this text. The ranks of neuroscientists con-
tinue to grow, with membership in the Society for Neuroscience expanding from 500
members in 1969 to approximately 38,000 members in 90 countries as of 2016 (Society
for Neuroscience [SfN], 2016).
INTE RIM SUMMARY 1 .1
Highlights in the Neuroscience Timeline
Historical Period Significant Highlights and Contributions
Ca. 3000 BCE ● Egyptians discard brain during mummification process;

however, published case studies indicate accurate
observations of neural disorders.
Ca. 400 BCE−200 CE ● Hippocrates recognizes that epilepsy is a brain disease.

● Galen makes accurate observations from dissection; however,
he believed erroneously that fluids transmitted messages.
(continued)
Historical Period Significant Highlights and Contributions
1600–1800 ● René Descartes suggests mind-body dualism.

● Anton van Leeuwenhoek invents the light microscope.
● Galvani and du Bois-Reymond discover that electricity
transmits messages in the nervous system.
1800–1900 ● Bell and Magendie determine that neurons communicate

in one direction and that sensation and movement are
controlled by separate pathways.
● Gall and Spurzheim make inaccurate claims about
phrenology, but their notion of localization of function in the
nervous system is accurate.
● Paul Broca discovers localization of speech production.
● Fritsch and Hitzig identify localization of motor function in
the cerebral cortex.
1900–Present ● Ramón y Cajal declares that the nervous system is composed

of separate cells; he shares the 1906 Nobel Prize with Camillo
Golgi.
● John Hughlings Jackson explains brain functions as a
hierarchy, with more complicated functions carried out by
higher levels of the brain.
● Otto Loewi demonstrates chemical signaling at the synapse.
● Charles Sherrington coins the term synapse; he wins the
Nobel Prize in 1932.
● Sir John Eccles, Andrew Huxley, and Alan Hodgkin share
the 1963 Nobel Prize for their work in advancing our
understanding of the way neurons communicate.
● Bernard Katz receives the 1970 Nobel Prize for his work on
chemical transmission at the synapse.
● Society for Neuroscience counts about 38,000 members
in 2016.
Summary Points
1. Neuroscience is the field that explores the structures, functions, and development of
the nervous system in illness and in health. Behavioral neuroscience is the branch of
the neurosciences that studies the correlations between the structures and functions
of nervous system and behavior. (LO1)
2. Although some periods of enlightenment regarding the relationship between the
nervous system and behavior emerged among the Egyptians and Greeks, the major
advances in behavioral neuroscience have been relatively modern and recent. (LO2)
3. Highlights in the neuroscience timeline include discoveries regarding the electrical
and chemical nature of neural communication, the control of sensation and motor
functions by separate nerves, the role of single cells as building blocks for the ner-
vous system, and the localization of functions in the brain. (LO2)
Review Questions
1. How would you describe the goals and methods of the interdisciplinary field of
neuroscience?
2. What historical discoveries contributed to our modern understanding of the brain and
behavior? Which concepts actually led us in the wrong direction?
Behavioral Neuroscience Research Methods

