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Discovering Behavioral Neuroscience
AN INTRODUCTION TO BIOLOGICAL PSYCHOLOGY
Fourth Edition
Laura A. Freberg
California Polytechnic State University,
San Luis Obispo
,
-
·-
# CENGAGE
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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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TO MY FAMILY
Roger, Kristin, Karen, and Karla
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Editorial review has deemed that any suppressed concent does no< materially affect the overall learning experience. Ccngagc Leaming reserves the rig.ht 10 remove additional contcn1 at any time if subsequent righ[S restrictions require ii.
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 ofMind 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, 0. Ivar Lovaas,
Larry Butcher, Jackson Beatty, John Libeskind, Donald Novin, Frank Krasne, and
F. Nowell Jones. She was privileged to study neuroanatomy with Arnold Scheibe!, 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
20 1 1, 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-US.
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 ofZelda.
iv
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 1-1
Subject Index/Glossary 1-12
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.
vi
Preface xix
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
Interim Summary 1.2 21
Research Ethics in Behavioral Neuroscience 23
Human Participant Guidelines 23
Animal Subjects Guidelines 24
Interim Summary 1 .3 25
Chapter Review • T hought 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
Functional Neuroanato my 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
Copyright 2019 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, In whole or i n part. WCN 02-200-202
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
Chapter Review• Thought Questions• Key Terms 65
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
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
Leaming reserves lhc right 10 remove additionaJ contcnl at any time if subsequent rights restrictions require ii.
Editorial review has deemed that any suppressed content docs not materially affect the overall learning experience. Ccngagc
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
II Psycho pharmacology
Neurotransmitters, Neuromodulators, and Neurohormones

Identifying Neurochemicals 109
107
108
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
Stimulants133
Opioids 137
Cannabis 138
LSD 139
Alcohol 141
Chapter Review • T hought Questions • Key Terms 144
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
The Genetic Bases of Behavior 146

From Genome to Trait 146
Sources of Genetic Variability 148
Heritability 151
Epigenetics 153
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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
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
Copyright 2019 Cengage Learning. All Rights Reserved. May not be copied, scanned, or duplicated, In whole or in part. WCN 02·200-202
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
Audition 226
Sound as a Stimulus 226
The Structure and Function of the Auditory System 229
Auditory Perception
235
Hearing Disorders 23 7
The Body Senses 241
The Vestibular System 241
Touch243
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
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 Cerebellum283
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
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
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
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
Interim Summary 1 1.1 387
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
Interim Summary 11 .3 412
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
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
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
• Emotion, Aggressio n, and Stress
Emotion 490
The Evolution and Adaptive Benefits of Emotion
489
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
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
Editorial review has deemed that any suppressed contentdocs not materially affect the overall learning experience. Ccngagc Leaming reserves lhc right 10 remove additionaJ contcnl at any time if subsequent rights restrictions require ii.
Contents xvii
Recovery and Treatment in Neurocognitive Disorders 550

Plasticity and Recovery 550
Cognitive Reserve 551
Rehabilitation for Neurocognitive Disorders 552
Chapter Review • Thought Questions • Key Terms 554
BEHAVIORAL NEUROSCIENCE GOES TO WORK: Preparing to Be a Clinical

Neuropsychologist 525
THINKING ETHICALLY: Apoe E4 in Younger Adults 529
531
BUILDING BETTER HEALTH: Recognizing the Signs of a Stroke
CONNECTING TO RESEARCH: Stanley Prusiner and the Prion 537
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
Interim Summary 1 6. 1 566
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 (MOD) 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 P TSD 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
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toista taiteenhaaraa, maalausta, josta ei sitten tullut kuitenkaan
mitään.
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Olimme ihan samaa mieltä, varsinkin kun tahdoimme päästä taas

käsiksi näyttämötaiteeseen. Tiedustelimme senvuoksi taiteilijattaren
ensimmäistä näyttämöllistä esitystä ja sen tekemää vaikutusta
häneen.
Eräässä tuttavaperheessä lapset olivat keskenään leikkineet ja

