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Copyright 2019 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. WCN 02-200-202
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
Copyright 2019 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, in whole or in part. Due to electronic rights, some third party content may be suppressed from the eBook and/or eChapter(s).
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An Introduction to Biological Psychology,
Fourth Edition Unless otherwise noted, all content is © Cengage.
Laura A. Freberg
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Print Number: 01 Print Year: 2017
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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and she smiled immensely at everybody, she fitted in with complete
harmony to the young jollity of the morning and the month.
Christopher thought she looked like a good-natured young shark.
‘What an awful pity,’ she said. ‘We might have gone for excursions
together.’
And she said it with such a heartiness of chagrin that they all laughed.
The end of it was that when the sandwiches came and the Moncktons
went, the Jerrolds, still talking, went with them, first to the entrance of the
hotel garden, then into the road, then, still talking, along the road.
The Jerrolds not having talked for a week were unable now to leave off.
Mr. Jerrold found himself wishing to tell the small agreeable lady who he
was, and why, and how, and did so with a completeness that surprised
himself. His daughter, striding on ahead with the young man—they seemed
naturally to shoot ahead together, the two young ones, the minute they got
on to the road—explained just as completely to her companion, who
appeared at once to tumble to it, the dreadful feeling of being about to burst
after a week’s flopping round with somebody who couldn’t be left while
one rushed all over the island, and couldn’t, owing to age and infirmity and
being a father and all that, rush too.
‘Just look at those youngsters forging along,’ said Mr. Jerrold, smiling
complacently at the two figures in front, at their four worsted-stockinged
legs moving so quickly in step, at their swinging arms, and their bare heads
turned to each other while they talked and laughed.
His companion looked, but said nothing. He wondered what relation she
was to the young man that she too should be a Monckton, and decided that
she must be either his father’s second wife or his aunt by marriage. Not a
blood relation, clearly; they were too much unlike.
‘Is there anything more delightful in the world,’ said Mr. Jerrold, gazing
benevolently at the pair ahead, ‘than a wholesome English boy and girl?’
At this his companion murmured something that he understood meant
that the two would soon be out of sight, which they certainly would be if
they went on at that pace, and he said, ‘Yes—quite. Hi, there,’ he called,
‘you youngsters! Steady—we can’t keep up!’
But the wind was against him, and they strode on unheeding.
‘Not that,’ said Mr. Jerrold, turning gallantly to the lady, ‘it would be
anything but my gain if they did go on. Why not let them? And you and I sit
down somewhere and talk.’
‘But Christopher has got the sandwiches,’ said his new friend.
‘So he has. Christopher. Delightful name. Attractive youth. Well, let
Christopher eat them, sharing them with Billy somewhere at the end of their
first twenty miles or so, and allow me the pleasure of offering you lunch in
the hotel.’
‘How very kind of you. But Christopher——’
‘Well, you see he doesn’t hear,’ said Mr. Jerrold. ‘I don’t suppose he’ll
even notice that we’re not following. When young people get together.... I
assure you it would give me the greatest pleasure to entertain you, and we
can sit afterwards in that nice garden in comfortable chairs till they come
back.’
Mr. Jerrold paused persuasively in the road. After a week of not
speaking to a soul but his daughter he ached to entertain. The two on in
front turned a corner and disappeared. Mr. Jerrold, whose wife had been
dead some years, felt the situation unfolding romantically. This was a dear
little lady, and she looked as though she needed taking care of. He had a
strong wish to give her lunch.
‘But——’ she began.
‘We will consider no buts,’ said Mr. Jerrold, even more gallantly. ‘Your
nephew—is he your nephew?—won’t notice——’
‘He’s my husband,’ said the little lady, flushing a very bright scarlet.
Well; what a surprise. Mr. Jerrold was really most surprised. Not that the
lady wasn’t, he was sure, a most agreeable wife for any man to have, but
that the young man seemed so very young to have one at all; and if at all
she ought, to match him, be a mere slip of a girl,—somebody about Billy’s
age.
Yes; it was a surprise. Mr. Jerrold didn’t quite know what to say.
‘In that case——’ he began.
But really he hadn’t an idea what to say, and stood in the middle of the
road staring down at the little lady through his monocle—he wore a
monocle—and she stared back at him, while the flush slowly ebbed away
out of her face.
III
Her nephew. So that was what Christopher seemed to be to this impartial
stranger. It gave Catherine more than a shock, it made her heart feel as if it
stood still. And his surprise, his humiliating surprise, when she said
Christopher was her husband, and her own discomfort when she told him....
Was it so much marked, then, the difference between them? It hadn’t
been in London. Why, in London before they married they had often stood
arm in arm in front of a glass and laughed to see how no one would guess,
really no one could possibly guess, that they were not very nearly of an age.
Besides what about all those bus-conductors and people calling her Miss?
One of them had even called her Missie—‘Take care, now, Missie,’ he had
said, catching her by the arm, ‘don’t you go jumping off before we’re
stopped and breaking your neck and getting us into trouble with your young
man’—but he, she was afraid, had been drinking. It must be because she
was so tired now always that she looked older. To-day she was tired,
yesterday she had been tired—oh, but so tired, so tired.
