Physiology of Behavior, Global Edition

Neil Carlson
Visit to download the full and correct content document:
https://ebookmass.com/product/physiology-of-behavior-global-edition-neil-carlson/
 This is a special edition of an established title widely used by colleges and
GLOBAL universities throughout the world. Pearson published this exclusive edition
for the benefit of students outside the United States and Canada. If you
GLOBAL
EDITION purchased this book within the United States or Canada, you should be aware EDITION

EDITION
GLOB AL
that it has been imported without the approval of the Publisher or Author.

Physiology of Behavior
The thirteenth edition of Physiology of Behavior offers a comprehensive and systematic approach
to behavioral neuroscience. Neil Carlson and Melissa Birkett draw upon their experience of
teaching and working with students to create this accessible portrait of the dynamic interaction
between biology and behavior. The textbook presents not only the advances made in research
methods but also the discoveries these methods have revealed. Further, this edition includes a
new chapter, “Disorders of the Developing Nervous System,” which highlights disorders such
as autism spectrum disorder and attention-deficit/hyperactivity disorder.

Key Features
• Chapter-opening vignettes describe the experience of people whose lives are impacted
by various disorders related to—among others—behavior, learning, sleep, and ingestion.
These vignettes set the tone and introduce students to a key concept discussed in the
chapter.
• Thought Questions at the end of each module provide students with real-world
behavioral neuroscience challenges. Students determine appropriate courses of action by
applying concepts learned, which develops their critical thinking abilities.
• End-of-chapter Review Questions help students reinforce connections across topics and

THIRTEENTH
EDITION
allow instructors to monitor class progress, and intervene when necessary.

Also available separately for purchase is Revel for Physiology of Behavior. Revel is an
interactive learning environment that deeply engages students and prepares them for class.
Media and assessment integrated directly within the authors’ narrative lets students read,
explore interactive content, and practice in one continuous learning path. Thanks to the
dynamic reading experience in Revel, students come to class prepared to discuss, apply, and

Physiology of Behavior
Carlson • Birkett
learn from instructors and from each other. Critical Concepts features have been added to
each chapter in this Revel, with goals of highlighting important topics in neuroscience and
providing opportunities to explore them in greater depth.

THIRTEENTH EDITION

Neil R. Carlson • Melissa A. Birkett

CVR_CARL0287_13_GE_CVR_Neografia.indd 1 18/11/21 5:05 PM

 Physiology
of Behavior
Thirteenth edition
Global edition

NEIL R. CARLSON
University of Massachusetts, Amherst

MELISSA A. BIRKETT
Southern Oregon University

Harlow, England • London • New York • Boston • San Francisco • Toronto • Sydney • Dubai • Singapore • Hong Kong
Tokyo • Seoul • Taipei • New Delhi • Cape Town • São Paulo • Mexico City • Madrid • Amsterdam • Munich • Paris • Milan

A01_CARL0287_13_GE_FM.indd 1 01/12/21 3:21 PM

Please contact https://support.pearson.com/getsupport/s/contactsupport with any queries on this content.

Acknowledgments of third-party content appear on the appropriate page within the text.

Pearson Education Limited

KAO Two
KAO Park
Hockham Way
Harlow
Essex
CM17 9SR
United Kingdom

and Associated Companies throughout the world

Visit us on the World Wide Web at: www.pearsonglobaleditions.com

© Pearson Education Limited 2022

The rights of Neil R. Carlson and Melissa A. Birkett to be identified as the authors of this work have been asserted
by them in accordance with the Copyright, Designs and Patents Act 1988.

Authorized adaptation from the United States edition, entitled Physiology of Behavior, 13th Edition, ISBN 978-0-
135-70983-2 by Neil R. Carlson and Melissa A. Birkett, published by Pearson Education © 2021.

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in
any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without either the prior
written permission of the publisher or a license permitting restricted copying in the United Kingdom issued by the
Copyright Licensing Agency Ltd, Saffron House, 6–10 Kirby Street, London EC1N 8TS. For information regarding
permissions, request forms, and the appropriate contacts within the Pearson Education Global Rights and
Permissions department, please visit www.pearsoned.com/permissions/.

All trademarks used herein are the property of their respective owners. The use of any trademark in this text does
not vest in the author or publisher any trademark ownership rights in such trademarks, nor does the use of such
trademarks imply any affiliation with or endorsement of this book by such owners.

This eBook is a standalone product and may or may not include all assets that were part of the print version. It also
does not provide access to other Pearson digital products like Revel. The publisher reserves the right to remove
any material in this eBook at any time.

British Library Cataloguing-in-Publication Data

A catalogue record for this book is available from the British Library

ISBN 10: 1-292-43028-1

ISBN 13: 978-1-292-43028-7
eBook ISBN 13: 978-1-292-430256

Typeset in by B2R Technologies Pvt. Ltd.

 Brief Contents
1 Introduction 15

2 Structure and Functions of Cells of the Nervous System 36

3 Structure of the Nervous System 68

4 Psychopharmacology and Neurotransmitters 91

5 Methods and Strategies of Research 119

6 Vision 153

7 Audition, the Body Senses, and the Chemical Senses 191

8 Control of Movement 235

9 Sleep and Biological Rhythms 266

10 Reproductive and Parental Behavior 301

11 Emotion 337

12 Ingestive Behavior 375

13 Learning and Memory 414

14 Human Communication 456

15 The Developing Nervous System 491

16 Neurological Disorders 512

17 Schizophrenia and the Affective Disorders 548

18 Stress and Anxiety Disorders 580

19 Substance Abuse 604

A01_CARL0287_13_GE_FM.indd 3 01/12/21 3:21 PM

 This page is intentionally left blank

A01_CARL0287_13_GE_FM.indd 4 01/12/21 3:21 PM

 Contents
Preface11 Activation of Receptors 59
Postsynaptic Potentials 60
Effects of Postsynaptic Potentials:
Neural Integration 61
Termination of Postsynaptic Potentials 62
Autoreceptors63
Other Types of Synapses 64
Nonsynaptic Chemical Communication 65

1 Introduction 15
Foundations of Behavioral Neuroscience 17
The Goals of Research 17
Roots of Behavioral Neuroscience 18
Natural Selection and Evolution 23
Functionalism and the Inheritance of Traits 23
Evolution of Human Brains 25
Ethical Issues in Research with Humans
and Other Animals 28 3 Structure of the Nervous System 68
Research with Animals 28
Basic Features of the Nervous System 69
Research with Humans 29
Anatomical Directions 70
The Future of Neuroscience: The Meninges and Ventricular System 72
Careers and Strategies for Learning 31
Structure and Function of the Central
Careers in Neuroscience 31
Nervous System (CNS) 74
Strategies for Learning 31
The Forebrain: Telencephalon 75
The Forebrain: Diencephalon 80
The Midbrain: Mesencephalon 82
The Hindbrain: Metencephalon and
Myelencephalon83
The Spinal Cord 83
Structure and Function of the Peripheral Nervous
System (PNS) 85
Cranial Nerves 85
2 Structure and Functions of Spinal Nerves 86
Cells of the Nervous System 36 The Autonomic Nervous System 87

Cells of the Nervous System 38

The Nervous System: An Overview 38
Neurons39
Supporting Cells 44
The Blood–Brain Barrier 46
Communication Within a Neuron 48
Neural Communication: An Overview 48
Electrical Potentials of Axons
The Membrane Potential
49
49
4 Psychopharmacology and
The Action Potential 51
Neurotransmitters 91
Conduction of the Action Potential 53 Principles of Psychopharmacology 93
Communication Between Neurons 56 An Overview of Psychopharmacology 93
Structure of Synapses 56 Pharmacokinetics94
Release of Neurotransmitters 58 Drug Effectiveness 96
5

A01_CARL0287_13_GE_FM.indd 5 01/12/21 3:22 PM

 6 Contents

Effects of Repeated Administration 97

Placebo Effects 98
Sites of Drug Action 99
Effects on Production of Neurotransmitters 100
Effects on Storage and Release of
Neurotransmitters101
Effects on Receptors 101
Effects on Reuptake and Deactivation
of Neurotransmitters 102
Neurotransmitters and Neuromodulators
Amino Acids
103
103
6 Vision153
Acetylcholine (ACh) 106 The Eye 155
The Monoamines 108 Introduction to Sensation and Perception 155
Peptides115 The Stimulus: Light 156
Lipids116 Anatomy of the Eye 156
Photoreceptors158
Transduction159
Central and Peripheral Vision 160
Overview of the Visual Pathway 161
Other Retinal Pathways 161
Brain Regions Involved in Visual Processing 163
Lateral Geniculate Nucleus 163
Striate Cortex 163
Extrastriate Cortex 166
Perceiving Color 168
5 Methods and Strategies of Research 119 Role of the Retinal Ganglion Cells 168
Role of the Retina 169
Experimental Ablation 121 Role of the Striate and Extrastriate Cortex 171
Evaluating the Behavioral Effects of Brain Perceiving Form 175
Damage121
Role of the Striate Cortex 175
Producing Brain Lesions 122
Role of the Extrastriate Cortex 176
Stereotaxic Surgery 123
Perceiving Spatial Location 182
Histological Methods 125
Role of the Retina 182
Tracing Neural Connections 127
Role of the Striate and Extrastriate Cortex 182
Studying the Structure of the Living
Human Brain 131 Perceiving Orientation and Movement 185
Recording and Stimulating Neural Activity 136 Role of the Striate Cortex 185
Recording Neural Activity 136 Role of the Extrastriate Cortex 185
Recording the Brain’s Metabolic and
Synaptic Activity 138
Stimulating Neural Activity 140
Neurochemical Methods 144
Finding Neurons That Produce
Particular Neurochemicals 144
Localizing Particular Receptors 145
Measuring Chemicals Secreted in
the Brain 146
Genetic Methods 148
7 Audition, the Body Senses,
Twin Studies 148
and the Chemical Senses 191
Adoption Studies 148
Genomic Studies 149 Audition193
Targeted Mutations 149 The Stimulus: Sound 193
Antisense Oligonucleotides 150 Anatomy of the Ear 194
CRISPR-Cas Methods 150 Auditory Hair Cells Transduce Auditory Information 196

A01_CARL0287_13_GE_FM.indd 6 01/12/21 3:22 PM

 Contents 7

The Auditory Pathway 197 Control of Reaching and Grasping:

Perceiving Pitch 200 Role of the Parietal Cortex 261
Perceiving Loudness 201 Deficits of Skilled Movements:
Perceiving Timbre 202 Apraxias and Dyspraxia 262
Perceiving Spatial Location 202 Limb Apraxia 262
Perceiving Complex Sounds 205 Constructional Apraxia 263
Perceiving Music 207 Dyspraxia263
Vestibular System 210
Anatomy of the Vestibular Apparatus 210
The Vestibular Pathway 212
Somatosenses214
The Stimuli 214
Anatomy of the Skin and Its Receptive Organs 214
Perceiving Cutaneous Stimulation 214
The Somatosensory Pathways 218
Perceiving Pain 220 9 Sleep and Biological Rhythms 266
Gustation226 What Is Sleep? 268
The Stimuli 226 Stages of Sleep 268
Anatomy of the Taste Buds and Gustatory Cells 227 Brain Activity During Sleep 271
Perceiving Gustatory Information 227 Why Do We Sleep? 273
The Gustatory Pathway 228 Functions of Sleep 273
Olfaction230 Functions of Slow-Wave Sleep 274
The Stimulus and Anatomy of the Olfactory Functions of REM Sleep 275
Apparatus230
Sleep and Learning 276
Transducing Olfactory Information 232
Physiological Mechanisms of Sleep and Waking 278
Perceiving Specific Odors 232
Neural Control of Sleep 278
Neural Control of Arousal 279
Neural Control of Sleep/Waking Transitions 283
Neural Control of Transition to REM 286
Disorders of Sleep 289
Insomnia289
Narcolepsy290
REM Sleep Behavior Disorder 291