The methods described in this section have helped neuroscientists discover the struc-
ture, connections, and functions of the nervous system and its components. From the
level of single molecules to the operation of large parts of the nervous system, we now
have the ability to make detailed observations that would likely astonish the early pio-
neers of neuroscience. The choice of methods depends very much on the goals and
research questions of the neuroscientist. Each method has its shares of strengths and
weaknesses, but the use of multiple methods to answer a single research question can
compensate for any gaps in a particular method.
Microscopic Methods
Microscopic, or histological, methods provide means for observing the structure, orga-
nization, and connections of individual cells. The first investigation of nerve tissue
under a microscope was conducted by Anton van Leeuwenhoek in 1674. However, due
to the technical challenges of viewing structures as small and complex as those found
in the nervous system, most of the advances in microscopy occurred following the
development of stronger, clearer lenses during the 1800s.
The naked eye can perceive objects that are at least 0.2 mm in size. To see anything
smaller requires magnification (see ● Figure 1.6). Magnification alone does not guar-
antee a clear image, however, as you probably have noticed when you’ve zoomed in on
an image on your computer. You just get the same blurry image with bigger pixels. Our
Naked eye
100mm
Rat brain
Light microscope
10mm
Brain
Electron microscope
1mm
100mm Cell body
Dendrites
10mm
1mm
Synapses
100nm Dendritic spines
Synaptic vesicles
10nm
Ion channels
1nm
DNA
Ions and
0.1nm atoms
● Figure 1.6 Size of Objects Visible
Using Various Microscopy Methods
histology The study of cells and
tissues at the microscopic level.
ability to see small structures also depends on resolution, or the ability to tell two points
of light apart.
Today’s light microscope can magnify an image 1,000 times while maintaining ade-
quate resolution, allowing scientists to observe a feature as small as a synapse. Because
of the ways our visual systems process visible light, discussed in Chapter 6, using a light
microscope to magnify images beyond about 1,000 times compromises resolution to an
unacceptable degree. Fortunately, other methods have been discovered that maintain
both high levels of magnification and resolution. Electron and fluorescent microscopy
can resolve images at the level of the nanoscopic scale, or between 1–100 nanometers.
A nanometer is one billionth of a meter. In other words, these methods are about one
million times more powerful than the naked eye.
Fluorescent microscopy works by labeling a specimen with a particular fluorphore,
a chemical that emits light when excited by light. The specimen is then illuminated
with a different wavelength, which causes it to emit fluorescence that is then analyzed.
The downside to this process is that you have to work quickly. The specimen does not
emit fluorescence indefinitely.
Electron microscopy uses beams of electrons rather than beams of visible light
(photons; see Chapter 6) to illuminate a specimen. Because electrons have much shorter
wavelengths than photons, resolution is much improved. Electron microscopy allows
us to see features as small as a single molecule of neurochemical! The main disadvan-
tage of electron microscopy is that it cannot be used on living cells. Specimens require
dehydration and chemical treatments that are incompatible with life.
Tissue to be studied under the microscope must be prepared for viewing in a series
of steps. Tissue must be made thin enough to allow light (or electrons) to pass through
it. Brain tissue is fragile and somewhat watery, which makes the production of thin
enough slices impossible without further treatment. To solve this problem, the first step
in the histological process is to “fix” the tissue, either by freezing it, dehydrating it, or
treating it with formalin, a liquid containing the gas formaldehyde. Formalin not only
hardens the tissue, making it possible to produce thin slices, but it also preserves the
tissue from breakdown by enzymes or bacteria. Freezing the tissue accomplishes these
objectives as well. The resulting samples can be embedded in gelatin, paraffin wax, or
plastic to provide further stability and ease of slicing.
Once tissue is fixed, it is sliced or “sectioned” by a special machine known as a
microtome (see ● Figure 1.7). Some microtomes look and work like a miniature ver-
sion of the meat slicers found in most delicatessens. The tissue is pushed forward while
● Figure 1.7 Using a Microtome

to Section Patient H. M.’s
Brain Researchers at UC San Diego
broadcast the careful sectioning of
the brain of the late Henry Molaison,
otherwise known as the famous
amnesic patient H. M., via streaming
video on the Internet. Molaison’s tem-
poral lobe surgery and the resulting
memory deficits he experienced are
familiar to all students of psychology.
The Brain Observatory
microtome A device used to

make very thin slices of tissue for
histology.
a sliding blade moves back and forth across the tissue, producing slices. Other micro-
tomes use a vibrating knife, similar to your electric toothbrush. These are particularly
useful when sectioning tissue that is not frozen. For viewing tissue under the light
microscope, tissue slices between 10 and 80 μm (micrometers) thick are prepared. A
micrometer is one one-millionth of a meter or one one-thousandth of a millimeter.
Electron microscopes require slices of less than 1 μm. Sectioning a single rat brain pro-
duces several thousand slides. The fragile slices are stored in a saline solution and then
mounted on slides for viewing.
Even when fixed and mounted on slides, nerve tissue would appear nearly trans-
parent under the microscope if it were not for a variety of specialized stains. Researchers
select particular stains depending on the features they wish to examine. For example, to
make a detailed structural analysis of a small number of single cells, the best choice is
a Golgi stain, named after its discoverer, Camillo Golgi. On the other hand, you might
be more interested in identifying clusters of cell bodies, the major bulk of the nerve cell,
within a sample of tissue. In this case, you would select a Nissl stain. A myelin stain
would allow you to follow pathways carrying information from one part of the brain to
another by staining the insulating material that covers many nerve fibers. If you know
Courtesy of Siemens Healthcare

where a pathway ends but would like to discover its point of origin, you should use
horseradish peroxidase. When this enzyme is injected into the end of a nerve fiber, it
travels backward toward the cell body. Antibodies, proteins normally produced by the
immune system to identify invading organisms, can be combined with a variety of dyes
to highlight particular proteins found in cells in a process known as immunohisto-
chemistry (IHC). As mentioned previously, a wide variety of fluorophores can be used
to label structures, genes, and proteins for examination under fluorescent microscopes.
Viewing the nervous system at the cellular level continues to provide considerable
insight into its functioning, but viewing cells and their connections in the context of the
entire brain is pushing our understanding to new levels. Newly developed methods in
GJLP, CNRI/Science Source