She stared up at Mr. Jerrold, while the flush faded out of her face, and
thought how dreadful it was going to be if every time she was tired people
took her for Christopher’s aunt. What a humiliation. And inevitably sooner
or later he would notice it himself, and hear it too from strangers, just as she
was hearing it from this stranger.
‘Let us sit down,’ said Mr. Jerrold sensibly, ‘and wait for them to come
back.’
And a day or two afterwards, when Christopher, impelled by his desire
for movement, by a terrific longing to do something, anything, that wasn’t
lying in grass reading poetry to Catherine—if he didn’t read poetry to her
she was surprised and asked him why, because at the beginning he had
wanted to do nothing else—hired a two-seater and drove her round the
island, stopping for the night at a little place on the west side where there
was a small hotel they liked the look of, on their going in and asking if they
could be put up for the night the young lady in the office, glancing at them,
said she was very sorry but she had only one room vacant.
‘But we only want one,’ said Christopher, surprised at this answer. ‘We
want a double room, that’s all.’
‘Oh, I’m sorry——’ said the young lady, turning red and bending over
her ledger to hide her confusion. ‘Yes——’ she said, running her finger
down the page, ‘I can give you No. 7.’
‘Do you think she thought we weren’t married?’ said Christopher,
amused, when they were in No. 7 undoing their suit-cases. ‘Or do you think
she thought we were so grand that we couldn’t do without a sitting-room?’
Catherine, very busy it seemed with her suit-case, said nothing. She
couldn’t have. She felt sick, as if some one had hit her head. Again she had
been taken for Christopher’s aunt. Or even for his—no, her mind swerved
aside from that word; it simply refused to look at it.
They saw no more of the Jerrolds, though Christopher had talked of long
and violent scrambles with the eagerly acquiescent Billy while they sprinted
on ahead that morning before he realised that Catherine had been left
behind out of sight. When he did discover this he had turned back at once.
‘Come on,’ he had said to the surprised Billy, seizing her wrist, ‘we must
run.’
And he had run; and she had run, thinking it great fun but wondering
why they should be running; and after that, when they all joined up again,
her father had taken her back to the hotel and the Moncktons had gone for
their picnic by themselves, and she had never set eyes on them again nor
heard anything more of the promised scrambles.
But one thing she had heard, and with astonishment, from her father, and
that was that Mr. Monckton was Mrs. Monckton’s husband.
‘No!’ cried Billy, her eyes very round; adding, after a silence, ‘Good
Lord.’
‘Quite,’ said her father.
A few days more and the honeymoon was at an end. Christopher had not
attempted again to leave Catherine, for she didn’t seem well, though she
assured him she was,—assured him eagerly, almost painfully eagerly, and
that it was only the spring. He wasn’t quite able to believe this, and stayed
with her and petted her. She loved to lie quite quiet in his arms, out of doors
or anywhere, while he read to her or they both snoozed. He suppressed his
fidgetings, because he knew if he said he wanted to walk she would want to
walk with him, and then she would be tired out and he after all not
exercised. It struck him once as odd how little they talked. They used to talk
and talk before they were married. Now they hardly said anything, except
when they began to whisper, and then it wasn’t talk, but emotion clothing
itself scantily in words. Still, it had been a heavenly, heavenly time;
something to remember joyfully all one’s days.
‘When we’re old,’ he said, the last evening, ‘how we shall think of this.’
Just as if, she thought, pressing close to him so as to hide from the
thought, when he had got to the stage of being old she wouldn’t, far ahead
of him, be long past thinking at all.
He had to be back at his office the end of the second week, and the last
night in their abode of bliss they hardly slept at all, so loth were they to lose
any minutes of what was left of their honeymoon in unconsciousness; and
the effect of this was that in the morning, while Christopher was as blithe as
a lark and breakfasted cheerfully and packed up with zeal, Catherine could
hardly move for fatigue, and was really shocked by her leaden face when
she saw it in the glass.
Luckily he noticed nothing that time; he was too busy packing up, too
much pleased in the fresh morning to be doing something different, to be
starting on a journey. Besides, wasn’t he going to work like a navvy now?
Hadn’t he got something to work for,—responsibilities, the sweetest, most
wonderful in the world? He itched to be at it, to do well, make her proud of
him, earn money for her as that old George had earned money for her.
With gusto he swept his scattered things into his suit-case, whistling the
love music out of The Immortal Hour as he packed, with gusto he settled
the bills and tipped the woman, with gusto he walked, his arm through
Catherine’s, down the path to the gate for the last time, and waved to the
buttercup-field in which he would not again have to lie. He was in high
spirits. It was jolly getting back to work, beginning it again in these new
delicious conditions, with Catherine to speed him in the morning and
welcome him back at night. Now he would have variety; now he would
have work and love, absence and presence, in their right proportion.