8 Control of Movement 235

Problems Associated with Slow-Wave Sleep
Biological Clocks
292
293
Skeletal Muscle 237 Circadian Rhythms and Zeitgebers 293
Anatomy237 The Suprachiasmatic Nucleus 294
The Physical Basis of Muscular Contraction 237 Control of Seasonal Rhythms: The Pineal Gland
Sensory Feedback from Muscles 239 and Melatonin 298
Changes in Circadian Rhythms: Shift Work and Jet Lag 298
Control of Movement by the Spinal Cord 241
The Monosynaptic Stretch Reflex 241
The Gamma Motor System 243
Polysynaptic Reflexes 243
Control of Movement by the Brain 245
Cortical Structures 245
Planning and Initiating Movements:
Role of the Motor Association Cortex
Subcortical Structures
246
250
10 Reproductive and Parental Behavior 301
Cortical Control of Movement: Descending Pathways 255 Sexual Development 303
Complex Motor Behavior 258 Production of Gametes and Fertilization 303
Imitating and Comprehending Movements: Development of the Sex Organs 303
Role of the Mirror Neuron System 258 Sexual Maturation 307

A01_CARL0287_13_GE_FM.indd 7 01/12/21 3:23 PM

 8 Contents

Control of Sexual Behavior by Hormones and Pheromones 309

Hormonal Control of Female Reproductive Cycles 309
Hormonal Control of Sexual Behavior
of Laboratory Animals 310
Organizational Effects of Androgens on Behavior:
Masculinization and Defeminization 312
Human Sexual Behavior 312
Effects of Pheromones 315
Neural Control of Sexual Behavior 319
Male Sexual Behavior 319
Female Sexual Behavior 321
Formation of Pair Bonds 323
Sexual Orientation 324
Activational and Organizational Effects of Hormones 325
Role of Steroid Hormones 325 12 Ingestive Behavior 375
Sexual Orientation and the Brain 326 Drinking377
Role of Prenatal Environment in Physiological Regulatory Mechanisms 377
Sexual Orientation 328
Two Types of Thirst 378
Heredity and Sexual Orientation 328
Neural Mechanisms of Thirst 381
Parental Behavior 330
What Is Metabolism? 382
Maternal Behavior of Rodents 330
The Short-Term Reservoir 383
Hormonal Control of Maternal Behavior 331
The Long-Term Reservoir 384
Neural Control of Maternal Behavior 332
Fasting Phase 384
Neural Control of Paternal Behavior 334
Absorptive Phase 385
What Starts a Meal? 386
Environmental Factors 386
Gastric Factors 386
Metabolic Signals 387
What Stops a Meal? 389
Short-Term Satiety 390
11 Emotion337 Environmental Factors 391
Sensory Factors 391
Fear339
Gastric Factors 391
Components of Emotional Response 339
Intestinal Factors 391
Research with Laboratory Animals 339
Liver Factors 392
Research with Humans 343
Insulin393
Aggression345 Adipose Tissue Factors 393
Research with Laboratory Animals 345
Brain Mechanisms 395
Research with Humans 346
Brain Stem 395
Hormonal Control of Aggressive Behavior 347
Hypothalamus396
Impulse Control 353
Obesity402
Role of the vmPFC 353
Possible Causes 403
Brain Development and Impulse Control 355
Treatment405
Serotonin and Impulse Control 355
Eating Disorders 409
Moral Decision Making 356
Possible Causes 410
Communication of Emotions 358
Treatment412
Facial Expression of Emotions: Innate Responses 358
Neural Basis of the Communication
of Emotions: Recognition 359
Neural Basis of the Communication
of Emotions: Expression 365
Feeling Emotions 369
The James-Lange Theory 370
Feedback from Emotional Expressions 372

A01_CARL0287_13_GE_FM.indd 8 01/12/21 3:23 PM

 Contents 9

13 Learning and Memory 414 Disorders of Language Production: Broca’s Aphasia

Disorders of Language Comprehension:
465

Overview of Learning and Memory 417 Wernicke’s Aphasia 467

Types of Learning 417 Conduction Aphasia 473
Types of Memory 420 Aphasia in People Who Are Deaf 475
Stimulus-Response Learning 423 Stuttering476
Classical Conditioning 423 Disorders of Reading and Writing 479
Operant Conditioning 425 Pure Alexia 479
Motor Learning 430 Toward an Understanding of Reading 481
Role of the Cortex 430 Toward an Understanding of Writing 487
Role of the Basal Ganglia 430
Perceptual Learning 431
Role of the Cortex 431
Retaining Perceptual Information in
Short-Term Memory 432
Relational Learning 434
Role of the Hippocampus 434
Role of the Cortex 439
Amnesia439
Role of the Hippocampus 440
Stimulus-Response Learning 442
Motor Learning 442
Perceptual Learning 443
Relational Learning
Long-Term Potentiation
443
447
15 The Developing Nervous System 491
Induction of Long-Term Potentiation 447 Development of the Nervous System 492
Role of NMDA Receptors 448 An Overview of Brain Development 493
Role of AMPA Receptors 450 Prenatal Brain Development 493
Role of Synaptic Changes 451 Postnatal Brain Development 497
Disorders of Development 499
Toxic Chemicals 499
Inherited Metabolic Disorders 499
Down Syndrome 501
Autism Spectrum Disorder 503
Symptoms503
Genetic and Environmental Factors 504
Brain Changes 504
Attention-Deficit/Hyperactivity Disorder 508
Symptoms508
Genetic and Environmental Factors 509
Brain Changes 510

14 Human Communication456
Language Production and Comprehension:
Brain Mechanisms 458
Lateralization458
Language Production and Comprehension
in the Brain 460
16 Neurological Disorders 512
Bilingualism461 Tumors and Seizures 514
Prosody462 Tumors514
Voice Recognition 463 Seizures516
Disorders of Language Production Cerebrovascular Accidents 520
and Comprehension 465 Causes520

A01_CARL0287_13_GE_FM.indd 9 01/12/21 3:23 PM

 10 Contents

Treatments521
18 Stress and Anxiety Disorders 580
Traumatic Brain Injury 524
Causes525 Stress581
Treatments525 Physiology of the Stress Response 582
Health Effects of Long-Term Stress 583
Degenerative Disorders 527
Effects of Stress on the Brain 584
Transmissible Spongiform Encephalopathies 527
Psychoneuroimmunology587
Parkinson’s Disease 528
Posttraumatic Stress Disorder 591
Huntington’s Disease 533
Symptoms591
Amyotrophic Lateral Sclerosis 536
Genetic and Environmental Factors 591
Multiple Sclerosis 537
Brain Changes 592
Dementia538
Treatment593
Korsakoff’s Syndrome 542
Anxiety Disorders 595
Disorders Caused by Infectious Diseases 544
Symptoms595
Encephalitis545 Genetic and Environmental Factors 595
Meningitis546 Brain Changes 596
Treatment596
Obsessive-Compulsive Disorder 599
Symptoms599
Genetic and Environmental Factors 600
Brain Changes 600
Treatment601

17 Schizophrenia and the Affective

Disorders548
Schizophrenia550
19 Substance Abuse 604
Description550
Genetic Factors 551 Common Features of Substance Abuse 606
Environmental Factors 553 Positive Reinforcement 607
Anomalies in Schizophrenia 556 Negative Reinforcement 613
The Mesolimbic Dopamine Pathway: Genetic Factors 616
Positive Symptoms 559 Alcohol617
The Mesocortical Dopamine Pathway: Nicotine617
Negative and Cognitive Symptoms 561 Stimulants618
Affective Disorders 566 Brain Mechanisms Associated with
Description566 Commonly Abused Drugs 618
Genetic Factors 566 Opiates619
Biological Treatments 567 Stimulants620
Role of the Frontal Cortex 571 Nicotine621
The Monoamine Hypothesis 573 Alcohol625
Cannabis628
Role of the 5-HT Transporter 573
Role of Neurogenesis 574 Treatment for Substance Abuse 630
Opiates630
Role of Circadian Rhythms 575
Stimulants632
Nicotine632
Alcohol633
Brain Stimulation 634
Glossary636
References651
Name Index 704
Subject Index 722

A01_CARL0287_13_GE_FM.indd 10 01/12/21 3:23 PM

 Preface
I
wrote the first edition of Physiology of Behavior over
30 years ago. When I did so, I had no idea I would
someday be writing the thirteenth edition. I’m still hav-
ing fun, so I hope to do a few more. The interesting work
coming out of my colleagues’ laboratories—a result of their
creativity and hard work—has given me something new
to say with each edition. Because there was so much for
me to learn, I enjoyed revising this edition just as much as
writing the first one. That is what makes writing new edi-
tions interesting: learning something new and then trying
to find a way to convey the information to the reader.
In this edition, Melissa Birkett updated content to re-
flect new research developments, and formulated a separate
chapter on disorders of the developing nervous system. To-
gether, we drew upon our teaching and experience work-
ing with students to create a comprehensive and accessible
guide for students of behavioral neuroscience.
The first part of the book is concerned with founda-
tions of behavioral neuroscience: the history of the field, the
structure and functions of neurons, neuroanatomy, psycho-
pharmacology, and research methods. The second part is
concerned with inputs and outputs that guide behavior: the
sensory systems and the motor system. The third part deals
with classes of species-typical behavior: sleep, reproduction,
emotional behavior, and ingestion. The chapter on reproduc-
tive behavior includes parental behavior as well as courting
and mating. The chapter on emotion includes a discussion of
fear, anger and aggression, communication of emotions, and
feeling emotions. The chapter on ingestive behavior includes
the neural and metabolic bases of drinking and eating. The
fourth part of the book explores learning, including research
on synaptic plasticity, the neural mechanisms that are re-
sponsible for perceptual learning and stimulus-response
learning (including classical and operant conditioning), hu-
man amnesia, and the role of the hippocampal formation in
relational learning. The final part of the book examines the
neural basis of human communication as well as neurologi-
cal, mental, and behavioral disorders. Behavioral disorders
are addressed in four chapters; the first is a new chapter
combining information about development of the nervous
system with information about disorders of development,
autism spectrum disorders, and attention-deficit/hyper-
activity disorder; the second discusses schizophrenia and
the affective disorders; the third discusses stress and anxi-
ety; and the fourth discusses substance abuse. Each chapter
begins with a Case Study, which describes the experience of
people whose lives are impacted by an important issue in
neuroscience. Other case studies are included within the text
of the chapters. Learning Objectives to guide your reading are
A01_CARL0287_13_GE_FM.indd 11
found at the beginning of each major section of the text. The
learning objectives can help you identify and understand the
key points from each section and are also summarized at the
end of each module. Thought Questions are also located at
the end of each module and are designed to stimulate your
thinking about what you have learned. Chapter Review Ques-
tions conclude each chapter. They provide useful reviews of
each chapter and a more comprehensive opportunity to test
your understanding. In Revel, Critical Concepts features have
been added to each chapter, with goals of highlighting im-
portant topics in neuroscience and providing opportunities
to explore them in greater depth.
New to This Edition
The research reported in this edition reflects both the enor-
mous advances made in research methods and the dis-
coveries these methods have revealed. In neuroscience, as
soon as a new method is developed in one laboratory, it is
adopted by other laboratories and applied to a wide range
of problems. Researchers are combining techniques that
converge upon the solution to a problem and use many
methods, often in collaboration with other laboratories.
The art in this book continues to evolve. For this thir-
teenth edition, the art has been updated to improve acces-
sibility, as well as to keep up with the latest findings and
studies in the field. We have always striven to be as up to
date and as accurate as possible. We hope the new art in this
edition reflects that ongoing effort.
You’ll notice that many of the chapters contain new
headings and subheadings, as well as more concise learn-
ing objectives. We believe that this approach will help the
reader to more easily identify main themes and concepts.
The following list summarizes some of the updates
new to this edition.
Chapter 1: Introduction
• New research on adult neurogenesis has been added.
• Epigenetics is included as an important concept in
behavioral neuroscience.
• New media content has been incorporated into Revel.
Chapter 2: Structure and Functions of Cells of the
Nervous System
• New media content has been incorporated into Revel.
Chapter 3: Structure of the Nervous System
• Figures were revised.
• Information about development of the nervous
system was moved to Chapter 15.
  11
01/12/21 3:23 PM
 12 Preface
Chapter 4: Psychopharmacology and
Neurotransmitters
• A new case study has been added to the beginning
of the chapter.
Chapter 5: Methods and Strategies of Research
• A new section about CRISPR techniques has been
included.
Chapter 6: Vision
• An example of flat vision following damage to the
parieto-occipital cortex has been added.
Chapter 7: Audition, the Body Senses, and the
Chemical Senses
• An additional case study has been added to the
beginning of the chapter.
Chapter 8: Control of Movement
• A new section on dyspraxia has been added.
Chapter 9: Sleep and Biological Rhythms
• Research on lucid dreaming has been included.
Chapter 10: Reproductive and Parental Behavior
• Several new topics have been added: research on
5α-reductase deficiency, research on changes
across the menstrual cycle, information about
trace amine-associated receptors, and research on
paternal behavior.
Chapter 11: Emotion
• Research on heredity of aggression, and testosterone
and environment have been added.
Chapter 12: Ingestive Behavior
• Information about food deserts has been added.
• New research related to leptin and reinforcement
in weight loss has been added.
• New treatments for binge eating disorder have been
included.
Chapter 13: Learning and Memory
• Organization and descriptions have been updated
throughout the chapter.
Chapter 14: Human Communication
• New research on brain regions involved in multiple
languages, tip of the tongue phenomenon, and stut-
tering have been added.
NEW! Chapter 15: Disorders of the Developing
Nervous System
• Information about development of the nervous
system and disorders of development was moved
to this chapter.
• Information about autism spectrum disorder and
attention-deficit/hyperactivity disorder was
moved to this chapter.
A01_CARL0287_13_GE_FM.indd 12
Chapter 16: Neurological Disorders
• This chapter was renumbered.
• Information about disorders of development was
moved to Chapter 15.
• New surgical techniques have been added.
• Research on possible treatments for Huntington’s
disease has been added.
Chapter 17: Schizophrenia and the Affective Disorders
• This chapter was renumbered.
• Information about neurodevelopmental disorders
was moved to Chapter 15.
• Research on genetic factors involved in schizophrenia
has been added.
• A new section on marijuana and schizophrenia has
been added.
• Descriptions of new treatment options for postpar-
tum depression have been added.
Chapter 18: Stress and Anxiety Disorders
• This chapter was renumbered.
• New research on the role of the hippocampus in
chronic pain has been added.
Chapter 19: Substance Abuse
• This chapter was renumbered.
• Research on epigenetic factors related to cocaine
abuse has been added.
• Information about opiate abuse interventions has
been added.
• Research on e-cigarettes has been added.
Resources for Instructors
REVEL™ Revel is an interactive learning environment that
deeply engages students and prepares them for class. Me-
dia and assessment integrated directly within the authors’
narrative lets students read, explore interactive content,
and practice in one continuous learning path. Thanks to
the dynamic reading experience in Revel, students come to
class prepared to discuss, apply, and learn from instructors
and from each other.
Learn More About Revel
www.pearson.com/revel
Instructor’s Manual (ISBN 9781292430225) Each chapter
includes an Integrated Teaching Outline with teaching
objectives, lecture material, demonstrations and activities,
videos, and student handouts. The Instructor’s Manual is
avai­lable online at www.pearsonglobaleditions.com.
Test Bank (ISBN 9781292430232) Includes over 2,500
thoroughly reviewed multiple-choice, completion, short
answer, and essay questions, each with answer feedback,
correlations to both text learning objectives and APA
01/12/21 3:23 PM