optical imaging allow researchers to literally construct a “see through brain” (Deisseroth,
2016). By removing fats (lipids) from a sample, scatter of light is reduced, and the remain-
ing sample becomes essentially transparent. Using this method, researchers were able to
show that populations of individual cells in an animal’s frontal lobe that were active in
either positive or negative experiences were wired quite differently (Ye et al., 2016).
● Figure 1.8 CT Scans
Hounsfield’s original ma-
Imaging chine took several hours to
obtain data for a single slice
Modern imaging techniques provide significant advantages over autopsy, the study (above). Modern scanning
of the body after death. With current imaging technologies, we can watch the living equipment is much faster
brain as it engages in processes such as reading (Chapter 13) or emotional response and can produce detailed
(Chapter 14). We can identify differences in the ways the brains of serial murderers 3-D images (bottom).
function compared with the brains of typical people (Chapter 16).
COMPUTERIZED TOMOGRAPHY (CT) The groundwork for brain imaging was laid
by German physicist Wilhelm Röntgen, who discovered X-rays in 1896. Röntgen was
astonished to learn that X-rays could move through the human body and that they Golgi stain A stain developed by
would produce a negative photographic image of the body’s major structures. The first Camillo Golgi used to observe
single neurons.
X-ray ever taken was an image of Röntgen’s wife’s hand. Nissl stain A stain used to view
Normal X-rays do not do a very good job of imaging soft tissue. If you have ever populations of cell bodies.
seen an X-ray of your head taken by your dentist or orthodontist, you probably saw myelin stain A stain used to trace
neural pathways.
bones and teeth, but not much brain. However, with adaptations made possible by horseradish peroxidase A stain
more modern computers, X-rays can be used to image previously unseen anatomical used to trace axon pathways from
structures. The first commercial computerized tomography (CT) scanner was intro- their terminals to points of origin.
optical imaging A method for
duced in 1971 by Godfrey Hounsfield, based on previous work by William Oldendorf preparing a brain for examination
and Allan Cormack (see ● Figure 1.8). “Tomography” comes from the Greek words that literally makes it transparent.
tomos, or slice, and graphia, to write or describe. CT technology provided the first computerized tomography (CT)
An imaging technology in which
high-resolution look at a living brain. More modern CT technology allows for the con- computers are used to enhance
struction of highly detailed three-dimensional images. X-ray images.
● Figure 1.9 PET Scans Show

Patterns of Brain Activation PET
Courtesy Dept. of Energy Office of Public Affairs

scans do not provide much struc-
tural detail, but they do offer a
clear picture of brain activity. Red
and yellow areas are most active,
whereas blue and black areas are
least active. These three images
show different patterns of brain
activity during a visual task (left), (a) Visual Task (b) Listening Task (c) Problem-solving Task
a listening task (center), and a
problem-solving task (right).
However useful CT scanning may be for medical imaging, the technology does
have drawbacks for research purposes. Although it provides excellent structural infor-
mation, a CT scan cannot distinguish between a living brain and a dead one. In other
words, the CT scan provides no information regarding activity levels in the brain. This
limits the usefulness of CT in helping us answer questions about behavior.
POSITRON EMISSION TOMOGRAPHY (PET) The next major breakthrough in

imaging technology was the development of the positron emission tomography, or
PET, scan, which allowed researchers to observe brain activity for the first time (see
● Figure 1.9). PET scans do not provide good information about brain structure, however.
PET scans were made possible by the invention of the gamma camera, which is
used to detect radiation released by radioactive atoms that are decaying or breaking
up. Beginning in the mid-1970s, Michael Phelps and Edward Hoffman of Washington
University began to apply this basic technique to the study of brain function (Hoffman,
Phelps, Mullani, Higgins, & Ter-Pogossian, 1976; Phelps, Hoffman, Mullani, Higgins,
& Ter-Pogossian, 1976).
PET brain studies combine radioactive tracers with a wide variety of molecules,
including oxygen, water, neurochemicals, and drugs. Data from gamma rays resulting
from the breakdown of the tracer are recorded by detectors and fed to a computer, by
which the data are reconstructed into images. Typically, programmers have assigned
red and yellow to areas of high activity and green, blue, and black to areas of low activ-
ity. Newer PET machines can take images of adjacent slices at the same time, which
allows for three-dimensional reconstruction of brain activity. A closely related proce-
dure, single photon emission computed tomography (SPECT), is less expensive than
PET but provides less visual detail.
For most research purposes, PET and SPECT have been replaced by functional
MRI (fMRI), discussed in an upcoming section. Compared to fMRI, however, PET has
a unique advantage in its ability to provide information about the levels of activity of
particular neurochemicals in the brain. For example, administering a tracer that binds
to serotonin receptors can identify their location and levels of activity in a living person.
MAGNETIC RESONANCE IMAGING (MRI) Magnetic resonance imaging, or MRI,