näytelleet, ruotsinkielellä, Z. Topeliuksen pienen kappaleen "Ole
armelias köyhille", jossa taiteilijattaremme 15-vuotiaana oli esittänyt
Brita muorin osan. Hauskaahan se etupäässä oli ollut, mutta kovin
vakavasti se oli otettu, täydellisellä hartaudella ja antaumuksella
esitettiin esitettävä.
— Ei siis tämä johtanut ajatusta teatteriin, synnyttänyt edes

hämärää aavistusta tulevasta toimialasta?
— Eipä tietenkään, koska vuotta myöhemmin koulusta erottuani

rupesin erään suuren tamperelaisen liikkeen kassanhoitajattareksi ja
siinä toimessa sitten pysyin kaksi vuotta.
— Entäs sitten kahden vuoden kuluttua?
— Läksin silloin teatteriin.
— Ajatus siis kypsyi kassanhoitajattarena.
— Silloinpa tietenkin.
— Esiinnyittekö Tampereella missään seuranäytelmässä?
— Esiinnyin, kuten nuoret ihmiset yleensä esiintyvät heitä sellaisiin

tilaisuuksiin avustamaan pyydettäessä. Ensimmäinen osani tältä
ajalta oli Alman osa kappaleessa "Tätä nykyä." Tekijän salanimeä en
muista, mutta älkää kirjoituksessanne mainitko kuulemaanne oikeata
nimeä, sillä en tiedä lainkaan, haluaako hän esiintyä julkisuudessa.
Suomalainen teatteri kyllä esitti näytelmän aikoinaan, ja on se
painettuna kirjanakin ilmestynyt. Nämä esiintymiset eivät suinkaan
vaikuttaneet ratkaisevasti uranvaihdokseeni, päätös siitä kypsyi
hiljalleen ensimmäisen siellä viettämäni vuoden kuluessa, ja
luultavasti ikävän ja työalaani kyllästymisen aiheuttamana.
Seitsentoistavuotiaana kirjoitin sitten äidilleni, että ehdottomasti tulen
pyrkimään teatteriin ja että minua on turha kieltää siitä, sillä kaikesta
huolimatta lähden kumminkin.
— Halu oli siis järkähtämätön ja vakava joka tapauksessa.
— Siitäpä se näyttää.
— Niin, siitä ja sen syistähän emme tosiaankaan puhu.

Kyselisimme kuitenkin unelmistanne ja odotuksistanne tälle uudelle
uralle astuessanne, tiedustelisimme noin vain ohimennen,
sivuuttaaksemme tuon arkaluontoisen syntyjen syvien tutkimisen.
— Kahdeksantoistavuotiaan unelmathan ovat aina loistavat ja
valoisat ja korkealle sitä tietysti taiteensa asetti, korkeammalle
kaikkea muuta maallista. Muistan hyvin, kuinka syvästi säälin
jokaista ihmistä, joka vain puhui tavallisista arkiasioista; ne raukat
eivät mitään tiedä oikeasta, todellisesta elämästä, mietiskelin
mielessäni. Nyt myöhemmin on sitten kyllä itsekin saanut paljon
arkiasioita aprikoida, ja hyvinhän sitä siihenkin perehtyy ja tottuu.
— Oliko polkunne tällä uudella uralla sitten aivan ohdakkeeton?

uskalsimme kysäistä.
— Eipä suinkaan, toista oli silloin kuin nykyään. Nykyisellä