learning objectives, difficulty rating, and skill type des-
ignation. The Test Bank is available online at www
.­pearsonglobaleditions.com.
Enhanced Lecture PowerPoint Slides with Embedded
Videos (ISBNs Lecture PPTs 9781292430201; Video PPTs
9781292430218) The enhanced lecture PowerPoints offer
detailed outlines of key points for each chapter supported
by selected visuals from the textbook, and include the
videos featured in Revel. Standard Lecture PowerPoints
without embedded videos are also available. Slides are
available for download from the Instructor’s Resource
Center at www.pearsonglobaleditions.com.
Acknowledgments
We would like to thank the many colleagues and review-
ers who have provided invaluable time, expertise, and
resources to making this the best book it can be.
John Agnew, Walden University
MaryBeth Ahlum, Nebraska Wesleyan University
Mark Basham, Regis University
Lora Becker, University of Evansville
Jessica Bodoh-Creed, California State University,
Los Angeles
Melissa Burns-Cusato, Centre College
Giuseppe Cataldo, Queens College–CUNY
Cynthia Cimino, University of South Florida
Ann Cohen, University of Pittsburgh
Deborah Conway, Community College of Allegheny
County
Patricia Costello, Walden University
Traci Craig, University of Idaho
Joseph DeBold, Tufts University
Darragh Devine, University of Florida
David Devonis, Graceland University
Jeannie DiClementi, Indiana University–Purdue
University Fort Wayne
Nick Dominello, Holy Family University
William Dragon, Cornell College
Michael Dudley, Southern Illinois University
Edwardsville
Jean Egan, Asnuntuck Community College
Marie-Joelle Estrada, University of Rochester
Claire Etaugh, Bradley University
Rebecca Foushee, Fontbonne University
Mary Fraser, Menlo College
Joseph Green, The Ohio State University, Lima
Aaron Godlaski, Centre College
John C. Hallock, Pima Community College
Julie Hanauer, Suffolk County Community College
A01_CARL0287_13_GE_FM.indd 13
Preface 13
Linda Lockwood, Metropolitan State University of
Denver
Euriel Merrick, South University
Heather Molenda-Figueira, University of Wisconsin,
Stevens Point
Brian Piper, Husson University School of Pharmacy
Trisha Prunty, Lindenwood University–Belleville
Christian Reich, Ramapo College of New Jersey
KatieAnn Skogsberg, Centre College
Suzanne Sollars, University of Nebraska at Omaha
Doug Wallace, Northern Illinois University
Matthew Will, University of Missouri, Columbia
Scott Wood, Azusa Pacific University
Several colleagues have reviewed previous editions
of this book and made excellent suggestions for improve-
ment. We thank:
Massimo Bardi, Marshall University
Kyle Baumbauer, Texas A&M University
Lora Becker, University of Evansville
Annie Cardell, Mountain State University
James Cherry, Boston University
Gary Dunbar, Central Michigan University
Walter Isaac, Georgia College & State University
Eric Jackson, University of New Mexico
Karen Jennings, Keene State College
Linda Lockwood, Metropolitan State College of Denver
Christopher May, Carroll University
Khaleel Razak, University of California, Riverside
Christian Reich, Ramapo College of New Jersey
Christopher Sletten, University of North Florida
Alicia Swan, Southern Illinois University
Lorey Takahashi, University of Hawaii
Sheralee Tershner, Western New England University
Charles Trimbach, Roger Williams University
Steve Weinert, Cuyamaca College
Erin Young, Texas A&M University
Finally, Neil thanks his wife, Mary, for her support, and
Melissa thanks her family and colleagues for their support.
Please write to tell us what you like and dislike about
the book at: nrc@psych.umass.edu.
Global Edition Acknowledgments
Pearson would like to thank Tatiana Novoselova, Middle-
sex University, for contributing to the Global Edition.
Global Edition Reviewers
Simon Evans, University of Surrey
Tom Lockheart, University of Portsmouth
01/12/21 3:23 PM
 14 Preface
About the Authors
Neil R. Carlson pursued his undergraduate studies at the
University of Illinois. He had planned to study nuclear
physics, but when he discovered in an introductory
psychology course that psychology was really a science, he
decided that was what he wanted to do. Before changing his
major, Carlson talked with several professors and visited their
laboratories, and when he saw what physiological psycholo-
gists do, he knew that he had found his niche. He stayed on
at Illinois and received his Ph.D. Then, after a two-year post-
doctoral fellowship at the University of Iowa, Carlson came to
the University of Massachusetts, where he taught throughout
his entire career. He retired from UMass in the fall of 2004 but
continues to keep up with developments in the field of behav-
ioral neuroscience and to revise this book.
As an undergraduate psychology major at Cornell Univer­
sity, Melissa A. Birkett discovered courses in biopsychology,
behavior, endocrinology, and evolutionary psychology.
There, she was introduced to interdisciplinary research
incorporating multiple perspectives in the challenging
task of understanding behavior. She became interested in
A01_CARL0287_13_GE_FM.indd 14
learning about behavior and its underlying mechanisms.
She worked as an undergraduate research a­ssistant in
several laboratories on projects ranging from insect behavior
to sleep in undergraduates. Those formative experiences
and interactions with several influential research mentors
convinced her to pursue a career in research.
Birkett completed her Ph.D. in the Neuroscience and
Behavior program at the University of Massachusetts
Amherst (where Neil Carlson was a faculty member at the
time). In 2007, she became a faculty member at Northern
Arizona University in the Department of Psychological
Sciences, and in 2018 joined the psychology department
at Southern Oregon University. Birkett currently conducts
research related to the stress response and teaches under-
graduate courses in psychology, research methods, statistics,
behavioral neuroscience, and psychopharmacology. Each
year, she supervises student researchers and seeks to provide
them with the kinds of opportunities she found valuable as
a student. Her work has been recognized with awards for
outstanding teaching and teaching innovation, and she
has contributed to several publications on best practices in
teaching neuroscience.
01/12/21 3:23 PM
 Chapter 1
Introduction

Vitstudio/Shutterstock
The human nervous system contains billions of neurons.

Chapter Outline
Foundations of Behavioral Neuroscience Ethical Issues in Research with Humans
The Goals of Research and Other Animals
Roots of Behavioral Neuroscience Research with Animals
Research with Humans
Natural Selection and Evolution
Functionalism and the Inheritance of Traits The Future of Neuroscience: Careers and
Strategies for Learning
Evolution of Human Brains
Careers in Neuroscience
Strategies for Learning

Learning Objectives
LO 1.1 Compare the roles of generalization and LO 1.2 Summarize historical and contemporary
reduction in behavioral neuroscience contributions to behavioral neuroscience
research. from various scientific disciplines.

15

M01_CARL0287_13_GE_C01.indd 15 25/11/21 8:05 AM

 16 Chapter 1
LO 1.3 Describe the role of natural selection in
the evolution of behavioral traits.
LO 1.4 Identify factors involved in the evolution
of human brains.
LO 1.5 Outline reasons for the use of animals in
behavioral neuroscience research.
LO 1.6 Identify mechanisms for oversight of
animal research.
One day, while playing with her father in the garden, five-year-old
Anika suddenly stopped as her right arm twitched a little. For the
next few days, Anika’s parents observed her closely and noticed
the occurrence of similar episodes, which she had no memory
of. An examination by a doctor revealed Anika had s­ eizures, a
phenomenon when electrical energy which neurons use to com-
municate, gets out of control (you will read more about seizures
in Chapter 16). When this happens, people are briefly unable to
control their behavior and often develop involuntary movements.
Sometimes, if neurons that make up the motor system are in-
volved, a seizure can cause a convulsion, which is uncontrollable
activity of the muscles. When a person experiences more than
one seizure, the condition is termed as epilepsy.
In Anika’s case, the doctor concluded, after examining
her brain’s electrical activity and the brain images, that she had
­Rasmussen encephalitis. This is a rare inflammatory condition that
typically impairs only one of the brain hemispheres, and it usually
affects young children. Rasmussen encephalitis responds poorly to
medications and the most effective strategy is to surgically remove
the affected hemisphere to prevent catastrophic effects of the
At the end of the twentieth century, many researchers be-
lieved that the brain was incapable of change in adulthood.
However, some neuroscientists suggested the cells and con-
nections of the adult brain were flexible, or plastic, and at-
tempted to change the view of the brain that had been held
for more than a century. Changing this widely held view
was not an easy process. Though they were equipped with
revolutionary new data, the researchers were criticized
for years, and their data and methods were questioned.
Eventually, the data accumulated, and the scientific consen-
sus became that the adult brain continues to experience neu-
ral changes. This change in understanding about the brain
has been met with optimism and excitement. Therapies for
brain injury and mental illness have been developed based
on this new understanding of brain changes.
The adult brain modifies connections between the cells
in the brain, called neurons, throughout a lifetime. Dozens
of researchers are making new discoveries every year about
neurogenesis, the generation of new neurons particularly
in specific regions of the adult brain. New debates about
M01_CARL0287_13_GE_C01.indd 16
LO 1.7 Discuss ethical considerations in research
with human participants.
LO 1.8 Identify mechanisms for oversight of
human research.
LO 1.9 Identify careers in behavioral
neuroscience.
LO 1.10 Describe effective learning strategies for
studying behavioral neuroscience.
epileptic activity on the rest of the brain. Anika’s seizure activity was
concentrated in the left hemisphere of her brain and the treatment
of choice for her would be to remove the left hemisphere. Anika’s
parents were anxious about the consequences of such a drastic
treatment; however, the doctor assured them that this strategy
had proven to be highly beneficial in the long run. Initially, the sur-
gery leads to several motor and cognitive deficits but with intense
­rehabilitation, such patients can restore the lost functions and avoid
gross cognitive abnormalities. Such remarkable recovery is possible
due to one of the key features of the brain’s p ­ hysiology—­plasticity,
when networks of brain cells change through growth and
­reorganization forming new connections and circuits. ­Incremental
training of the affected body part helps the brain learn the lost
­motor functions. Plasticity occurs throughout life as a r­esult of
learning, experiences, and memory formation. However, it is most
effective in the early years of one’s life, making surgical interven-
tions for conditions such as Rasmussen encephalitis successful
and rendering patients both seizure free and adapted to life.
Source: Adam L. Hartman and J. Helen Cross, Timing of Surgery in ­Rasmussen
Syndrome: Is Patience a Virtue?, Epilepsy Currents, 2014.
neurogenesis have also arisen and understanding of this
phenomenon remains incomplete (Kempermann et al.,
2018). Some researchers have reported little to no evidence
of neurogenesis in the dentate gyrus of the hippocampus
(an area of the brain typically associated with neurogenesis)
of adult humans, and suggested that neurogenesis in this re-
gion declines throughout childhood in our species (Sorrells
et al., 2018). In contrast, other researchers report evidence
that neurogenesis in this region continues throughout the
human lifespan, well into older adulthood in healthy indi-
viduals (Moreno-Jiménez et al., 2019; Boldrini et al., 2018).
Researchers previously believed that the adult brain
also lost a large number of neurons during aging. These
losses were thought to underlie the inevitable cognitive
decline of older adulthood. Re-examination of this idea,
along with additional data and development of new
methods and technology, has produced a new consensus
for a relatively modest (2-4 percent) decline in neurons in
typical aging. Researchers are also currently r­ e-evaluating
long-held ideas about differences in the number of neurons
25/11/21 8:05 AM