He was very happy. It seemed incredible to him, as he fondly looked
down at her when they were in the bright warm sunshine on the ferry, that
he had actually attained his heart’s desire and got Catherine. Life was
splendid,—packed with possibilities, a thing of the utmost magnificence.
The waters of the Solent danced and sparkled; white wings flashed out of
the deep blue of the sky; the sun lay hot across the back of his neck; the
wind was fresh and salt in his face; the world looked as it must have looked
on the morning of its first day. Old Lewes—he would go and dig out old
Lewes to-morrow, and make him come and lunch in one of those jolly little
restaurants where the food was good and ladies didn’t go, and yarn his head
off. And on Saturday he would take Catherine down and introduce her to
his uncle, who would certainly adore her, and she would wander about the
garden and enjoy her darling little self while he gave the old boy the round
of golf he knew he was thirsting for. So was he thirsting for it. His
honeymoon had been wonderful, but a fortnight is enough. It wouldn’t of
course be enough, and one would never then want it to end, if one were
going to be torn from one’s beloved when it was over. But here they were,
he and she, entering into the joys, the varied joys of married life, with him,
the male, girding up his loins in the morning and going forth to labour until
the evening, as men from time immemorial had girded themselves and gone
forth, and coming back at night to his nest and his mate. And this after all
was better in the long run than a honeymoon, just as real good bread and
butter was better than everlasting cake.
‘I’m so happy,’ he said, slipping his arm round her and giving her a
quick hug when no one was looking. He couldn’t see her face; she was
sitting too close to him, besides having put on a gauze motor-veil.
‘Darling Chris,’ she murmured, smiling through her veil.
But she would have liked it better if he hadn’t been quite so exuberantly
happy on that particular morning. After all, it was the finish of their
honeymoon, and they would never have one again. Yet perhaps it was as
well it was finished. Once at home and he at work, she would be able to
sleep at least all day....
In London Mrs. Mitcham, anxious to do the right thing, had filled
George’s different bits of china with white flowers, and the drawing-room
with its heavy black furniture looked more like a carefully kept-up
memorial than ever. She herself was very clean and spruce in her best
apron, and her face was wreathed in the proper welcoming smiles, though
she was in fact excessively nervous and embarrassed. She had always liked
Christopher, with the indulgent liking of the elderly female servant for the
irrepressible young gentleman, and she was thoroughly aware that marriage
was marriage, yet she couldn’t help turning a little red when he marched
into her mistress’s bedroom as if it were his. So it was his; but for years and
years it hadn’t been his, and for years and years before that it not only
hadn’t been his but had been poor Mr. Cumfrit’s. It was her vivid
recollection of poor Mr. Cumfrit—the times she had taken him and Mrs.
Cumfrit their morning tea into that very room, and they always so pleasant
and content together in their double bed—that gave her this feeling of shock
when she saw Christopher walking in.
Mrs. Mitcham, who was so well trained that her very thoughts were
respectful, wouldn’t have dreamed of comparing Christopher to a cuckoo,
but she had heard of the bird’s habits in regard to nests not originally its
own, and deep down in the vast dark regions of her subconsciousness,
where no training had ever set its foot and simple candour prevailed, was
the recognition of the fact that her mistress was the natural prey of cuckoos,
—first Stephen, turning her out of her original nest, now Christopher, taking
possession of this one.
She could only hope that all would be well. What wasn’t well, plain
enough for any one to see, was her mistress. Mrs. Mitcham couldn’t have
believed such a change possible in that short time. ‘It’s them honeymoons,’
she said to herself, shaking her head over her saucepans. They did no good
to a woman, she thought, not after a certain age. You had to be very strong
to put up with them at any time. No rest. No regular hours. Never knew
where you were. He looked all right, the young gentleman did, and it wasn’t
for not being happy that her mistress didn’t, for already, in the quarter of an
hour since they had arrived, Mrs. Mitcham had seen more love about in the
flat than she could remember during the whole of poor Mr. Cumfrit’s time
in it. She couldn’t help wondering what that poor gentleman would say if he
could see what was happening in his flat. He wouldn’t much like it, she was
afraid; but perhaps hardly anybody who was dead would much like what
they would see, supposing they were able to come back and look. Even
Mitcham wouldn’t. What Mitcham wouldn’t like seeing, if there was
nothing else for him to grumble about, would be how well she had got on
without him.
She carried the asparagus into the dining-room. Once again, as she
opened the door, she felt uncomfortable. How should she have thought of
knocking first? She had never had to knock at any except bedroom doors
since she had been in service. Whenever she happened to go into a room
everything would be just as nice and what you’d expect as possible: Mr.
Cumfrit at the head of the table, taking a sip of claret, Mrs. Cumfrit on his
right hand, quietly eating her toast. ‘I’ve done a good stroke of business to-
day,’ Mr. Cumfrit would be saying. ‘I’m so glad, George dear,’ Mrs.