nuorella polvella on ihan liian hyvät päivät. Teatterikoulussa saadaan
valmis näyttämötottumus ja tekotapa, ei tarvitse paljon itse
ponnistella, vähälahjaisimmatkin voivat jo heti alussa esiintyä
jonkinlaisella valmiudella ja suoriutua tehtävistään melkoisella
menestyksellä. Me saimme väliin vuosikaudet olla statisteina ja
vuorotellen hoitaa kuiskaajan tointa. Kuiskaajana oli kurjinta. Halu
paloi näyttämölle, mutta kohtalo viskasi näyttämön alle. Muistan
hyvin, kuinka kerran Turussa vierailtaessa näyteltiin "Kylän heittiö"
nimistä laulukappaletta. Minulla oli tässä ilossa kuiskaajan onneton
osa. Joku edellinen uhri, joka nyt luultavasti liikkui jo lavalla, oli
piirtänyt suorin kirjaimin luukun eteen sanat: Sufflera, din fan
(Kuiskaa, senkin piru)! Tämä tietysti ärsytti yhä enemmän jo
ennestään ärtynyttä mieltä, ja laulukohtauksien aikana painuin aina
kuiskaajan koppiin ja itkin kiukusta ja katkeruudesta. Varsinaisia
vastuksia minulla ei kuitenkaan ollut, ei suuria suruja eikä
pettymyksiä, olivathan kaikki muutkin ennen saaneet samaa kokea
kuin minäkin, ei siis mitään valittamista. Ja siitä huolimatta oli koko
teatterissaoloni minulle kuin uhkaava, pakottava ruoska.
— Mistä se sitten johtui?
— Siitä ankarasta kunnioituksesta työtä kohtaan, jonka minulle

antoi neiti Avellan, joka sattui joutumaan opettajakseni. Siihen
aikaan sai kukin vanhempi näyttelijä jonkun vasta-alkajan
opastettavakseen, ja minä jouduin nauttimaan, osaksi hyvien
tuttavienkin toimesta, neiti Avellanin nerokasta ja sytyttävää
ohjausta. Häntä saan kaikesta kiittää. Ensinnäkin jo ehkä teatteriin
pääsystäni. Tohtori Bergbom ei näet kiinnittänyt mitään erikoista
huomiota minuun, mutta neiti Avellan uhkasi, että ei lukisi kenenkään
alottelevan kanssa vastaisuudessa, ellei "Finneä oteta teatteriin". Ja
sitten on hänen ansiotaan koko se kunnioitus taidetta ja vakavaa
työskentelyä kohtaan, mikä minussa mahdollisesti on ollut ja vieläkin
on.
— Tämä mainitsemanne neiti Avellanin lausuntoko aiheutti sitten

pääsynne vakinaiseksi näyttelijäksi?
— Ehkäpä välillisesti, mutta oikeastaan sen aiheutti suoranainen

sattuma. Muistattehan Molièren "Luulosairaassa" pienen Louison
nimisen tytön, joka m.m. saa vitsaa isältään? No niin, minun
tehtävänäni oli harjoittaa siihen osaan pieni kansakoulutyttö, joka
oppikin sen hyvin, mutta oli niin kaino, että ei saanut sanaakaan
suustaan ratkaisevana hetkenä. Harmistuneena toht. Bergbom
määräsi minut näyttelemään oppilaani osaa, ja siinä onnistuinkin niin
hyvin, että sain muitta mutkitta näyttelijävälikirjan, tarvitsematta
antaa koenäytäntöä johtokunnalle. Tämä tapahtui kevättalvella 1886,
siis oltuani jo kokonaisen vuoden teatterin palveluksessa.
— Silloinko esiinnyitte ensi kerran vakinaisen teatterin lavalla?

— En, olin jo hiukan aikaisemmin näytellyt Sinipiikaa "Kullervossa"
Kivi-juhlassa tammikuun 1 päivänä 1886. Se oli samalla minulle
suuri ilonpäivä, sillä silloin oli nimeni ensi kerran painettu ohjelmaan.
Kysyimme sitten taiteilijattaren ensimmäistä suurta iloa taiteellisen

voiton ja menestyksen johdosta.
Se oli tapahtunut hänen esittäessään Boriskan osaa samassa