has become a standard medical diagnostic tool and a valuable research asset. Raymond
Damadian, Larry Minkoff, and Michael Goldsmith produced the first MRI image in
positron emission tomography 1977. This imaging technology uses powerful magnets to align hydrogen atoms within
(PET) An imaging technique that
provides information regarding a magnetic field. Next, radio frequency (RF) pulses are directed at the part of the body
the localization of brain activity. to be imaged, producing “resonance,” or spinning, of the hydrogen atoms. When the RF
magnetic resonance imaging pulses cease, the hydrogen atoms return to their natural alignment within the magnetic
(MRI) An imaging technique
that provides very high resolution field. As the atoms “relax,” each becomes a miniature radio transmitter, emitting a
structural images. characteristic pulse that is detected by the scanner.
MRI takes advantage of the fact that different types of brain tissue are made of dif-
ferent components. “Gray matter,” or tissue largely made up of the cell bodies of nerve
cells, contains large amounts of protein and carbohydrates. Nerve fibers, or “white mat-
ter,” are covered by insulating material that is mostly fat. The fluid surrounding the cells
of the brain is essentially salt water. These components, protein/carbohydrates, fats,
and water behave differently when exposed to MRI methods, leading to stronger sig-
nals from one type relative to another. The stronger signals are assigned a darker color
in the resulting image. Each small area of tissue is assigned a voxel, which is a three-di-
mensional version of a pixel. The darkness or coloration of each voxel represents the
level of pulse activity in an area.
Functional MRI (fMRI) allows us to correlate brain activity with the presentation
of a stimulus, the presence of an emotional state, or the performance of a particular
task in living humans. The first fMRI of the brain was conducted by Belliveau et al.
(1991). Functional MRI takes advantage of the fact that active neurons require more
oxygen than less active neurons, and that variations in blood flow to a particular area
will reflect this need.
The use of fMRI to track blood flow in the brain
was previewed in the nineteenth century by America’s
first official psychologist, William James, who was
impressed by the observations of Italian physiologist
Angelo Mosso on patients with head injuries. Due
to the nature of these injuries, in which some of the
patients’ skull bones were missing or damaged, Mosso
was able to measure and correlate blood flow with
the patients’ mental activities (Mosso, 1881). James’s
reflections on Mosso’s work sound very contempo-
rary: “Blood very likely may rush to each region of
the cortex according as it is most active, but of this
Courtesy of Laura Freberg

we know nothing” (James, 1890, vol. 1, p. 99). Mosso’s
observations were confirmed by Roy and Sherrington
(1890), who reported the existence of “an automatic
mechanism by which the blood supply of any part of
the cerebral tissue is varied in accordance with the
activity of the chemical changes which underlie the ● Figure 1.10 Functional Magnetic Resonance Imaging
functional action of that part” (p. 105). (fMRI) Tracks Cerebral Blood Flow This image demonstrates
How does fMRI track cerebral blood flow? Hemo- the use of fMRI to identify parts of the brain (the red and
globin, the protein molecule that carries oxygen yellow areas) that became selectively active when the author
within the blood, has different magnetic properties engaged in a “finger tap” exercise (touching each digit of her
right hand one by one with her thumb).
when combined with oxygen or not (Ogawa, Lee, Kay,
& Tank, 1990). Consequently, signals from a voxel will
change depending on the oxygenation of the blood
in that area, known as the Blood Oxygenation Level
Dependent (BOLD) effect. Let’s look at an example of the author’s own results of a
standard demonstration conducted at the Brain Imaging Laboratory at the University
of California, Santa Barbara (see ● Figure 1.10). Scans were taken as the author alter- voxel Short for “volume
pixel.” A pixel is the smallest
nated 20 second intervals of remaining very still with 20 second intervals of touching distinguishable square part of a
her right thumb to each of the other digits of her right hand one at a time. The image two-dimensional image. A voxel
highlights the voxels that showed changes in activity correlated with movement and is the smallest distinguishable
box-shaped part of a three-
stimulation. dimensional image.
The same machinery used for MRI and fMRI can also produce images using functional MRI (fMRI) A
diffusion tensor imaging (DTI). This technique allows researchers to map connec- technology using a series of MRI
images taken one to four seconds
tivity in the brain by tracking the movement of water in the fiber pathways of the apart in order to assess the activity
nervous system (Le Bihan & Breton, 1985; Moseley et al., 1990). (see ● Figure 1.11). of the brain.
DTI cannot, however, tell us about the direction of information flow. Using this tech- diffusion tensor imaging (DTI)
Use of MRI technology to trace
nique, researchers have observed evidence of greater than normal connectivity in the fiber pathways in the brain by
frontal lobes of toddlers with autism spectrum disorder (Solso et al., 2016). tracking the flow of water.
Recording
Although perhaps less dramatic than the imaging tech-
niques, methods that allow researchers to record the
electrical and magnetic output from the brain continue
to be useful. As we will see in greater detail in Chapter 3,
Living Art Enterprises/Science Source