Cumfrit would be answering. Things like that would be going on in the
room, things you’d expect. Or, in the drawing-room, Mr. Cumfrit would be
one side of the fire, Mrs. Cumfrit the other, reading out bits of the evening
paper to each other. ‘Seems to me this wretched Government doesn’t know
what it wants,’ Mr. Cumfrit would be saying. ‘It does seem so, doesn’t it,
George dear,’ Mrs. Cumfrit would be answering, quiet and ladylike.
But this....
Impossible for Mrs. Mitcham not to start and draw back when she
opened the door. She all but scattered the lovely asparagus over the carpet.
She wasn’t used—and her mistress, too—she couldn’t have believed——
‘Come in, come in, Mrs. Mitcham,’ the young gentleman called out
gaily, picking up his napkin which had fallen on to the floor. ‘We’re
married, you know. It’s all right. You signed the certificate yourself.’
Mrs. Mitcham smiled nervously. The sauce-boat rattled in its dish as she
handed it to her mistress. She wasn’t used to this sort of thing.
IV
That evening, sentimentally, they went to The Immortal Hour. In this very
place, only two months back, they had been sitting apart hardly aware of
each other, hardly more than looking at each other out of the corners of their
eyes.
‘Do you remember the night I first moved up next to you?’ Christopher
whispered.
‘Don’t I,’ she murmured.
‘Oh, Catherine—isn’t it wonderful to think we’re married!’ he
whispered.
‘Sh-sh-,’ hissed the audience, still sparse and still ferocious.
She was in bliss again. He loved her so. He had been so utterly charming
in Hertford Street, boyishly delighted with everything, filling the dull little
flat with youth, and all that youth trails with it of clouds of glory—laughter,
happiness, radiant confidence. Amazing to have this there after George,
after the quiet years since George.
By the evening she was tired, horribly tired, and knew she looked like a
ghost; but she didn’t mind as long as it was dark and he couldn’t see her
silly white face and smudged, haggard eyes. There was only one interval,
and her hat would hide her then. The Immortal Hour was such a nice dark
opera: pitch dark for ages in the first act,—so restful, so soothing.
She went sound asleep, her head against his arm. He didn’t know she
was asleep, and was thinking all the time of how they were both thinking
and feeling the same things exactly, he and she who owed each other to the
for-ever-to-be-adored Immortal Hour.
‘Darling, darling,’ he murmured, stooping and trying to kiss her at the
darkest moment. This bliss of unity with the perfect love, this end of
loneliness, this enveloping joy....
She slept profoundly.
However, she woke when the curtain went down before the second part
of the act, and those of the audience who were new to it clapped in spite of
the music going on, and those who weren’t new indignantly hissed at them,
and sat up and pulled her hat straight. It was the same funny little
extinguishing hat she used to wear at the beginning; he had specially asked
her to put it on.
‘Yes, we must be proper now,’ said Christopher, smiling at her.
‘Sh-sh-,’ hissed the outraged audience.
How familiar it all was; how happy they were. She was glad he didn’t
know she had been asleep. It was awful to have gone to sleep on such an
occasion, but then she was so appallingly tired. Never in her life had she
been tired like this. Ah, here was the love scene beginning ... she wouldn’t
go to sleep now....
Her hand slid into his; his shut tight over it; they sat close, close, thrilled
by memories, by all that the music meant to them; and in the most beautiful
part Catherine felt her thrills grow fainter and fade away and go out, and
again her head drooped against his shoulder and again she went sound
asleep.
‘Oh, I love you, love you,’ whispered Christopher, putting his arm round
her, sure her drooping head was the gesture of abandonment to irresistible
emotion.
‘Sh-sh-,’ hissed the audience.
Afterwards he wanted to take her somewhere to supper.
‘Supper?’ echoed Catherine faintly, who was dying with fatigue.
‘Yes. We must celebrate—drink the health of our home-coming,’ said
Christopher, drawing her hand through his arm and proudly walking her off
to a taxi. His wife. Marvellous. No more slipping away in the crowd and
escaping him now, thank you. ‘Let’s go somewhere where we can dance. I
shall blow up if I don’t let off steam somehow.’
‘Dance?’ echoed Catherine again, still more faintly, as she was swept up
into the taxi.
‘Do you realise we’ve never danced together once yet?’
‘But we can’t go anywhere like that in these clothes.’
That was true. He hadn’t thought of that. Well then, they would dress
properly the next night and go and dance and dance.
Catherine sat back in the seat. Dance? She hadn’t danced for years, not
since before her marriage with George—never since.
She told Christopher this, and he only laughed and said it was high time
she did dance; he adored dancing; he longed to dance with her; they would
often go.
‘Oh, Christopher,’ said Catherine, sliding close up to him, ‘the best thing
of all will be being alone together at home, you and I, in your precious
evenings. Won’t we go there now? Do we really want supper?’
‘Tired, darling?’ he asked, instantly anxious, stooping to look under her
hat.
‘Oh no—not a bit. Not in the least. Really not,’ said Catherine quickly.
‘But—our first evening—it’s so lovely at home——’
He hung out of the window and redirected the driver. ‘Yes. Of course,’
he said, taking her in his arms. ‘That’s far and away the best of all——’
And they began to whisper.