in parts of the brain between men and women, the effects
of chronic alcohol use on damage to neurons, and changes
in neurons associated with mental illness and neurologi-
cal disorders. The neuroscience community is actively
engaged in understanding these apparently conflicting
results, and as one neuroscientist has summarized the sit-
uation, “It is important to keep an open mind and to be
inquisitive and creative, in order to separate truths from
myths” (von Bartheld, 2018, p. 12).
Behavioral neuroscience is a dynamic and ever-
changing field. As you read, consider not only the facts,
but also the research process used to obtain those facts,
and the exciting possibility that there is still much to learn
about the brain and the nervous system. The last frontier in
this world—and perhaps the greatest one—lies within us.
The human nervous system makes possible all that we can
do, all that we can know, and all that we can experience.
Its complexity is immense, and the task of studying it and
understanding it dwarfs all previous explorations our spe-
cies has undertaken.
Foundations of Behavioral
Neuroscience
Behavioral neuroscience was formerly known as physiologi-
cal psychology, and it is still sometimes referred to by that
name. In fact, the first psychology textbook, written by
Wilhelm Wundt in the late nineteenth century, was titled
Principles of Physiological Psychology. In recent years, the ex-
plosion of information from experimental biology, chem-
istry, animal behavior, psychology, computer science, and
other fields has contributed to creating the diverse inter-
disciplinary field of behavioral neuroscience. This united
effort is due to the realization that the ultimate function of
the nervous system is behavior.
When we ask our students what they think the ulti-
mate function of the brain is, they often say “thinking,”
or “logical reasoning,” or “perceiving,” or “remembering
things.” The nervous system does perform these func-
tions, but they all support a single primary function: con-
trol of movement. (Note that movement includes speech
and other forms of communication, an important category
of human behavior.) The basic function of perception is
to inform us of what is happening in our environment so
that our behaviors will be adaptive and useful: Perception
without the ability to act would be useless. Once percep-
tual abilities evolved, they could be used for purposes
other than guiding behavior. For example, we can enjoy a
beautiful sunset or a great work of art without our percep-
tion causing us to do anything in particular. And thinking
can often take place without causing any overt behavior.
However, the ability to think evolved because it permits us
M01_CARL0287_13_GE_C01.indd 17
Introduction 17
to perform complex behaviors that accomplish useful self-
preserving goals. And whereas reminiscing about things
that happened in our past can be an enjoyable pastime, the
ability to learn and remember evolved—again—because it
permitted our ancestors to profit from experience and per-
form behaviors that were useful to them.
The growing field of behavioral neuroscience has
been formed by scientists who have combined the
experimental methods of psychology with those of phys-
iology and have applied them to the issues that concern
researchers in many different fields. Research in neuro-
science includes topics in perceptual processes, control
of movement, sleep and waking, reproductive behav-
iors, ingestive behaviors, emotional behaviors, learning,
and language. In recent years we have begun to study
the neuroscience underlying significant human health
concerns, such as substance abuse and neurological and
mental disorders. These topics are discussed in subse-
quent chapters.
The Goals of Research
LO 1.1 Compare the roles of generalization and
reduction in behavioral neuroscience research.
The goal of all scientists is to explain the phenomena
they study. But what do we mean by explain? Scientific
explanation takes two forms: generalization and reduc-
tion. Generalization refers to explanations as examples
of general laws, which are revealed through experiments.
Reduction refers to explanations of complex phenomena
in terms of simpler ones.
Behavioral neuroscientists seek to explain behavior
by studying the physiological processes that control it.
But behavioral neuroscientists cannot just be reduction-
ists. It is not enough to observe behaviors and correlate
them with physiological events that occur at the same
time. We also need to understand the function of a given
behavior. For example, mice, like many other mammals,
often build nests. Behavioral observations show that mice
will build nests under two conditions: when the air tem-
perature is low and when the animal is pregnant. A non-
pregnant mouse will build a nest only if the temperature is
cool, whereas a pregnant mouse will build one regardless
of the temperature. The same behavior occurs for differ-
ent reasons. Nest-building behavior is controlled by two
different physiological mechanisms. Nest building can be
studied as a behavior related to the process of temperature
regulation, or it can be studied in the context of parental
behavior. Although the same set of brain mechanisms will
control the movements that a mouse makes in building a
nest in both cases, these mechanisms will be activated by
different parts of the brain. One part receives information
from the body’s temperature detectors, and the other part
is influenced by hormones that are present in the body
25/11/21 8:05 AM
 18 Chapter 1

during pregnancy. It is not enough to observe behaviors

Figure 1.1 Galen (130–200 c.e.)
and correlate them with physiological events that occur at
the same time. We must understand the overall function of
a given behavior.
Sometimes, physiological mechanisms can tell us
something about psychological processes such as lan-

Photo Researchers/Science History Images/Alamy Stock Photo

guage, memory, or mood. For example, damage to a par-
ticular part of the brain can cause very specific impair-
ments in a person’s language abilities. The nature of these
impairments suggests how these abilities are organized in
the brain. When the damage involves a brain region that
is important in analyzing speech sounds, it also produces
deficits in spelling. This finding suggests that the ability to
recognize a spoken word and the ability to spell it rely on
related brain mechanisms. Damage to another region of the
brain can produce extreme difficulty in reading unfamiliar
words by sounding them out, but it does not impair the
person’s ability to read words with which they are already
familiar. This finding suggests that reading comprehension
can take two routes: one that is related to speech sounds
Not all ancient Greek scholars agreed with Hippocrates.
and another that is primarily a matter of visual recognition
Aristotle (384–322 b.c.e.), for example, thought the brain
of whole words.
served to cool the passions of the heart. Galen (130–200 c.e.)
In practice, the research efforts of behavioral neuro-
dissected and studied the brains of cattle, sheep, pigs, cats,
scientists involve both forms of explanation: generaliza-
dogs, weasels, monkeys, and apes (Finger, 1994), and con-
tion and reduction. Ideas for experiments are stimulated
cluded that Aristotle’s theory about the brain’s role was “ut-
by the investigator’s knowledge both of psychological
terly absurd, since in that case Nature would not have placed
generalizations about behavior and of physiological
the encephalon [brain] so far from the heart, . . . and she would
mechanisms. A good behavioral neuroscientist must
not have attached the sources of all the senses [the sensory
therefore be an expert in the study of behavior and the
nerves] to it” (Galen, 1968 translation, p. 387). (See Figure 1.1.)
study of physiology.
CONTRIBUTIONS FROM PHILOSOPHY: THE MIND–
BODY QUESTION From the earliest historical times,
Roots of Behavioral Neuroscience human beings have believed that they possess something
LO 1.2 Summarize historical and contemporary intangible that animates them: a mind, or a soul, or a spirit.
contributions to behavioral neuroscience from We each also have a physical body, with muscles that
various scientific disciplines. move it and sensory organs such as eyes and ears that per-
ceive information about the world around us. Within our
This section traces some of the discoveries of the past that
bodies the nervous system plays a central role, receiving
have contributed to the field of behavioral neuroscience
information from the sensory organs and controlling the
today and have helped advance our understanding of
movements of the muscles. But what role does the mind
mind, brain, and behavior.
play? Does it control the nervous system? Is it a part of the
CONTRIBUTIONS FROM THE ANCIENT WORLD: ROLE nervous system? Is it physical and tangible, like the rest of
OF THE HEART AND BRAIN Study of (or speculation the body, or is it a spirit that will always remain hidden?
about) the physiology of behavior has its roots in antiquity. This puzzle has historically been called the mind–body
A papyrus scroll from around 1700 b.c.e. contains surgical question. Philosophers have been trying to answer it for
records of head injuries and the oldest surviving descrip- many centuries, and more recently scientists have taken up
tions of the brain, cerebrospinal fluid, meninges, and skull the task. In general, people have followed two different ap-
(Feldman and Goodrich, 1999). proaches: dualism and monism. Dualism is a belief in the
Because its movement was necessary for life and be- dual nature of reality, which means that mind and body
cause emotions caused it to beat more strongly, ancient are separate. From a dualist perspective the body is made
Egyptian, Indian, and Chinese cultures considered the of ordinary matter, but the mind is not. Monism is a belief
heart to be the seat of thought and emotions. The ancient that everything in the universe consists of matter and en-
Greeks did too, but Hippocrates (460–370 b.c.e.) concluded ergy and that the mind is a phenomenon produced by the
that this role should be assigned to the brain. workings of the nervous system. (See Figure 1.2.)