nerve cells are capable of generating small electrical
charges across their membranes, much like miniature
batteries. Any electrical current will generate magnetic
fields. Although small in scale, this electrical and mag-
netic activity can be recorded using electrodes either on
the surface of the skull or brain or imbedded within the
brain tissue itself.
● Figure 1.11 Diffusion Tensor Imaging (DTI) Using
MRI technology, the flow of water down the length of THE ELECTROENCEPHALOGRAM (EEG) The first
nerve fibers can be imaged to construct maps of the fiber recordings of the human brain’s electrical activity,
pathways of the brain. measured through electrodes placed on the scalp,
were made by a German psychiatrist, Hans Berger, in
1924. Berger noted that the recordings varied during
wakefulness, sleep, anesthesia, and epilepsy. Chapters 11 and 15 investigate the
relationships between the EEG and these states of consciousness in greater detail.
For many years, EEG technology did not change much. Although it was useful in
the study of sleep and the diagnosis of epilepsy, EEG did not offer anything further to
our understanding of brain function. With the advent of more powerful computers,
however, new quantitative methods for analyzing EEG recordings became possible.
Computerized EEG brain tomography can be used to generate maps of activity, mak-
ing it possible to pinpoint the source of abnormal activity. EEG brain tomography can
be used to follow a patient through withdrawal from psychoactive drugs or during a
coma. The technique can aid in diagnoses of many disorders, including schizophrenia,
electroencephalogram (EEG) The dementias, epilepsy, and attention deficit hyperactivity disorder (see Chapters 15 and
recording of the brain’s electrical 16). Computerized analysis of EEG recordings can be used to generate animations of
activity through electrodes placed
on the scalp. activity over time and for the construction of three-dimensional maps of brain activity.
These analysis tools have breathed new life into EEG technology.
THINKING Ethically
CAN WE READ MINDS WITH BRAIN IMAGING?
I n the 2002 film Minority Report, Tom Cruise plays a

police officer who arrests people for crimes they have
not yet committed. How close are we to knowing what
to detect deception (Langleben et al., 2016). As we will
see in our discussion of emotion in Chapter 14, typical
lie detection technologies, which rely on indirect mea-
people are planning to do before they actually do it? sures of arousal, are so poor that they typically are not
Perhaps closer than you think. admissible as court evidence in the United States. The
Current imaging technologies, such as fMRI, do not ability to detect deception directly through brain imag-
exactly allow us to “read” the human mind, but we are ing promises enormous benefits to criminal justice and
getting ever closer to that ability (Tong & Pratte, 2012). national security, but raises equally substantial concerns
For example, researchers were able to use fMRI record- about ethics. Not only can we envision the legislative
ings taken while a participant watched Steve Martin need to expand our constitutional right to privacy to
playing Inspector Clouseau in the movie The Pink Pan- include “mental privacy,” but legislators will also have
ther to reconstruct the image (Nishimoto et al., 2011). to consider that the new technology may acquire infor-
Of even greater importance for society is the fact mation from brain scans that is irrelevant to the purpose
that researchers are closing in on the ability to use fMRI of the investigation in the first place.
EVENT-RELATED POTENTIALS (ERPS) A variation of basic EEG technology is

the recording of event-related potentials (ERPs). This technique allows researchers
to correlate the timing of activity of cortical neurons recorded through scalp
electrodes with stimuli presented to the participant. The brain’s electrical activity
in response to a stimulus, such as a tone, is quite small compared to the activity
normally recorded in an EEG, so responses to many presentations of a stimulus are
averaged (see ● Figure 1.12). This type of analysis can be helpful in cases in which a
person’s behavior does not provide a clear indication of whether a particular stimulus
has been perceived. For example, young children with autism spectrum disorder (see
Chapter 16) often behave as though their hearing were impaired. When spoken to
by parents or others, a child with autism spectrum disorder often shows no reaction
at all. Through observations of event-related potentials to sound, we can determine
whether the sound information has reached the brain.
More recently, improvements in the ability to analyze ERP data have allowed
researchers to document the precise timing and localization of brain activity in ways
that are often superior to fMRI (Cacioppo et al., 2013; Cacioppo, Weiss, Runesha, &
Cacioppo, 2014). Using this approach, researchers were able to show that lonely people
distinguish between negative social stimuli (words like alone and excluded) and nega-
tive nonsocial stimuli (words like angry and sad) faster than people who are not lonely
(Cacioppo, Balogh, & Cacioppo, 2015).
MAGNETOENCEPHALOGRAPHY (MEG) Magnetoencephalography (MEG) allows