Next day he went back to work. When he left at ten o’clock Catherine
was still in bed.
‘Do you mind my not being at breakfast?’ she had asked him when Mrs.
Mitcham very gingerly beat, or, more accurately, delicately patted the gong.
Mrs. Mitcham had had some moments of painful indecision before doing
anything with the gong. It was altogether a most awkward morning for her.
She had never yet been placed in such a position. Husband and wife, of
course, and all that—she knew all that; but still it did feel awkward, and she
had a queer reluctance to rousing them—almost as if they were dangerous,
as well as embarrassing, to have loose about the flat. Yet it was breakfast
time. Orders were for nine sharp. She did finally get herself to the gong and
timidly tapped it, divided between duty and her odd reluctance to see her
mistress and the young gentleman come out of that room, to have to face
them....
‘Stay there, my darling love,’ said Christopher, smoothing the pillows
and tucking Catherine up as tenderly as if she were a baby. ‘I’ll bring you
your breakfast.’
‘I—never do get up to breakfast,’ she said, after a moment’s hesitation,
smiling at him as he bent over her—she, who had not once during the whole
of George’s time missed being down on the stroke of half-past eight to pour
out his coffee for him and kiss him good-bye on the door-mat. ‘Good-bye,
little woman,’ George used to say, waving to her before the lift engulfed
him. In those days good husbands of good wives frequently called them
little women.
Here now was her chance. She would establish a custom that might save
her. And if she never had got up to breakfast it wouldn’t worry Christopher
that she never did, and he wouldn’t, frowning with concerned perplexity,
ask her searching questions as to being not well. So, by sleeping on into the
mornings after he had gone to work, she might catch up with rest and dodge
those horrid furrows exhaustion was dragging down her face.
So the habit was started, and Mrs. Mitcham learnt not to expect to see
her till lunch-time. Sometimes she even slept later, and once or twice stayed
in bed all day, not getting up till just in time to dress for dinner. This,
however, only happened during the first two or three weeks. As time went
on Mrs. Mitcham began to be able to count on her mistress’s having her
bath at twelve o’clock and being ready by one.
Mrs. Mitcham was all for her resting and taking care of herself, for she
was much attached to Catherine, but she couldn’t help feeling—she didn’t
permit herself to think it, but she couldn’t help feeling—that there was
something unbecoming in this turning of day into night. There was plenty
of night, Mrs. Mitcham thought, for those who chose to take it, but of
course if——
Mrs. Mitcham, folding up her mistress’s garments, shook her head. And
the garments too—she shook her head at them. Such things had not hitherto
been part of Mrs. Cumfrit’s outfit. Good things she had had, as good of their
kind as one would wish to see,—lawn, silk, fine embroideries,—but never
what Mrs. Mitcham called flimsies. These were flimsy, and not only flimsy
but transparent. Every time Mrs. Mitcham saw them she was shocked
afresh. She couldn’t get used to them. Mrs. Cumfrit—she corrected herself,
and said Mrs. Monckton—had gone out and bought them the first afternoon
of her return from the Isle of Wight; and she so careful about coals, and
turning the electric light out. There were six nightgowns that you could pull
through a wedding-ring, they went so into nothing. Chiffon nightgowns.
Different colours. Pink, lemon-colour, and so on; and all of them you could
see through as plain as daylight. It was a mercy, thought Mrs. Mitcham, that
it was dark at night. She, who prided herself on Catherine and had always
thought her the ideal of what a lady should be, was much perturbed by these
nightgowns. And the bathroom too—such a litter there now of scented
dusting powder, and scented crystals, and flagons of coloured liquid that
smelt good but improper, thought Mrs. Mitcham, furtively sniffing; what
would poor Mr. Cumfrit say to his bathroom now, he who had never had a
thing in it but a big sponge and a piece of Pears’ soap?
It was after the visit of the Fanshawes that Mrs. Mitcham first found a
lip-stick on Catherine’s dressing-table. She was immensely upset. No lady
she had had to do with had ever had such a thing on her dressing-table.
Powder was different, because one needed powder sometimes for other
things besides one’s face, and also one powdered babies, and they, poor
lambs, couldn’t be suspected of wanting to appear different from what God
had made them. But a lip-stick! Red stuff. What actresses put on, and those
who were no better than they should be. Her mistress and a lip-stick—what
would Miss Virginia say?
The Fanshawes, who were the immediate cause of the buying of the lip-
stick, came to tea the second Sunday after the end of the honeymoon,—
Ned, his mother and sister. They had been extraordinarily taken aback by
Catherine’s appearance. The flat rang with their exclamations and laments.
Catherine, who had been looking sixteen when they last saw her, Catherine
the bright-eyed, the quick of movement, Catherine with her lovely skin and
unruffled brow—they couldn’t get over it.