"Kylän heittiössä", jonka kappaleen kuiskaajana hän kerran Turussa
oli niin katkerasti itkenyt.
Kaarle Halme.
— Aiotte siis lopettaa sarjanne ja tehdä minusta viimeisen miehen,

virkkoi Kaarle Halme häneltä haastattelua pyytäessämme. Kiitos
vain kunniasta! Mutta olettehan jo haastatellut niin monta suuruutta,
että mitäs tällaisista pojista…
— Poika kuin poika — ja mitäs turhista masentua, voihan sitä

miehestä vastaisuudessa vielä suuruuskin sukeutua, kuten kaikista
muistakin… Ensimmäiset tulevat viimeisiksi ja viimeiset
ensimmäisiksi… Mutta tosissa puhuen olen valinnut juuri teidät
viimeiseksi senvuoksi, että voisin lopettaa sarjan johonkin
suurempaan, merkitsevämpään pisteeseen, jollaisella sen
alotinkin…
— Minunko siis pitäisi ruveta menneitä miettimään; sehän on kuin

testamentin tekoa, kuolemaa. Olen niin keskellä työtäni, näen kaiken
vielä niin edessäni, etten tiedä mennyttä olleeksi, en mitään tähän
saakka tapahtuneeksi.
— Ja kuitenkin on niin paljon tapahtunut.
— Mitä vielä. Kaikki kävi kuin itsestään, 23-vuotiaana menin

Suomalaiseen teatteriin…
— Miten sinne jouduitte?
— Menin vain, kuten jo sanoin. Olin siihen aikaan Turun

lääninhallituksessa, jossa sain toimen v. 1883. Teatteri vieraili
kaupungissa syksyllä 1887, minä menin tarjolle ja kelpasin.
— Muitta mutkittako?
— Täytyi tietysti ensin esiintyä Turussa ja antaa vielä koenäytäntö

johtokunnalle Helsingissä, sitten oli kaikki sitä myöten valmista.
— Missä kappaleessa ensin esiinnyitte Turussa?
— Näyttelin Marokon prinssin osan Shakespearen Venezian

kauppiaassa, Portian erään kosijan. Katsomossa istuivat kaikki
virkatoverini virnistellen, mikä minua hermostutti niin, että
ratkaisevassa silmänräpäyksessä unohdin koko osani. Kuiskaaja
pani parastaan. Portia, rouva Rautio, koetti hänkin supattaa jotakin
minulle, mutta mikään ei auttanut, ennenkuin ennätin rauhoittua ja
itse muistaa vuorosanani. No niin, otin sitten viransijaisen kahdeksi
viikoksi, lääninsihteeri Rydman, jonka täydellisessä suosiossa olin,
antoi minulle oikeuden olla poissa kokonaista 2 vuotta paikkaani
menettämättä, ja suori minut matkalle Helsinkiin, vaikka ei erittäin
kehottanut teatteriin liittymään, mutta ei tahtonut estääkään.
— No, sille Helsinginmatkalle jäitte?
— Sille jäin, aluksi 100 markan kuukausipalkalla, joka ei vieläkään

niin ruhtinaallinen ole. Silloisista virkatovereistani istuu usea nyt
20000 vuosituloilla, jotka kai minullakin olisi, ellen olisi antautunut
tuolle epäkiitolliselle taiteilijauralle. Olinhan jo nuoruudestani
huolimatta ollut Liedon nimismiehen sijaisena; alku oli siis hyvä.
—. Minkä osan esititte sitten johtokunnalle Helsingissä?
— Daniel Hjortin. Heti esityksen päätyttyä nousi Jaakko Forsman

paikaltaan, muutoin vanha kotipitäjäläinen, Hämeenkyrön poika, ja
lausui: "Tervetuloa Helsinkiin!" Tiesin sen kohtaloni ratkaisuksi, ja
heti sitten sainkin välikirjan.
— Olitte tietysti näytellyt jo usein ennen teatteriin liittymistänne.

Minkä ikäisenä ja missä osassa ensi kerran eläissänne esiinnyitte?
— Hämeenkyrössä esiinnyin ensi kerran ja olin silloin

kymmenvuotias. Joku vanhempi koulupoika oli kirjoittanut Saul
nimisen kappaleen, jonka sai myös pitäjäläisten suureksi
ihastukseksi lavalle, ja pisti sievät rahat taskuunsa. Tässä
kappaleessa näyttelin Davidin osan. Muistan, että minulla oli m.m.
pitkä yksinpuhelu, jonka tahdoin piirtää hiilellä esiripun yläosaan,
mukavuutta rakastava kun olen aina vähin ollut, mutta siihen eivät
muut suostuneet.
— Tästäkö sitten teatteri-intonne alkoi, vai oliko se jo ennenkin