researchers to record the brain’s magnetic activity (Cohen, 1972). Active neurons put
out tiny magnetic fields. By “tiny,” we mean that the fields generated by neural activity
are about one billion times smaller than Earth’s magnetic field and about 10,000
times smaller than the field surrounding a typical household electric wire. The major
advantage of recording magnetism rather than electrical activity from the brain relates
to the interference of the skull bones and other tissues separating the brain from the
electrodes, which prevents a large amount of the brain’s electrical activity from being
recorded using EEG. In contrast, the skull bones and tissues allow magnetism to pass
through without any reduction. In addition, recordings of the magnetic fields produced
by the brain can be taken much faster than either fMRI or PET scans, providing a
Single EEG
+ ● Figure 1.12 Event-Related
recording
Potentials (ERPs) The analysis
of event-related potentials
Electrode allows researchers to map
Amplifier
the brain’s EEG response to
20 mv
environmental stimuli. In
this example, a characteristic
waveform emerges when
responses to the presentation
of a tone are averaged over
– 100 trials.
100 200 300 400 500 600 700
Stimulus Time (msec)
ERP
+
(average
of 100 or
2 mv
more EEG
event-related potential (ERP) An
recordings) alteration in the EEG recording
produced in response to the
– application of a particular
100 200 300 400 500 600 700
stimulus.
Stimulus Time (msec) magnetoencephalography (MEG)
A technology for recording the
magnetic output of the brain.
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“WHY.”
I’m sad, for, loving thee, I know full well

That this world’s talk, with its calumnious spell,
Will never spare thy fresh youth’s opening flower.
For every happy day and sunny hour,
Fate will exact in grief and tears his pay.
I’m sad because I see my loved one gay!
MOSCOW.
Moscow, I love thee with a filial love,

Strong, burning, tender, which a Russian knows!
I love the holy gleam thy brows above,
Thy battled Kremlin in its calm repose.
In vain the foreign Potentate[6] essayed,
Great Russian giant of a thousand years,
To cope with thee, and, by deceit betrayed,
To make thee bow thy soul to craven fears.
In vain the Stranger spurred; you reared; he fell!
The world grew silent ’neath his mighty spell;
Whilst thou alone didst live, my stately one,
Thou heir to glories ours, and ours alone!
Thou livest still, and every stone of thine
Doth tell of generations tales divine.
FOOTNOTES:
[6] Napoleon.
FROM PUSHKIN.
⁂
I wander down the noisy streets,

I enter crowded fanes,
I join in youthful revelries,
I give my fancy reins.
I say, “The years are flying fast,

And seen we scarce are here,
Before we reach eternal tombs;
For each the hour is near.”
I glance upon the lonely oak,

The patriarch of the wood,
And think, “He’ll live through my brief day,
He through my father’s stood.”
I fondly kiss the little child,

And, kissing, think, “Good-bye!
I’m giving up my place to you.
You bloom; ’tis mine to die.”
Thus every day, thus every hour,

I’m wont with thought to spend,
And strive to guess the birthday-date
Of my approaching end.
Ah! where will Fate send Death to me?

Abroad? in war? on deep?
Or will a neighbouring valley hold
My cold dust in its keep?
Yet, though I know my lifeless form
Must rot where’er I die,
I’d fondly wish near my loved home,
In my own land, to lie.
There, round the entrance to the grave,

Let young life freely play,
And careless Nature calmly smile
With ageless beauty gay!
ANACREONTIC.
We know the steed of mettle

By the breed-marks branded on it;
We know the haughty Highlander
By his plumed and towering bonnet;
And I know the happy lovers
By the love-light in their eyes,
Where, its tale of joyance telling,
The languid flame doth rise.
(TO HIS WIFE.)
No! not for me the wild tumultuous gladness,

The rapturous rush, the transports, and the madness,
The moans, the cries, the young Bacchante makes,
When, clinging close in coilings like a snake’s,
With wounding kiss, and gush of hot caresses,
For the last moments’ thrills she quiveringly presses.
Far dearer thou, my gentle one, to me,

And happy I—distracted more by thee—
When yielding to long prayers with gentle grace,
You press me softly in your meek embrace;
Modestly cold, to love with passion fraught
You scarce respond; you conscience seem of naught;
Yet warm and warmer glowing, till at last,
As ’twere against your will, you share my blast.
⁂
Let me not lose my senses, God;