Christopher had gone for a walk in the Park after lunch, straining at his
leash, angry at being kept in London this beautiful afternoon because the
Fanshawes insisted on coming and thrusting their inquisitive noses where
they weren’t wanted, and he hadn’t got back when they arrived, so that
Catherine had them to herself at first.
‘Damn those women,’ he had remarked when, after persistent
telephoning and letters and impassioned inquiries as to what had happened
to her and when they might come and see her, Catherine had felt she had
better face it and wrote and told them she was married and asked them to
tea that Sunday to meet her husband. And when he heard that it wasn’t only
women but Ned too, he damned him particularly, on the ground that he had
a silly nose and wore a fur rug up to his chin; and, expressing extreme
disgust at being kept in on his and Catherine’s only real afternoon by a
blighter like that, he went off for a quick turn in the Park, promising
faithfully to be back in time.
He wasn’t; and the Fanshawes got there first, and the flat was echoing
with exclamations when he opened the door.
Catherine was sitting on the sofa, wedged between the two female
Fanshawes, whose arms encircled her and whose free hands stroked her,
and that worm Ned was looking on from his, Christopher’s, special chair.
‘I’ve had influenza—that’s why,’ Catherine was lying as he came in.
‘But fancy not telling us you were married! Fancy not telling us a word!’
‘Of course it’s what we’ve been dying to have happen for ages—isn’t it,
Ned?’
‘You sweet little thing, we’re so delighted—aren’t we, Ned?’
‘Do tell us all about it. Isn’t he off his head with happiness? How
pleased Virginia must be——’
‘So nice for her to have a father again——’
‘Are they devoted to each other?’
‘Have you been down to Chickover with him yet?’
‘We’re simply aching to see him——’
And there in the doorway he stood.
‘Here is Christopher,’ said Catherine, flushing and half getting up.
They all turned their heads. For a moment nobody spoke. He advanced
on them with outstretched hand, doing his best to smile broadly, to be the
welcoming host.
That young man. That boy. The boy they had found with Catherine one
day, who had rushed out the minute they came in, and Catherine had
laughed when they asked who on earth he was, and said all she knew about
him was that he was certainly mad. That fellow. The youth who had glared
through the window of the car and almost shook his fist....
The Fanshawes couldn’t speak. They couldn’t move either. They were
stunned.
V
At Chickover there had been the most painful consultations between
Stephen and Mrs. Colquhoun as to the best thing to do under the deplorable
circumstances. Should they or should they not tell Virginia? Could they,
indeed, help telling her? Not all, of course; she must never be told all. The
night spent somewhere between Chickover and London—they both felt that
the entire stretch of country between those two points was from henceforth
polluted—the night that made the scandalous marriage a necessity, must be
kept from Virginia for ever. But it became clear after a week that she must
be told something, if only to account for her not hearing from her mother.
Stephen couldn’t bring himself to let her have the letters. They came at
first, as he had expected, one after the other and all very thick. He
wondered, turning them over in his hand, whether it wasn’t his duty to open
them, but he resisted the strong leaning towards his duty that lifelong
practice in doing it had induced in him, and took the more dignified course
of sending them back unopened. Much more punitive too, he felt,—leaving
the wretched woman completely in the dark as to what was happening at
Chickover and what Virginia was feeling.
Then, when the letters at last left off coming, he watched for telegrams;
he rather expected telegrams.
None came.
Then he was on the look-out for an unannounced arrival; he quite
thought there would be one.
Nothing happened. Just silence.
At the end of the week Virginia said, ‘I can’t think why mother doesn’t
write’—and began to look worried, and write letters herself.
Stephen took them out of the box in the hall and burnt them. ‘Painful,
painful necessities,’ he said to Mrs. Colquhoun; for this letter business went
against the grain—the gentleman grain, he told his mother, who hardly left
him, comforting and advising him as best she could.
At the end of another week Virginia sent a telegram, or rather was going
to send it but was stopped by Stephen. Clearly she must be told something.
She had said, while writing it: ‘If I don’t get an answer to this I shall go up
to London myself and see if anything is wrong.’
‘Poor child, poor child,’ murmured Mrs. Colquhoun, the moment for
enlightenment having manifestly come. ‘Would you like me to be with
you?’ she whispered in Stephen’s ear.
‘Better not, I think,’ he whispered back.
Alone with Virginia he took her on his knee. She was holding the
telegram she had just written, and was in a hurry to go and send it off.
‘Yes, Stephen—what is it?’ she asked, fretted at being held back, and
worried by this strange silence of her mother’s to the point of being unlike
herself.
‘I am but a clumsy creature,’ he began, overwhelmed by the thought of
the blow about to be delivered—and delivered by his hand, too, his own
loving hand.
He laid his head on her breast, his arms round her, as she sat on his knee.
This beginning made Virginia still more uneasy; Stephen had never
called himself a clumsy creature before. ‘What is it, darling?’ she asked,
very anxious.
‘What is it not,’ groaned Stephen, holding her tight. To think it was he,
he who so deeply loved her....
‘Oh, Stephen’—Virginia was thoroughly frightened—‘mother?’