M01_CARL0287_13_GE_C01.indd 18 25/11/21 8:05 AM


Figure 1.2 The Mind–Body Question
Monism and dualism pose two possible answers to the mind–body question.
Monism
Mind is
produced by the
brain
The French philosopher René Descartes’s (1596–1650)
speculations concerning the roles of the mind and brain in
the control of behavior provide a good starting point in the
modern history of behavioral neuroscience. To Descartes,
animals were mechanical devices. He believed their behav-
ior was controlled by environmental stimuli. His view of
the human body was much the same: It was a machine. As
Descartes observed, some movements of the human body
were automatic and involuntary. For example, if a person’s
finger touched a hot object, the arm would immediately
withdraw from the source of stimulation. Reactions like
this did not require participation of the mind; they occurred
automatically. Descartes called these actions ­reflexes. (See
Figure 1.3.)
Like most philosophers of his time, Descartes was a
dualist and believed that each person possessed a mind—
a uniquely human attribute that was not subject to the
laws of the universe. But his thinking differed from that
of his predecessors in one important way: He was the first
to suggest that a link exists between the human mind and
its purely physical housing, the brain. He believed that
the mind controlled the movements of the body, while the
body, through its sense organs, supplied the mind with in-
formation about what was happening in the environment.
In particular, he hypothesized that this interaction took
place in the pineal body, a small organ situated on top of
the brain stem, buried beneath the cerebral hemispheres.
He noted that the brain contained hollow chambers (the
ventricles) that were filled with fluid, and he hypothesized
that this fluid was under pressure. When the mind decided
to perform an action, it tilted the pineal body in a particu-
lar direction like a little joystick, causing fluid to flow from
M01_CARL0287_13_GE_C01.indd 19
Introduction 19
Dualism
Mind is
separate from the
brain
the brain into the appropriate set of nerves. This flow of
fluid caused muscles to inflate and move. As you’ll learn in
the rest of this section, it did not take long for biologists to
disprove Descartes’s belief about the brain using pressur-
ized fluid to control behavior.
Speculating about the nature of the mind can get us
only so far. If we could answer the mind–body question
simply by thinking about it, philosophers would have
done so long ago. Behavioral neuroscientists take an em-
pirical, monistic approach to the study of human nature.
Most neuroscientists believe that once we understand
the workings of the human body—and, in particular, the
Figure 1.3 Descartes’s Model
Descartes believed that the “soul” (what we now call the mind)
controls the movements of the muscles through its influence on the
pineal body. According to his theory, the eyes sent visual information
to the brain, where it could be examined by the soul. When the
soul decided to act, it would tilt the pineal body (labeled H in the
diagram), which would divert pressurized fluid through nerves to the
appropriate muscles.
Oxford Science Archive/Heritage Images/The Print
Collector/Alamy Stock Photo
25/11/21 8:05 AM
 20 Chapter 1
workings of the nervous system—the mind–body ques-
tion will be resolved. We will be able to explain how we
perceive, how we think, how we remember, and how we
behave. We will even be able to explain the nature of our
own self-awareness.
CONTRIBUTIONS FROM PHYSIOLOGY: ELECTRICAL
COMMUNICATION IN THE NERVOUS SYSTEM Luigi
Galvani (1737–1798), an Italian physiologist, found that
electrically stimulating a frog’s nerve contracted the mus-
cle to which it was attached. Contraction occurred even
when the nerve and muscle were detached from the rest
of the body, so the ability of the muscle to contract and
the ability of the nerve to send a message to the muscle
were characteristics of these tissues themselves. Contrary
to Descartes’s description, the brain did not inflate muscles
by directing pressurized fluid through the nerve. Galvani’s
experiment prompted others to study the nature of the
message transmitted by the nerve and the means by which
muscles contracted. One of the most important figures
in the development of experimental physiology was Jo-
hannes Müller (1801–1858), a German physiologist. Müller
applied experimental techniques to physiology. Previously,
most natural scientists had been limited to observation
and classification. Although these activities are essential,
Müller insisted that major advances in our understand-
ing of the workings of the body would be achieved only
by experimentally removing or isolating animals’ organs,
testing their responses to various chemicals, and other-
wise altering the environment to see how the organs re-
sponded. His most important contribution to the study of
the physiology of behavior was his doctrine of specific
nerve ­energies. Müller observed that although all nerves
carry the same basic message—an electrical impulse—we
perceive the messages of different nerves in different ways.
For example, messages carried by the optic nerves produce
sensations of visual images, and those carried by the audi-
tory nerves produce sensations of sounds. How can differ-
ent sensations arise from the same basic message?
Müller’s answer was that the messages occur in dif-
ferent channels. The portion of the brain that receives
messages from the optic nerves interprets the activity as
visual stimulation, even if the nerves are actually stimu-
lated mechanically. (For example, when we rub our eyes,
we see flashes of light.) Because different parts of the brain
receive messages from different nerves, the brain must be
functionally divided: Some parts perform some functions,
while other parts perform others.
In 1870, German physiologists Gustav Fritsch (1838–
1927) and Eduard Hitzig (1838–1907) used electrical stim-
ulation as a tool for understanding the physiology of the
brain. They applied weak electrical current to the exposed
surface of a dog’s brain and observed the effects of the
stimulation. They found that stimulating different portions
M01_CARL0287_13_GE_C01.indd 20
of a specific region of the brain caused specific muscles to
contract on the opposite side of the body. We now refer to
this region as the primary motor cortex, and we know that
nerve cells there communicate directly with those that
cause muscular contractions. We also know that other re-
gions of the brain communicate with the primary motor
cortex to control behaviors. For example, regions of the
brain involved in speech communicate with the portion of
the primary motor cortex that controls the muscles of the
lips, tongue, and throat, which we use to speak.
German physicist and physiologist Hermann von
Helmholtz (1821–1894) studied many physiological pro-
cesses and was the first scientist to attempt to measure the
speed of conduction through nerves. Scientists had previ-
ously believed that such conduction was identical to the
conduction that occurs in wires, traveling at approximately
the speed of light. But Helmholtz found that neural con-
duction was much slower—only about 27.4 meters per
second. This measurement proved that neural conduction
was more than a simple electrical message, as we will see
in Chapter 2.
CONTRIBUTIONS FROM ANATOMY: STRUCTURE OF
THE NERVOUS SYSTEM Müller’s advocacy of experi-
mentation and the logical deductions from his doctrine of
specific nerve energies set the stage for other scientists to
perform experiments directly on the brain. Pierre Flourens
(1794–1867), a French researcher, did just that. Flourens re-
moved various parts of animals’ brains and observed their
behavior. By seeing what the animal could no longer do,
he could infer the function of the missing portion of the
brain. This method is called experimental ablation. Flou-
rens claimed to have discovered the regions of the brain
that control heart rate and breathing, purposeful move-
ments, and visual and auditory reflexes.
Soon after Flourens performed his experiments, Paul
Broca (1824–1880), a French surgeon, applied the principle
of experimental ablation to the human brain. He did not
intentionally remove parts of human brains to see how
they worked but observed the behavior of people whose
brains had been damaged by strokes. In 1861 he performed
an autopsy on the brain of a man who had had a stroke that
resulted in the loss of the ability to speak. Broca’s obser-
vations led him to conclude that a portion of the cerebral
cortex on the front part of the left side of the brain per-
forms functions that are necessary for speech. This came
to be known as Broca’s area. (See Figure 1.4.) Other physi-
cians soon obtained evidence supporting his conclusions.
As you will learn in Chapter 14, the control of speech is not
localized to only one particular region of the brain. Speech
requires many different functions, which are organized
throughout the brain. Nonetheless, the method of experi-
mental ablation remains important to our understanding
of the brains of both humans and laboratory animals.
25/11/21 8:05 AM
 Introduction 21

Figure 1.4 Broca’s Area Figure 1.5 Golgi-Stained Neurons

This region of the brain is named for French surgeon Paul Broca, Drawing of Neurons by Santiago Ramón y Cajal
who discovered that damage to a part of the left side of the brain
disrupted a person’s ability to speak.

Top
Broca’s
area

Front

Jan Purkinje (1787–1869), a Czech physiologist, stud-

ied both the central and peripheral nervous systems in the
middle of the nineteenth century. He discovered Purkinje
fibers—neurons terminating on cardiac cells responsible
for controlling contractions of the heart. He also used the
microscope to investigate the structure of neurons in many
regions of the brain, which included discovering Purkinje
cells in the cerebellum (Chvátal, 2017).
Late in the nineteenth century, Spanish anatomist
Santiago Ramón y Cajal (1852–1934) used the Golgi stain-
ing technique (described in Chapter 5) to examine individ-
ual neurons of the brain. His drawings of neurons (made Science Source
under magnification from a microscope) from the brain,
spinal cord, and retina depicted the detailed structures of
of behaviors, such as the mirror neuron system for coor-
these cells for the first time. Cajal proposed that the ner-
dinating social behavior (described in Chapter 8). Other
vous system consisted of billions of discrete, individual
developments contributed to new brain-based treatments
neurons, in opposition to the predominant idea of the time
for disorders such as severe depression and Parkinson’s
that the nervous system was a continuous network. In
disease (Chapters 16 and 17).
1906, he was awarded the Nobel Prize for his work describ-
The twenty-first century has already witnessed
ing the structure of the nervous system. Figure 1.5 shows
several important advances and discoveries. As re-
one of his drawings.
searchers continue to refine their understanding of the
CONTEMPORARY RESEARCH CONTRIBUTIONS structures and functions of the brain, new discoveries
Twentieth-century developments in experimental physiol- about pathways and circuits abound. For example, the
ogy included many important inventions, such as sensitive 2014 Nobel Prize was awarded to John O’Keefe (1939–),
amplifiers to detect weak electrical signals, neurochemi- May-Britt Moser (1963–), and Edvard Moser (1962) for
cal techniques to analyze chemical changes within and work on spatial positioning systems in the brain (also
between cells, and histological techniques to visualize called the brain’s global positioning system, or GPS). In
cells and their constituents. These and many other impor- 2017, Jeffrey Hall (1945–), Michael Rosbash (1944–), and
tant developments are discussed in detail in subsequent Michael Young (1949–) received the Nobel Prize for their
chapters. work describing the molecular mechanisms controlling
Briefly, highlights in contributions to neuroscience dur- circadian rhythms.
ing the twentieth century include discoveries ranging from New genetic techniques have spurred many excit-
the electrical and chemical messages used by neurons, to ing discoveries in neuroscience as well. The development
the circuits and brain structures involved in a wide variety of optogenetics provides researchers with the ability to

M01_CARL0287_13_GE_C01.indd 21 25/11/21 8:06 AM

 22 Chapter 1

selectively activate single neurons and observe changes
in behavior—using light! (See Chapter 5.) The develop-
ment of CRISPR-Cas9 techniques have enabled precise
editing of genetic material (DNA). This technique uses an
enzyme (Cas9) to cut out pieces of DNA paired with a set
of replacement directions (guide RNA) to create modified
genes. The impact of this technique on behavioral neu-
roscience is just beginning to be understood. The field of
epigenetics focuses on the role of the environment in the
expression of genes. Researchers continue to learn more
about how environmental-dependent gene expression can
have a profound impact on an individual’s behavior.
As behavioral neuroscience continues to progress as an
interdisciplinary field, efforts such as the European Human
Brain Project, which is working to develop a computer
simulation of the brain, and the Brain Research through
Advancing Innovative Neurotechnologies (BRAIN) ini-
tiative in the United States will continue to bring together
groups of researchers from biology, chemistry, engineering,
psychology, physiology, and other fields. Behavioral neu-
roscience, after all, has its roots—and its future—in inter-
disciplinary research.
DIVERSITY IN NEUROSCIENCE Neuroscience is a
diverse interdisciplinary field whose researchers work
around the globe. The Society for Neuroscience was
founded in 1969, with 500 members committed to devel-
oping a professional organization for scientists and phy-
sicians devoted to understanding the brain and nervous
system. This international organization now has approxi-
mately 37,000 members from over 90 different countries.
Reviewing the list of Nobel Prizes related to neuroscience
research in Table 1.1, you’ll notice the names of men and
women from several different countries. The field is cur-
rently striving to increase diversity through inclusivity of
women and underrepresented groups in the sciences.

Table 1.1 Selected Nobel Prizes for Research Related to Neuroscience

Year Recipients (country) Field of Study
1906 Camillo Golgi (Italy) and Santiago Ramón y Cajal (Spain) Structure of the nervous system
1936 Sir Henry Hallett Dale (U.K.) and Otto Loewi (Austria) Chemical transmission of nerve impulses
Sir John Carew Eccles (Australia), Sir Alan Lloyd Hodgkin (U.K.), and Sir Andrew
1963 Ionic mechanisms of nerve cell membrane
Fielding Huxley (U.K.)
Julius Axelrod (U.S.), Sir Bernard Katz (Germany, U.S.), and Ulf Svante von Euler
1970 Neurotransmitters
(Sweden)
1981 David Hubel (Canada, U.S.), Torsten Wiesel (Sweden, U.S.), and Roger Sperry (U.S.) Functions of the nervous system
2000 Arvid Carlsson (Sweden), Paul Greengard (U.S.), and Eric Kandel (U.S.) Neural communication
2014 John O’Keefe (U.S., U.K.), Edvard Moser (Norway), and May-Britt Moser (Norway) Spatial positioning system in the brain
2017 Jeffrey Hall (U.S.), Michael Rosbash (U.S.), and Michael Young (U.S.) Molecular mechanisms controlling circadian rhythms

Module Review: Foundations of Behavioral Neuroscience

The Goals of Research
LO 1.1 Compare the roles of generalization and
reduction in behavioral neuroscience research.
To explain the results of behavioral neuroscience re-
search, generalization can be used to reveal general laws
of behavior. Reduction can be used to explain complex
phenomena in terms of smaller, discrete phenomena.
Both are critical to understanding human behavior.
Roots of Behavioral Neuroscience
LO 1.2 Summarize historical and contemporary
contributions to behavioral neuroscience from
various scientific disciplines.
Ancient scholars disagreed on the importance of
the brain in behavior, and some attributed thought
and emotion to the heart. The mind–body question
influenced both historical (dualist) and contemporary
(monist) views of the brain. French philosopher Des-
cartes described reflexes but believed that other behav-
ior was the product of pressurized fluid causing muscles
to contract. Early physiologists influenced the study of
electrical components of neural communication. Müller
proposed the doctrine of specific nerve energies while
Fritsch and Hitzig studied the effects of electrical stim-
ulation of different brain regions. Galvani discovered
that nerves convey electrical messages, and von Helm-
holtz refined that understanding to begin to account
for chemical communication between cells. Advances
in research revealed the structure of the brain and the
cells of the nervous system. Flourens and Broca studied
the functions of brain regions using ablation. Purkinje
and Cajal studied the structures and functions of spe-
cific sets of neurons. Contemporary research continues