Better the pilgrim’s scrip and rod,
Or toil and hunger sad.
Not that I prize this mind of mine,
Or that my reason to resign
I should not be right glad,
If only then they’d set me free.
At large! How sportively I’d flee
To where the dark wood gleams!
I’d sing in raving ecstasies,
Forgetting self in fantasies
Of changeful wondrous dreams.
To the wild waves I’d lend an ear,
And glancing upward, full of cheer,
Would scan the open sky;
And strong and free I’d rush amain,
A whirlwind sweeping o’er the plain,
Crashing through woods I’d fly.
But there’s the rub! You lose your sense—
Are dreaded like a pestilence,
And clapped in prison drear.
They chain you to the idiot’s yoke,
And, through the cage-bars, to provoke
The wild beast they draw near.
No more the nightingale to hear
At midnight singing sweet and clear,
Nor greenwood’s rustling strains,
But only brother-madmen’s cries,
The nightly keeper’s blasphemies,
And shrieks, and clang of chains.
⁂
I’ve overlived aspirings,

My fancies I disdain;
The fruits of hollow-heartedness,
Sufferings alone remain.
’Neath cruel storms of Fate,

Withers my crown of bay,
A sad and lonely life I lead,
Waiting my latest day.
Thus, struck by latter cold,

While howls the wintry wind,
Trembles upon the naked bough
The last leaf left behind.
PETER THE GREAT.
With autocratic hand

He boldly sowed the light;
He did not scorn his native land—
He knew her destined might.
A carpenter, a seaman,
A scholar, hero, he,
With mighty genius on the throne,
A labourer was incessantly.
THE PROPHET.
By spiritual thirst opprest,

I hied me to the desert dim,
When lo! upon my path appeared
The holy six-winged seraphim.
My brow his fingers lightly pressed,
Soothing my eyelids into rest:
Open my inward vision flies,
As ope a startled eaglet’s eyes.
He touched my ears, and they were filled
With sounds that all my being thrilled.
I felt a trembling fill the skies,
I heard the sweep of angels’ wings,
Beneath the sea saw creeping things,
And in the valleys vines arise.
Over my lips awhile he hung,
And tore from me my sinful tongue—
The babbling tongue of vanity.
The sting of serpent’s subtlety
Within my lips, as chilled I stood,
He placed, with right hand red with blood.
Then with a sword my bosom cut,
And forth my quivering heart he drew;
A glowing coal of fire he put
Within my breast laid bare to view.
As corpse-like on the waste I lay,
Thus unto me God’s voice did say—
“Prophet, arise! confess My Name;
Fulfil My will; submit to Me!
Arise! go forth o’er land and sea,
And with high words men’s hearts inflame!”
⁂
Play, my Kathleen;
No sorrow know.
The Graces flowers
Around thee throw.
Thy little cot
They softly swing,
And bright for thee
Dawns life’s fresh spring.
For all delights
Thou hast been born;
Catch, catch wild joys,
In life’s young morn!
Thy tender years
To love devote;
While hums the world,
Love my pipe’s note.
A MONUMENT.[7]
I’ve raised myself no statue made with hands;

The People’s path to it no weeds will hide.
Rising with no submissive head, it stands
Above the pillar of Napoleon’s pride.
No! I shall never die; in sacred strains
My soul survives my dust, and flies decay—
And famous shall I be, while there remains
A single Poet ’neath the light of day.
Through all great Russia will go forth my fame,
And every tongue in it will name my name;
And by the nation long shall I be loved,
Because my lyre their nobler feelings moved;
Because I strove to serve them with my song,
And called forth mercy for the fallen throng.
Hear God’s command, O Muse, obediently,
Nor dread reproach, nor claim the Poet’s bay;
To praise and blame alike indifferent be,
And let fools say their say!
FOOTNOTES:
[7] Like our Shakespeare, Pushkin knew his own merits.
THE POET.
Until Apollo calls the Bard

To share the holy sacrifice,
Plunged in the petty cares of life
The Poet’s spirit lies.
Silent and still his sacred lyre,

His soul to sleep a prey,
Amongst earth’s worthless sons he seems
More worthless, p’raps, than they.
But once the sacred summons rings

And strikes his eager ears,
The Poet’s soul, like eagle roused,
On upward pinion steers.
Then earthly pleasures cease to charm;

He scorns the babbling crowd;
No more beneath their Idol’s feet
His haughty head is bowed.
He flies—and wild and stern his moods,

His notes, now grave, now gay—
To shores where lonely billows play,
To depths of whispering woods.
FROM NADSON.
⁂
Pity the stately cypress trees;