‘Yes. Yes. Yours. And Virginia, my loved wife,’ he said, raising his head
and looking at her, ‘believe me I had rather, to spare you, it were mine.’
Virginia sat like a stone. Her face was stiff and set. The worst had
happened, then. Her little mother, her own sweet little mother, to whom she
had been unkind, unloving, and who had never once failed in kindness and
love to her, was dead.
‘She is dead,’ said Virginia, in a voice so toneless that it sounded
indifferent.
‘How much better,’ thought Stephen, ‘for everybody as well as herself if
she were.’
Aloud he said, his face buried in Virginia’s bosom. ‘No. She is not dead.
Quite the contrary. She is remarrying.’
And as Virginia said nothing, for her breath was taken away by these
blows and counter-blows, he went on: ‘Darling, I would have spared you if
I could. I have tried to spare you. I have tried all these days to find some
way of keeping it from you. Indeed, indeed I have tried——’
‘But, Stephen—why? Why shouldn’t mother marry again?’ asked
Virginia, with the irritability natural to people who have been frightened
without cause, but so unusual in her that Stephen could only account for it
by her physical condition. ‘I think it very strange of her not to tell me, but
why shouldn’t she remarry? Now there’ll be somebody to take care of her.
I’m glad.’
‘Darling——’
He pressed his face still closer to her bosom. He wished he could hide it
there for ever.
‘But I do think,’ said Virginia, reaction against her mother setting in now
she knew she wasn’t dead, as it had set in the day she saw her trotting safe
and sound up the avenue when she had been torn by fears of an accident, ‘I
do think she might have told me. I do think that.’
Her voice had tears in it. She strangled them, and held herself up very
straight, offering no real hospitality to Stephen’s head. She was deeply
wounded.
‘Ah, but there are some things one doesn’t tell,’ said Stephen.
He was miserable. He would have given at least half Chickover to be
able to spare her.
‘Remarrying isn’t one of them,’ she said; and for the first time he caught
a glimpse of another Virginia, a Virginia who perhaps, ten years ahead,
might argue.
He raised his head from her bosom and looked at her again.
‘I know what I am talking about, my child,’ he said. ‘This remarrying is.
She is marrying the young man with whom she motored up to London. You
saw him yourself. Perhaps you will now agree that there are some things
one does not willingly tell one’s daughter.’
Virginia stared at him a moment, her eyes very wide open. Then, without
speaking a word, she got up off his knee and walked over to a window and
stood at it with her back to him.
How strange of her, he thought. What a strange way of meeting trouble
—to go away from him like that, to turn from the love that longed to help
her.
He didn’t know what to do or say next. He sat watching her in the utmost
perplexity. His Virginia, getting up off his knee, withdrawing herself from
his loving arms——
‘Yes,’ she said after a long silence. ‘I can imagine there may be such
things. But I don’t think’—she turned and faced him—‘this is one of them.’
He got on to his feet and went towards her, his arms outstretched.
‘My darling, my wounded darling,’ he cried, all understanding and
pitifulness, ‘you are generous and young——’
‘So is Mr. Monckton,’ was her unexpected answer.
Really it was like a blow in the face. It stopped him short. It must be her
condition. But all that day the attitude continued, the strange, almost defiant
attitude, and Stephen could only go into his study and pray that her heart
might be softened, and her eyes opened to see things as they truly were.
Such a grave misfortune—she did not of course know how grave, how
terrible it was—the first in their married life, and to take it this way,
hardening her heart against the sympathy and understanding he and his
mother offered her in such boundless measure, and persisting, with an
obstinacy he wouldn’t have dreamed her capable of, in upholding what her
mother had done! True she said very little, but that little was all obstinate.
She was quite unlike her usual self to his mother, too, whose one thought
was to comfort, and would not admit that there was anything to be
comforted about. And when that night he got into bed, and drew her to his
heart in the exquisite contact of the body that had always till then soothed
every trouble of the spirit, and she came apparently willingly, and clung to
him, and was his own dear wife as he thought, and, happily sure of this, he
whispered that he hoped she had remembered to pray for her poor mother, it
was a grievous shock to feel her shrink away from him and hear her say she
hadn’t—not more than usual, not more than her childhood’s ‘God bless my
mummy,’ and most grievous to have her ask him, just as if they weren’t in
bed but downstairs conversing in their clothes, just as if they weren’t in the
sacred place and at the sacred moment never yet profaned by talk of
anything but love, why he really thought it so dreadful for her mother to
marry again.
‘Do you not see it is terrible to marry some one young enough to be your
son?’ he had asked sternly—he couldn’t have believed he would ever have
to be stern with his own love in such a place, at such an hour.
And she had answered: ‘But is it any more terrible than marrying some
one young enough to be your daughter?’
Virginia had answered that. His Virginia. In bed. In his very arms.
VI
Stephen was completely crushed by this. It was like the things children say,
unanswerable in its simple rightness, and yet, the grown-up world being
what it is, all wrong.