M01_CARL0287_13_GE_C01.indd 22 25/11/21 8:06 AM


in these areas and others, particularly with regard to the
genetic and molecular bases of behavior. Today, behav-
ioral neuroscience is a diverse, international, and inter-
disciplinary field.
Thought Question
Today, behavioral neuroscience actively applies a
computational approach to complex topics in brain
Natural Selection and
Evolution
Following the tradition of Müller and von Helmholtz,
other biologists continued to observe, classify, and think
about what they saw, to arrive at new conclusions. One of
the most important and influential of these scientists was
Charles Darwin (1809–1882). (See Figure 1.6.) Darwin for-
mulated the principles of natural selection and the theory
of evolution, which revolutionized biology at the time and
continues to shape the field of behavioral neuroscience
today. In science, a theory is an explanation that is strongly
supported by multiple lines of research with many con-
verging results.
Functionalism and the Inheritance
of Traits
LO 1.3 Describe the role of natural selection in the
evolution of behavioral traits.
Darwin’s theory emphasizes that all of an organism’s
­characteristics—its structure, its coloration, its behavior—
have functional significance. For example, the strong tal-
ons and sharp beaks that eagles possess permit the birds
to catch and eat prey. Caterpillars that eat green leaves
are themselves green, and their color makes it difficult for
birds to see them against their usual background. Mother
mice construct nests, which keep their offspring warm
and out of harm’s way. The behavior itself is not inherited.
What is inherited is a structure—the brain—that causes the
behavior to occur. Darwin’s theory gave rise to functional-
ism, the principle that characteristics of living organisms
perform useful functions. So, to understand the physi-
ological basis of various behaviors, we must first under-
stand what these behaviors accomplish (their function).
This means that we must understand something about the
natural history of the species being studied so that the be-
haviors can be seen in context. To understand the workings
of something as complex as a nervous system, we should
know what its functions are. Organisms of today are the
result of a long series of changes due to genetic variabil-
ity. Strictly speaking, we cannot say that any physiological
M01_CARL0287_13_GE_C01.indd 23
Introduction 23
physiology and in regulation of behavior. The interface
of computational technologies with the human brain
may potentially expand human capabilities beyond bio-
logical norms as well as aid patients with devastating
brain disorders. Which behavioral “superpower” would
you want to have and where would you start if you were
to develop such a project?
Figure 1.6 Charles Darwin (1809–1882)
Darwin’s theory of evolution revolutionized biology and strongly
influenced early psychologists.
North Wind Picture Archives
mechanisms of living organisms have a purpose. But they
do have functions, and these we can try to determine. For
example, the forelimb structures shown in Figure 1.7 are
adapted for different functions in different species of mam-
mals. Adaptations also occur in brain structures. For ex-
ample, male songbirds such as the white-crowned sparrow
possess highly developed brain structures (such as the ro-
bust nucleus of the archistriatum, high vocal center, and Area X)
that differ from some of their close, non-songbird relatives.
(See Figure 1.7.) The songbirds’ unique structures allow
them to learn and produce songs in response to complex
social and environmental stimuli. The function of male
song behavior in these species is to attract a mate and deter
rivals. The non-songbirds lack these brain structures and
their associated functions (Beecher and Brenowitz, 2005).
Among the various songbirds, in species in which only the
males sing, males have larger song brain structures com-
pared to females. In species in which both sexes sing duets,
there is no difference between the size of the structures in
males and females (Brenowitz, 1997).
25/11/21 8:06 AM
 24 Chapter 1

Figure 1.7 Adaption of Structures for Different Functions

The figure shows the forelimb bones of (a) human, (b) bat, (c) whale, and (d) dog. Through the process of natural selection, these bones have
been adapted to suit many different functions. Songbird (e) and non-songbird (f) brain structures also differ, corresponding with the different
functions of song in these species. Various song-related brain regions and their output to the structures used to produce song (the syrinx and
respiratory system) are labeled.

(a) (b) (c) (d)

HVC
mMAN
F

IMAN
L
NI

L
Field

Field
RA X

DLM
DM
llts DM
nX llts
m nX m
RA
RAr
VR
rV
RG

Syrinx Respiratory
Syrinx Respiratory
system
system
(e) (f)

Darwin formulated his theory of evolution to explain
the means by which species acquired their adaptive char-
acteristics. The cornerstone of this theory is the principle
of natural selection. Darwin noted that members of a spe-
cies were not all identical and that some of the differences
they exhibited were inherited by their offspring. If an in-
dividual’s characteristics permit it to reproduce more suc-
cessfully, some of the individual’s offspring will inherit
the favorable characteristics and will themselves produce
more offspring. As a result, the characteristics will become
more prevalent in that species. He observed that animal
breeders were able to develop strains that possessed par-
ticular traits by mating together only animals that pos-
sessed the desired traits. If artificial selection, controlled
by animal breeders, could produce so many varieties of
dogs, cats, and livestock, perhaps natural selection could
be responsible for the development of species. Over the
course of time in the natural world, it was the environ-
ment, not the choices of the animal breeder, that shaped
the process of evolution.
Darwin and his fellow scientists knew nothing about
the mechanism by which the principle of natural selection
works. In fact, the principles of molecular genetics were
not discovered until the middle of the twentieth century.
Briefly, here is how the process works: Every sexually
reproducing multicellular organism consists of a large
number of cells, each of which contains chromosomes.
Chromosomes are large, complex molecules that contain
the recipes for producing the proteins that cells need to
grow and to perform their functions. In essence, the chro-
mosomes contain the blueprints for the construction (that
is, the embryological development) of a particular mem-
ber of a particular species. If the plans are altered, a differ-
ent organism is produced.
The plans do get altered from time to time; mutations
occur. Mutations are accidental changes in the chromo-
somes of sperm or eggs that join together and develop
into new organisms. For example, a random mutation
of a chromosome in a cell of an animal’s testis or ovary
could produce a mutation that affects that animal’s off-
spring. Most mutations are deleterious; the offspring ei-
ther fails to survive or survives with detrimental effects of
the mutation. Some mutations are beneficial and confer a
selective advantage to the organism that possesses them.
An individual with a selective advantage is more likely
than other members of its species to live long enough to

M01_CARL0287_13_GE_C01.indd 24 25/11/21 8:06 AM


reproduce and pass on its chromosomes to its own off-
spring. Many different kinds of traits can confer a selec-
tive advantage: resistance to a particular disease, the abil-
ity to digest new kinds of food, more effective weapons
for defense or for procurement of prey, and even a more
attractive appearance to potential mates.
The traits that can be altered by mutations are physi-
cal ones; chromosomes make proteins, which affect the
structure and chemistry of cells. But the effects of these
physical alterations can be seen in an animal’s behavior.
This means that the process of natural selection can act
on behavior indirectly. For example, if a particular mu-
tation results in changes in the brain that cause a small
animal to change its behavior and freeze when it per-
ceives a nearby movement, that animal is more likely to
escape undetected when a predator passes nearby. This
tendency makes the animal more likely to survive and
produce offspring, passing on genes related to freezing
behavior to future generations.
Other mutations are not immediately favorable, but
because they do not put their possessors at a disadvan-
tage, they are inherited by at least some members of the
species. As a result of thousands of such mutations, the
members of a particular species possess a variety of genes,
and are all at least somewhat different from one another.
Variety is a definite advantage for a species. Different en-
vironments provide optimal habitats for different kinds
of organisms. When the environment changes, species
must adapt or run the risk of becoming extinct. If some
members of the species possess assortments of genes that
provide characteristics permitting them to adapt to the
new environment, their offspring will survive, and the
species will continue.
An understanding of the principle of natural selec-
tion plays some role in the thinking of every scientist
who undertakes research in behavioral neuroscience.
Some researchers explicitly consider the genetic mecha-
nisms of various behaviors and the physiological pro-
cesses on which these behaviors depend. Others are
concerned with comparative aspects of behavior and its
physiological basis; they compare the nervous systems
of animals from a variety of species to make hypotheses
about the evolution of brain structure and the behav-
ioral capacities that correspond to this evolutionary de-
velopment. But even though many researchers do not
directly study evolution, the principle of natural selec-
tion guides the thinking of behavioral neuroscientists.
We ask ourselves what the selective advantage of a par-
ticular trait might be. We think about how nature might
have used a physiological mechanism that already ex-
isted to perform more complex functions in more com-
plex organisms. When we entertain hypotheses, we ask
ourselves whether a particular explanation makes sense
in an evolutionary perspective.
M01_CARL0287_13_GE_C01.indd 25
Introduction 25
Evolution of Human Brains
LO 1.4 Identify factors involved in the evolution of
human brains.
To evolve means to develop gradually. The process of ­evolution
is a gradual change in the structure and ­physiology of a spe-
cies as a result of natural selection. New species evolve when
organisms develop novel characteristics that can take advan-
tage of unexploited opportunities in the environment.
Appearance of the earliest humans can be traced back
to the Cenozoic period, when tropical forests covered much
of the land areas. In these forests our most direct ancestors,
the primates, evolved. The first primates were small and
preyed on insects and small cold-blooded vertebrates such
as lizards and frogs. They had grasping hands that permit-
ted them to climb about in small branches of the forest. Over
time, larger species developed, with larger, forward-facing
eyes (and the brains to analyze what the eyes saw), which
facilitated moving among the trees and capturing prey.
The evolution of fruit-bearing trees provided an op-
portunity for fruit-eating primates. In fact, the original
advantage of color vision (and the associated sensory re-
gions of the brain) was probably the ability to discriminate
ripe fruit from green leaves in order to eat the fruit before
it spoiled—or some other animals got to it first. And be-
cause fruit is such a nutritious form of food, its availability
provided an opportunity that could be exploited by larger
primates, which were able to travel farther in quest of food.
The first hominids (humanlike apes) appeared in
Africa. They appeared not in dense tropical forests but in
drier woodlands and in the savanna. Our fruit-eating an-
cestors continued to eat fruit, but they evolved characteris-
tics that enabled them to gather roots and tubers as well, to
hunt and kill game, and to defend themselves against other
predators. They made tools that could be used to hunt,
produce clothing, and construct dwellings; they discov-
ered the many uses of fire; they domesticated dogs, which
greatly increased their ability to hunt and helped warn
of attacks by predators; and they developed the ability to
communicate symbolically, by means of spoken words.
Figure 1.8 shows the primate family tree. Our closest liv-
ing relatives—the only hominids besides ourselves who have
survived—are the chimpanzees, gorillas, and orangutans.
DNA analysis shows that genetically, there is very little dif-
ference between these four species. For example, humans
and chimpanzees share almost 99 percent of their DNA.
The first hominid to leave Africa did so around
1.7 million years ago. This species, Homo erectus (“upright
man”), scattered across Europe and Asia. One branch of
Homo erectus appears to have been the ancestor of Homo
neanderthalis, which inhabited Western Europe between
120,000 and 30,000 years ago. Neanderthals resembled
modern humans. They made tools out of stone and wood
25/11/21 8:06 AM
 26 Chapter 1
Figure 1.8 Evolution of Primate Species
Source: Redrawn from Lewin, R. (1993.) Human evolution: An illustrated introduction (3rd ed.). Boston: Blackwell Scientific Publications.
Reprinted with permission by Blackwell Science Ltd.
Present
Tarsier
10
20
New W
Time (million years ago)
Pro
orld
ims
ian
30
mon
s
key
s
40
50
and discovered the use of fire. Our own species, Homo
sapiens, evolved in East Africa around 100,000 years ago.
Some of our ancestors migrated to other parts of Africa and
out of Africa to Asia, Polynesia, Australia, Europe, and the
Americas. (See Figure 1.9.)
Humans possessed several characteristics that allowed
them to compete with other species. Their agile hands en-
abled them to make and use tools. Their excellent color vi-
sion helped them to spot ripe fruit, prey, and dangerous
predators. Their mastery of fire enabled them to cook food,
provide warmth, and frighten nocturnal predators. Their
upright posture and bipedalism (ability to walk using
two rear limbs) made it possible for them to walk long
distances efficiently, with their eyes far enough from the
ground to see long distances across the plains. Bipedalism
also permitted them to carry tools and food with them,
which meant that they could bring fruit, roots, and pieces
of meat back to their tribe. Their linguistic abilities enabled
them to combine the collective knowledge of all the mem-
bers of the tribe, to make plans, to pass information on to
subsequent generations, and to form complex civilizations
that established their status as the dominant species. All of
these characteristics required a primate brain capable of
these complex abilities.
Sophisticated primate brains developed within the
constraints of the size of a mother’s birth canal, and an
M01_CARL0287_13_GE_C01.indd 26
Human
Orangutan Gorilla
Baboon
Spider
monkey Colobus
monkey Chimpanzee
s
ey
nk
s
id
mo
ino Gibbon
orld
m
Ho
Old W
upright posture limits the size of a woman’s birth canal.
A newborn primate’s head is about as large as it can
safely be. Because a baby’s brain is not large or complex
enough to perform the physical and intellectual abilities
of an adult, the brain must continue to grow after the
baby is born. In fact, all mammals (and all birds, for that
matter) require parental care for a period of time while
the nervous system develops. The fact that young mam-
mals (particularly young humans) are guaranteed to be
exposed to the adults who care for them means that a
period of apprenticeship is possible. Consequently, the
evolutionary process did not have to produce a brain
that consisted solely of specialized circuits of neurons
that performed specialized tasks. Instead, it produced a
primate brain with an abundance of neural circuits that
could be modified by experience. Adults would nourish
and protect their offspring and provide them with the
skills they would need as adults. Some specialized cir-
cuits were necessary (for example, those involved in ana-
lyzing the complex sounds we use for speech), but, by
and large, the primate brain is more similar to a general-
purpose, programmable computer.
What counts, as far as intellectual ability goes, is
having a brain with plenty of neurons that are available
for behavior, learning, remembering, reasoning, and
making plans. Herculano-Houzel and colleagues (2007)
25/11/21 8:06 AM
 Introduction 27