How freshly green they spring!
Ah! why amidst their branches, child,
Have you put up your swing?
Break not a single fragrant bough.
Oh, take thy swing away
To heights where thick acacias bloom;
Mid dusty olives play!
Thence you can see the Ocean,
And, as your swing ascends,
Through greening boughs a sunny glimpse
The sea in laughter sends
Of white sails in the distance dim,
Of white gulls far away,
Of white flakes foaming on the sands,
A fringe of snowy spray.
FROM NEKRASOF.
TE DEUM.
In our village there’s cold and there’s hunger;

Through the mist the sad morn rises chill;
Tolls the bell—the parishioners calling
From afar to the church on the hill;
Austere and severe and commanding
Pealed that dull tone thro’ the air.
I spent in the church that wet morning;
I can never forget the scene there.
For there knelt the village hamlet,
Young and old in a weeping crowd;
To be saved from the grievous famine
The people prayed aloud.
Such woe I had seldom witnessed,
Such agony of prayer,
And unconsciously I murmured,
“O God, the people spare!”
“Spare their friends, too, in Thy mercy!

Oh, hear our heartfelt cry!
For those who strove to free the serf
We lift the prayer on high;
For those who bore the battle’s brunt
And lived to win the day,
For those who’ve heard the serf’s last song,
To Thee, O God, we pray.”
THE PROPHET.
Ah! tell me not he prudence quite forgot;

That he himself for his own fate’s to blame.
Clearer than we, he saw that man cannot
Both serve the good and save himself from flame.
But men he loved with higher, broader glow;

His soul for worldly honours did not sigh;
For self alone he could not live below,
But for the sake of others he could die.
Thus thought he—and to die, for him, was gain.

He will not say that “life to him was dear;”
He will not say that “death was useless pain;”
To him, long since, his destiny was clear.
⁂
Offer my Muse a friendly hand,

For I can sing no other song.
Who feels no woe, nor flames at wrong,
Loves not his Fatherland.

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Discovering behavioral neuroscience:

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Laura A. Freberg is Professor of Psychology at California Polytechnic State University,

1 What Is Behavioral Neuroscience? 1

2 Functional Neuroanatomy and the Evolution of the

3 Neurophysiology: The Structure and Functions

5 Genetics and the Development of the Human Brain 145

7 Nonvisual Sensation and Perception 225

9 Homeostasis, Motivation, and Reward 301

10 Sexual Behavior 337

11 Sleep and Waking 375

12 Learning and Memory 415

13 Cognitive Neuroscience 455

14 Emotion, Aggression, and Stress 489

About the Cover

Photo by Dr. Sonja Pyott, University of North Carolina

1 What Is Behavioral Neuroscience? 1

CONNECTING TO RESEARCH: Thinking about Your Food as Healthy or

2 Functional Neuroanatomy and the Evolution

The Brain’s Blood Supply 34

CONNECTING TO RESEARCH: Linking the Brain and the Immune System 31

3 Neurophysiology: The Structure and Functions of the

CONNECTING TO RESEARCH: Astrocytes, HIV, and the Blood–Brain Barrier 70

CONNECTING TO RESEARCH: Otto Loewi and “Vagus Stuff” 110

5 Genetics and the Development of the Human Brain 145

Building a Brain 158

CONNECTING TO RESEARCH: Epigenetics, Gene Expression, and Stress 155

7 Nonvisual Sensation and Perception 225

BUILDING BETTER HEALTH: Earbuds and Hearing Loss 238

Motor Systems of the Brain 281

THINKING ETHICALLY: Gene Doping for Strength 271

9 Homeostasis, Motivation, and Reward 301

Reward Pathways 333

BUILDING BETTER HEALTH: Understanding the Benefits of Fever 306

10 Sexual Behavior 337

BEHAVIORAL NEUROSCIENCE GOES TO WORK: How Do Therapists Treat Gender

11 Sleep and Waking 375

CONNECTING TO RESEARCH: A Composite Scale of Morningness 377

12 Learning and Memory 415

Systems Supporting Classical Conditioning 436

BEHAVIORAL NEUROSCIENCE GOES TO WORK: What Is Neuro Education? 421

13 Cognitive Neuroscience 455

CONNECTING TO RESEARCH: Savants and Laterality 466

14 Emotion, Aggression, and Stress 489

CONNECTING TO RESEARCH: Facial Expressions Predict Assault 498

Recovery and Treatment in Neurocognitive Disorders 550

BEHAVIORAL NEUROSCIENCE GOES TO WORK: Preparing to Be a Clinical

Antisocial Personality Disorder (ASPD) 587

CONNECTING TO RESEARCH: Genetic Overlap in Five Disorders 557

▶ Building Better Health features provide an additional opportunity for students to

New Content for the Fourth Edition

Chapter 1 What Is Behavioral Neuroscience?