ilmennyt?
— Ei ennen, eikä tästäkään. Pääharrastuksenani oli

kirjoitteleminen, joka ei vieläkään ole sammunut, kuten tiedätte.
Turunkin sanomalehtien alakertoja täyttelin novelleilla, joista ei
maksettu paljon mitään.
— Harrastitte näyttämötaidetta tietysti Turussakin?
— Harrastinpa tietystikin, ja ihan teatterin ohjaajana. Nämä puuhat

saivat alkunsa siten, että muuan Vankka niminen toveri oli
puutteessa ja hänen hyväkseen panimme toimeen pienet
näyttäjäiset. Useita muitakin köyhiä tovereita autettiin siten. Niin,
ensi talvenahan vietän teatterinjohtajajuhlaani, jolloin näyttämölle
asettamieni ja ohjaamieni kappaleiden luku nousee sataan, Turun
näytännöt niihin luettuina.
— Kyselisin vielä ensimmäisiä näyttämötaiteellisia

menestyksiänne ja voittojanne.
— Ensimmäinen suurempi merkkitapaus taiteessani oli tohtori

Rellingin osa Ibsenin Villisorsassa. Silloin aloin hallita tekotapaani ja
järkiperäisesti eritellä osiani. Toinen voittoni oli Ahman osa Santeri
Ivalon Juho Vesaisessa, jolloin liitin pysyväisesti ohjelmistooni
kotimaisten aiheitten esittämisen. Kenraalikuvernööri Mörnerin osa
Arvid Järnefeltin Samuel Croëlissa merkitsi sitten viimein valmiutta
taiteessani. Sitä esitystä oli muun muassa katsomassa kirjailija
Birger Mörner, joka näytännön jälkeen tuli kiittämään minua siitä, että
oli saanut nähdä sukunsa kantaisää niin hienosti ja sielukkaasti
esitettävän. — Ja nyt kai pääsen rauhaan tärkeimmät tiedot
saatuanne? lopetti taiteilija.
Mikä kumartaen myönnettiin.

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Discovering Behavioral Neuroscience:

Discovering behavioral neuroscience: an introduction to

Introduction to Research Methods in Psychology, 4th ed

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An Introduction to Mechanical Engineering (4th Ed.) 4th

Encyclopedia of Behavioral Neuroscience, 2nd Edition

The Science of Psychology: An Appreciative View 5th

Behavioral Neuroscience 9th Edition S. Mark Breedlove

(Original PDF) Chemistry: An Introduction to General,

Discovering Behavioral Neuroscience: © 2019, 2016 Cengage Learning, Inc.

An Introduction to Biological Psychology.

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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

Name Index 1-1

Subject Index/Glossary 1-12

Neuroscience as an Interdisciplinary Field 2

CONNECTING TO RESEARCH: Thinking about Your Food as Healthy or

Functional Neuroanato my and the Evolution

Anatomical Directions and Planes of Section 28

The Brain's Blood Supply 34

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

Glia and Neurons 68

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

Neurotransmitters, Neuromodulators, and Neurohormones

Types of Neurochemicals 109

CONNECTING TO RESEARCH: Otto Loewi and "Vagus Stuff" 110

The Genetic Bases of Behavior 146

Building a Brain 158

Chapter Review• Thought Questions• Key Terms 181

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

From Sensation to Perception 184

BUILDING BETTER HEALTH: Earbuds and Hearing Loss 238

Types of Muscles 266

Motor Systems of the Brain 281

THINKING ETHICALLY: Gene Doping for Strength 271

Homeostasis and Motivation 302

Reward Pathways 333

BUILDING BETTER HEALTH: Understanding the Benefits of Fever 306

Sexual Behavior 337

Sexual Development 338

BEHAVIORAL NEUROSCIENCE GOES TO WORK: How Do Therapists Treat Gender

CONNECTING TO RESEARCH: A Composite Scale of Morningness 377

Categorizing Learning and Memory 416

Systems Supporting Classical Conditioning 436

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