Figure 1.9 Migration of Homo sapiens

The figure shows proposed migration routes of Homo sapiens after evolution of the species in East Africa.
Source: Redrawn with permission from Cavalli-Sforza, L. L. (1991.) Genes, peoples and languages. Scientific American, 265(5), p. 75.

15–35,000 years ago

40,000 years ago

60,000(?) years ago

100,000 years ago

Equator

50–60,000(?) years ago

compared the brains of several species of rodents and
primates and found that primate brains contain more
neurons per gram than rodent brains. (See Figure 1.10.)
Reflecting on their results, the researchers concluded
that “our standing among primates as the proud owners
of the largest living brain assures that, at least among
primates, we enjoy the largest number of neurons from
which to derive cognition and behavior as a whole”
(Herculano-Houzel, 2009, p. 10). Can you predict what
types of functions these additional neurons might be de-
voted to in humans?
What types of genetic changes were responsible for
the evolution of the human brain? This question will be
addressed in more detail in Chapter 15, but evidence sug-
gests that the most important principle is slowing the pro-
infantile characteristics, including their disproportionate
size relative to the rest of the body.
Figure 1.10 Comparison of Mammalian Brains
Species with more complex behaviors have brains with more
neurons that are available for behavior, learning, remembering,
reasoning, and making plans. Primate brains contain more neurons
per gram than rodent brains and more neurons in the cortex.
Source: Herculano-Houzel, S., and Marino, L. (1998.) A Comparison of
Encephalization between Odontocete Cetaceans and Anthropoid Primates.
Brain, Behavior and Evolution, 51(4), 230–238.
Number of cerebral cortex neurons (billions)

16
cess of brain development, allowing more time for growth.
As we will see, the prenatal period of cell division in the 14
brain is prolonged in humans, which results in a brain that 12
weighs an average of 350 g and contains approximately
10
86 billion neurons (Azevedo et al., 2009). After birth the
brain continues to grow. Production of new neurons al- 8
most ceases, but those that are already present grow and
6
establish connections with each other, and other brain cells,
which protect and support neurons, begin to proliferate. 4
Not until late adolescence does the human brain reach its 2
adult size of approximately 1,400 g—about four times the
weight of a newborn’s brain. This prolongation of matura- nt et ue a ee s
ha os aq rill nz an
tion is known as neoteny (roughly translated as “extended p rm c Go a
Hu
m
Ele Ma Ma imp
youth”). The mature human head and brain retain some Ch

M01_CARL0287_13_GE_C01.indd 27 25/11/21 8:06 AM

 28 Chapter 1
Module Review: Natural Selection and Evolution
Functionalism and the Inheritance of Traits
LO 1.3 Describe the role of natural selection in the
evolution of behavioral traits.
Natural selection is the process responsible for evolution
of structures with specific functions. Members of a species
possess a variety of structures. If the structures permit an
individual to reproduce more successfully, its offspring will
also have these structures, and they will become more prev-
alent in the population. An example of inherited structures
responsible for behavior is the set of brain structures respon-
sible for male song behavior in some species of songbirds.
Evolution of Human Brains
LO 1.4 Identify factors involved in the evolution of
human brains.
The evolution of specialized structures responsible
for functions such as color vision, fine motor control,
Ethical Issues in Research
with Humans and Other
Animals
This book contains many facts about what is currently
known about the structure and function of the nervous sys-
tem. Where do these facts come from? They are the result of
carefully designed experiments that can include computer
simulations, individual cells, and often humans and other
animals. Neuroscience research involving humans and other
animals is subject to important ethical considerations and
oversight. This section addresses these issues in more detail.
Research with Animals
LO 1.5 Outline reasons for the use of animals in
behavioral neuroscience research.
Much of the research described in this book involves ex-
perimentation on living animals. Any time we use another
species of animals for our own purposes, we should be sure
that what we are doing is both humane and worthwhile. It is
important that a good case can be made that research in
behavioral neuroscience qualifies on both counts. Humane
treatment is a matter of procedure. We know how to main-
tain laboratory animals in good health in comfortable, sani-
tary conditions. We know how to administer anesthetics
and analgesics so that animals do not suffer during or after
M01_CARL0287_13_GE_C01.indd 28
complex vision, and language required a more com-
plex primate brain. Primate brains contain many more
neurons per gram than other species. These additional
cells are responsible for behavior, learning, remember-
ing, reasoning, and making plans. Additional brain
development occurs after birth and throughout an
extended period of development and parental care in
humans.
Thought Question
Kavoi and Jameela (2011) reported that a part of the brain
responsible for olfaction, the olfactory bulb, is larger in
dogs than in humans, even after accounting for differ-
ences in overall brain size. Using the principles of natu-
ral or artificial selection, hypothesize how dogs came to
have this larger structure in their brain and predict how it
might impact their behavior.
surgery, and we know how to prevent infections with proper
surgical procedures and the use of antibiotics. Most indus-
trially developed societies have strict regulations about the
care of animals and require approval of the experimental
procedures that are used on them. There is no excuse for
mistreating animals in our care. In fact, the vast majority of
laboratory animals are treated humanely and many animal
researchers are also strong animal welfare advocates.
Whether an experiment is worthwhile can be difficult to
say. We use animals for many purposes. We eat their meat
and eggs, and we drink their milk; we turn their hides into
leather; we extract insulin and other hormones from their
organs to treat people’s diseases; we train them to do use-
ful work on farms or to entertain us. Even having a pet is
a form of exploitation; it is we—not they—who decide that
they will live in our homes. The fact is we have been using
other animals throughout the history of our species.
Pet ownership has the potential to cause much more
suffering among animals than scientific research does.
Pet owners are not required to receive permission from a
board of experts that includes a veterinarian to house their
pets, nor are they subject to periodic inspections to be sure
that their home is clean and sanitary, that their pets have
enough space to exercise properly, or that their pets’ diets
are appropriate. Scientific researchers are required to have
all those things. The disproportionate amount of concern
that animal rights activists show toward the use of ani-
mals in research and education is puzzling, particularly
because this is the one indispensable use of animals. We
25/11/21 8:06 AM

can survive without eating animals, we can live without
hunting, we can do without furs; but without using ani-
mals for research and for training future researchers, we
cannot make progress in understanding and treating dis-
eases. In not too many years scientists will probably have
developed a vaccine that will prevent the further spread
of diseases such as Ebola virus disease, malaria, or AIDS.
Even diseases that we have already conquered would
impact new lives if drug companies could no longer use
animals to develop and test new treatments. If they were
deprived of animals, these companies could no longer
extract hormones used to treat human diseases, and they
could not prepare many of the vaccines we now use to
prevent disease.
Our species is beset by medical, psychological, and
behavioral problems, many of which can be solved only
through biological research. Consider some of the major
neurological disorders. Strokes, like the one experienced
by Jeremiah at the beginning of this chapter, are caused
by bleeding or obstruction of a blood vessel within the
brain, and often leave people partly paralyzed, unable
to read, write, or converse with their friends and ­family.
Basic animal research on the means by which nerve cells
communicate with each other has led to important dis-
coveries about the causes of the death of brain cells. This
research was not directed toward a specific practical
goal; the potential benefits actually came as a surprise to
the investigators.
Experiments based on these results have shown that
if a blood vessel leading to the brain is blocked for a few
minutes, the part of the brain that is nourished by that ves-
sel will die. However, the brain damage can be prevented
by first administering a drug that interferes with a particu-
lar kind of neural communication. This research is impor-
tant, because it may lead to medical treatments that can
help to reduce the brain damage caused by strokes. But it
involves operating on a laboratory animal, such as a rat,
and pinching off a blood vessel. (The animals are anesthe-
tized.) Some of the animals will sustain brain damage, and
all will be euthanized so that their brains can be examined.
However, you will probably agree that research like this is
just as legitimate as using animals for food.
As you will learn later in this book, research with
laboratory animals has produced important discoveries
about the possible causes or potential treatments of neu-
rological and mental disorders, including Parkinson’s dis-
ease, schizophrenia, bipolar disorder, anxiety disorders,
­obsessive-compulsive disorder, anorexia nervosa, obesity,
and substance abuse. Although much progress has been
made, these problems persist, and they cause much human
suffering. Unless we continue our research with laboratory
animals, they will not be solved.
Some people have suggested that instead of using
laboratory animals in our research, we could use tissue
M01_CARL0287_13_GE_C01.indd 29
Introduction 29
cultures or computers. While these techniques can be used
to pursue some research questions, unfortunately, tissue
cultures or computers are not substitutes for complex, liv-
ing organisms. We have no way to study behavioral prob-
lems such as substance abuse in tissue cultures, nor can we
program a computer to simulate the workings of an ani-
mal’s nervous system. (If we could, that would mean we
already had all the answers.)
OVERSIGHT OF ANIMAL RESEARCH
LO 1.6 Identify mechanisms for oversight of animal
research.
In the United States, any institution that receives federal
research funding to use animals in research is required
to have an Institutional Animal Care and Use Committee
(IACUC). The IACUC is typically composed of a veteri-
narian, scientists who work with animals, non-scientist
members, and community members not affiliated with the
institution. This group reviews all proposals for research in-
volving animals, with the intent of ensuring humane and
ethical treatment of all animals involved. Even noninvasive
research with animals (such as fieldwork or observational
studies) must pass review and be approved by the IACUC.
This approval process ensures not only the welfare of the
animals, but also that the research is compliant with local,
state, and federal regulations.
Research with Humans
LO 1.7 Discuss ethical considerations in research with
human participants.
Not all neuroscience research is conducted with animal
models. Much of what we currently understand about
the brain and behavior is the result of research with
human participants. In addition to humane research
conditions, research with human participants must also
include informed consent and precautions to protect the
identity of the participants. Informed consent describes
the process in which researchers must inform any poten-
tial participant about the nature of the study, how any
data will be collected and stored, and what the antici-
pated benefits and costs of participating will be. Only
after obtaining this information can the participant make
an informed decision about whether to participate in a
study. Violating the informed consent process can have
ethical, legal, and financial consequences. In 2010, the
case of Havasupai Tribe v. Arizona Board of Regents was
settled, including the return of biological samples and
a payment of $700,000 to the Havasupai tribe after six
years of dispute. The settlement was issued in response
to a vague and incomplete informed consent process that
resulted in the use of blood samples originally intended
for research on diabetes being used in contested research
25/11/21 8:06 AM
 30 Chapter 1

involving factors related to schizophrenia (Van Assche et

Figure 1.11 Behavioral Neuroscience Research with
al., 2013). Protecting the identity of participants is cru- Human Participants
cial for all research with human participants, and par- Researchers work with volunteers to learn more about the brain
ticularly important in behavioral neuroscience research mechanisms responsible for functions such as emotion, learning,
investigating potentially sensitive topics (for example, memory, and behavior.
the use of illicit drugs in studies of brain changes in sub-
stance abuse).
An emerging interdisciplinary field, neuroethics, is
devoted to better understanding implications of and de-
veloping best practices in ethics for neuroscience research
with human participants. A 2014 report from a panel of
national experts explored the ethical challenges of neu-

Inpix Creative/Indiapicture/Alamy Stock Photo

roscience research by investigating (1) neuroimaging and
brain privacy; (2) dementia, personality, and changed pref-
erences; (3) cognitive enhancement and justice; and (4)
deep brain stimulation research and the ethically difficult
history of psychosurgery (Presidential Commission for the
Study of Bioethical Issues, 2014). The panel recommenda-
tions included integrating ethics and science through edu-
cation at all levels.