Virginia was nearer the fount of truth than he was. Where he stood, thirty
years further from it, its waters naturally were muddier; but there they were,
and had to be dealt with including their mud, and not as if they remained for
ever, as she so near the source supposed, crystal clear.
On the other hand he couldn’t do without his wife. He owed her
everything, and above all he owed her his return to youth. She had come
and released him from the darkening prison of deepening middle age. He
worshipped her more than he knew. For instance, up to this he hadn’t
known he was wax in her hands, he had imagined he led, guided, was in
absolute authority; now suddenly he knew he was wax. When his mother-
in-law had dared compare his marriage with hers, the thing had been the
deadliest outrage. Virginia pointed her finger at it, and instead of being
outraged he was crushed—crushed by the truth as she saw it, crushed by the
knowledge that in her clear young eyes he hadn’t, in his condemnation of
her mother’s action, a leg to stand on.
Yet how many legs did Stephen not know he really had to stand on.
Everybody, except children like Virginia, inexperienced and new, would
agree that the difference between the two cases was such that one was
accepted as natural and the other with derision. But Virginia said there was
no difference—he had an uneasy feeling that Christ might have said so too
—and declared that if her mother’s marrying some one so much younger
was terrible, then her marrying Stephen was terrible; more terrible, because
there were actually eight years greater difference in their case.
All this she said that first tragic night in bed, and inquired, when she had
finished, what was to be done about it.
He was wax. Not immediately; not for some black weeks of agonising
separation, of days spent avoiding each other, of nights spent with their
backs turned; but inevitably, before her stubbornness, he melted. She had
too many of the necessities of his life in her hands. To be out of harmony
with Virginia was worse, far worse—he shuddered as he admitted it, but
there it was—than being out of harmony with God.
And Virginia, though she kept up her stout exterior and went doggedly
through these painful days of April and May—the estrangement lasted all
that time—was most wretchedly unhappy. What was she to make of all
this? In her heart she was as much shocked as Stephen. But how could she
not stand up for and defend her mother? How could she deny her own
blood? Deeply she resented having to defend her mother; it shattered the
foundations of the whole of her childhood’s faith. Was there ever anything
more miserable, she thought, than to love some one and be horrified at them
at the same time? She who ought to have been putting up her feet and
resting more diligently than ever—‘The fifth month, dear child,’ Mrs.
Colquhoun anxiously reminded her, ‘we are in the fifth month now, you
know, and have to be most careful’—walked ceaselessly instead in the
garden, up and down, up and down, in all those paths where she was least
likely to be found by her mother-in-law, trying her hardest to see clear, to
think right, groping round for some way to get back to Stephen while at the
same time not deserting her mother, to get back to his arms, to his heart, to
the unclouded love without which she felt she couldn’t live.
‘You know, Virginia dearest,’ said Mrs. Colquhoun at last, whose only
wish was to console and be confided in, and who was much upset by this
marked and morbid avoidance, ‘you have nothing to be ashamed of.’
‘Has anybody?’ Virginia asked, stopping short in the middle of the path
she had been waylaid in, and looking her mother-in-law squarely in the
eyes.
‘That wretched, wretched mother of hers,’ groaned Mrs. Colquhoun to
Stephen, describing this little scene to him and how uncomfortable and hurt
she had been by the poor child’s want of frankness. ‘What misery she has
brought on us all.’
‘Nevertheless,’ said Stephen, looking up wearily from the sermon he
was trying to write—his head, his heart, every part of him seemed to ache
—‘we must countenance her.’
‘Countenance? Countenance her? Do you mean behave as if we
approved of her?’
‘I do, mother. I have been thinking it over very seriously. Virginia’s
health, and with her health her child’s health, is at stake. She——’ his voice
faltered, for he was most miserable—‘she weeps at night. She—she weeps
when she thinks I am asleep. If I try to console her it—it becomes
heartbreaking.’
He turned his face away and bent over the manuscript. Tears had come
out of his own eyes, and were wetting his face. Impossible for any one to
conceive the torments of his nights in bed with his beloved one and
estranged from her. That turning of backs, that cold space between their two
unhappy bodies....
‘Wretched, wretched woman,’ said Mrs. Colquhoun again, more bitterly
than ever, for she saw her son’s tears.
‘Yes. But we must think of Virginia.’
‘Has she been writing to her mother?’
‘I am sure she has not. She would do nothing without my knowledge.’
He dug the nib of his pen into the blotting-paper.
‘My wife is the soul of loyalty and straightness——’ he began, but his
voice quivering uncontrollably at the mention of her dear qualities, he broke
off.
‘Yes indeed, Stephen. Indeed I know it. She is the dearest child. Only at
these times a woman isn’t quite herself, and Virginia, I can see, has got into
a curious morbid state——’
‘I should leave her alone, mother,’ interrupted Stephen, his head bent so
that she couldn’t see his face.
Mrs. Colquhoun was hurt. All her affection and sympathy being thrown
back, as it were, at her,—told straight by her son, to whose welfare she had
devoted the whole of her life, that she was taking the wrong line with

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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