OVERSIGHT OF HUMAN RESEARCH

LO 1.8 Identify mechanisms for oversight of human
research.
Much like animal research, research with human volun-
teers is essential to advancing our knowledge of the brain
in health and disease. Also similar to animal research, work
with human participants is subject to strict regulation and
must be reviewed and approved by a board of experts and
laypeople. The Institutional Review Board (IRB) functions
similarly to the IACUC to ensure ethical treatment of vol-
unteers in research. (See Figure 1.11.) The IRB is typically
composed of scientific experts, laypeople, and members
of the community. This group is tasked with protecting
human research participants.

Module Review: Ethical Issues in Research with Humans

and Other Animals
Research with Animals
LO 1.5 Outline reasons for the use of animals in
behavioral neuroscience research.
Animals are used in behavioral neuroscience research to
improve understanding of the nervous system and de-
velop treatments for disease and injury. Animal models
are used when it is not possible or it is inappropriate to
conduct research with human participants and when cell
models or computer programs cannot simulate the com-
plexity of the nervous system.
LO 1.6 Identify mechanisms for oversight of animal
research.
The humane treatment of research animals is governed
by local, state, and federal regulations. The IACUC is
tasked with reviewing animal research proposals and
protecting the welfare of animals in research.
Research with Humans
LO 1.7 Discuss ethical considerations in research with
human participants.
Ethical considerations for research involving human par-
ticipants include protections such as informed consent
and confidentiality. The field of neuroethics is devoted
to better understanding implications of and developing
best practices in ethics for neuroscience research with hu-
man participants.
LO 1.8 Identify mechanisms for oversight of human
research.
The IRB is responsible for the protection of human re-
search participants. It is composed of scientific experts,
laypeople, and community members. The IRB reviews
proposals for research involving people.

M01_CARL0287_13_GE_C01.indd 30 25/11/21 8:06 AM

 Introduction 31

Thought Question by society. For example, if you wish to create a drug or a

­device that would eliminate aggression in humans, iden-
Behavioral physiology studies can lead to scientific and tify the ethical challenges you would encounter. Think
technological advances that would allow manipulat- about the main ethical issue in this case and ask your
ing certain behaviors that are often seen as unwanted friends if they agree or not.

The Future of using advanced brain-imaging techniques to study the

activity of various regions of the brain as a person partici-
Neuroscience: Careers and pates in various behaviors. This research is also carried
out by cognitive neuroscientists—researchers with a
Strategies for Learning Ph.D. (usually in psychology) and specialized training in
the principles and procedures of neurology.
What is behavioral neuroscience, and what do behavioral
Most professional behavioral neuroscientists have re-
neuroscientists do? What are the best ways to learn more
ceived a Ph.D. from a graduate program in psychology or
about this diverse and exciting field? By the time you fin-
from an interdisciplinary program. Programs can include
ish this book, you will have a much richer answer to these
faculty members from departments such as psychology, bi-
questions. The next section will describe the field—and ca-
ology, chemistry, biochemistry, or computer science. Most
reers open to those who specialize in it. Likewise, we want
professional behavioral neuroscientists are employed by
to provide you with some strategies to help you learn as
colleges and universities, where they are engaged in teach-
you study this fascinating discipline.
ing and research. Others are employed by institutions de-
voted to research—for example, in laboratories owned and
Careers in Neuroscience operated by national governments or by private philan-
LO 1.9 Identify careers in behavioral neuroscience. thropic organizations. A few work in industry, usually for
pharmaceutical companies that are interested in assessing
Behavioral neuroscience belongs to a larger field that
the effects of drugs on behavior.
is simply called neuroscience. Neuroscientists concern
To become a professor or independent researcher,
themselves with all aspects of the nervous system: its
one must receive a doctorate—usually a Ph.D., although
anatomy, chemistry, physiology, development, and func-
some people turn to research after receiving an M.D.
tioning. The research of neuroscientists ranges from the
Most behavioral neuroscientists spend two years or more
study of molecular genetics to the study of social be-
in a postdoctoral position after completing their gradu-
havior. Behavioral neuroscientists study all behavioral
ate degree, working in the laboratory of a senior scientist
phenomena that can be observed in humans and ani-
to gain more research experience. During this time they
mals. They attempt to understand the role of the nervous
write articles describing their research findings and sub-
system, interacting with the rest of the body (especially
mit them for publication in scientific journals. These pub-
the endocrine system, which secretes hormones), in
lications are an important factor in obtaining an indepen-
controlling behavior. They study such topics as sensory
dent position.
processes, sleep, emotional behavior, ingestive behavior,
Not all people who are engaged in neuroscience re-
aggressive behavior, sexual behavior, parental behavior,
search have doctoral degrees. Research technicians with
and learning and memory. They also study animal mod-
bachelor’s or master’s level degrees perform essential—
els of symptoms that humans experience, such as anxiety,
and intellectually rewarding—services working with se-
depression, phobia, and obsessions and compulsions.
nior scientists. Technicians can continue to gain experience
Although the original name for the field described in
and education on the job, enabling them to assume respon-
this book was physiological psychology, several other terms
sibility for managing and completing projects indepen-
are now in general use, such as biological psychology, bio-
dently. (See Figure 1.12).
psychology, psychobiology, and—the most common one—
behavioral neuroscience.
Two other fields often overlap with that of behav- Strategies for Learning
ioral neuroscience: neurology and cognitive neuroscience.
Neurologists are physicians who diagnose and treat LO 1.10 Describe effective learning strategies for
diseases of the nervous system. Most neurologists are studying behavioral neuroscience.
solely involved in the practice of medicine, but some The brain is a complicated organ. After all, it is responsible
engage in research. They study the behavior of people for all our abilities and all our complexities. Scientists have
whose brains have been damaged by natural causes, been studying this organ for many years and (especially in

M01_CARL0287_13_GE_C01.indd 31 25/11/21 8:06 AM

 32 Chapter 1

Figure 1.12 Pursuing a Research Career in Neuroscience

What kinds of training are required for a career in neuroscience? Where do neuroscientists work? Explore this timeline to learn more.

Time Period Description

High School Students interested in neuroscience may take courses
in biology, chemistry, psychology, or other sciences in
high school.

SDI Productions/E+/Getty Images

College Students interested in neuroscience may study biology,
chemistry, psychology, neuroscience, or other related
areas. Some students work as research assistants in
laboratories and develop mentored relationships with
researchers. College graduates interested in neurosci-
ence can work as research technicians or assistants.

WavebreakMediaMicro/Fotolia
Graduate Training Students can pursue advanced graduate t­raining for
one or more years after college. Graduate ­training
typically involves advanced coursework and more
­independent research. Graduate students are e ­ xpected
to conduct research (with the guidance of a research
mentor) and disseminate the results of their work.
After completing a graduate program, individuals may
teach in a secondary or postsecondary institution,
­conduct research, or work in industry.

Jean-Marie Guyon/123RF
Postgraduate Training Postgraduate positions are more independent and
often involve additional training in specialized research
areas or with specialized research techniques. After
completing postgraduate training, individuals may
teach in a secondary or postsecondary institution,
­conduct research, or work in industry.
Tyler Olson/123RF

M01_CARL0287_13_GE_C01.indd 32 25/11/21 8:06 AM


recent years) have been learning a lot about how it works.
It is impossible to summarize this progress in a few simple
sentences; therefore, this book contains a lot of informa-
tion. We have tried to organize this information logically,
telling you what you need to know in the order in which
you need to know it. (To understand some things, you
sometimes need to understand other things first.) We have
also tried to write as clearly as possible, making examples
as simple and as vivid as we can. Still, you cannot expect
to fully understand the information in this book by simply
giving it a passive read; you will have to do some work.
Learning about behavioral neuroscience involves much
more than memorizing facts. Of course, there are facts to be
memorized: names of parts of the nervous system, names of
chemicals and drugs, scientific terms for particular phenom-
ena and procedures used to investigate them, and so on. But
the quest for information is nowhere near completed; we
know only a small fraction of what we have to learn. And
almost certainly, many of the “facts” that we now accept will
someday be shown to be incorrect. If all you do is learn facts,
where will you be when these facts are revised?
Our goal is to offer some practical advice about studying.
You have been studying throughout your academic career,
and you have undoubtedly learned some useful strategies
along the way. Even if you have developed efficient and
effective study skills, at least consider the possibility that
there might be some ways to improve them. This section is
intended to provide you with suggestions to maximize your
learning about behavioral neuroscience. These suggestions
are supported by empirical research on learning.
• Write notes that organize information into meaning-
ful groups; don’t just highlight. Connecting new infor-
mation to prior knowledge is an important means for
learning. To do this will require actively thinking about
the new information at hand and finding ways to link
it to your current understanding. This is an active and
involved process that will take some time and effort.
Highlighting or underlining without combining the in-
formation into your own notes is passive and does not
facilitate learning and retention the way that writing
or typing your own notes does. Previous research has
demonstrated that highlighting and underlining alone
do not improve test scores, and in some cases may even
be detrimental to learning (Dunlosky et al., 2013).
• Teach yourself by teaching someone else. After read-
ing a section or chapter, consider how you would
teach the information to someone else—a classmate,
a friend, or maybe a curious family member. This ac-
tivity will help you to think about the most important
aspects of the section. Nestojko and colleagues (2014)
found that students who prepared to teach others
about the content of a complex reading assignment
performed better on a later test than students who had
prepared themselves for a test on the reading.
M01_CARL0287_13_GE_C01.indd 33
Introduction 33
• Study in the environment you will be tested in or vary
your study environments. State-dependent learning the-
ory says that information learned in one environment is
most readily recalled in the same environment. The ratio-
nale behind this performance-boosting effect of environ-
ment is that the context (e.g., the color of the walls, the
chair, the people around you) provides important cues
that help you recall what was previously learned in that
environment. If you’re not able to study in the same en-
vironment as you will be tested, you can try to incorpo-
rate as many of the same elements as possible (e.g., use
the same computer, pens, procedure for note taking, etc.)
or you can study in many different environments (e.g.,
at home, in the student union, in your residence hall) so
that you will not become dependent on any one single
cue or set of cues when you are tested. In an interesting
test of state dependent learning, Godden and Baddeley
(1975) tested college student scuba divers on information
they read while underwater or on land. Students recalled
information learned underwater the best in an underwa-
ter test. The students performed most poorly on the tests
of information in a different context (for example, infor-
mation learned underwater but tested on land).
• Study with the absolute minimum of distractions.
Your brain works best when it focuses on one chal-
lenging task (like learning about neuroscience!) at
a time (Hattie and Yates, 2014). Turn off televisions,
social media, and phones whenever possible, and try
to study in a quiet environment. Lee and colleagues
(2012) assigned college students learning about sci-
ence, history, and politics to three groups: reading in
silence, reading with a TV show playing in the back-
ground that students could ignore, and reading with
a TV show playing in the background that students
would later be tested on so that they would be sure to
pay attention to both the TV show and their assigned
study material. Students were instructed to read and
answer multiple choice questions. As you might have
guessed, students who tried to read and pay attention
to the TV show performed the worst on the test.
• Don’t cram! Spread out your study sessions.
Studying new information in two shorter but sepa-
rated sessions leads to more effective recall than
studying in one long session. Don’t cram. Instead,
plan to study something new once, then study it again
a different day before being asked to recall or apply
it on a final test or assignment. Though you should
plan your own study sessions around your schedule
and based on assignment or test due dates in your
class, some cognitive spacing has already been built
into this book for you. While there is no “one size fits
all” time period for spacing out reading and study
sessions, one to several days is a good rule of thumb
(Carpenter et al., 2012).
25/11/21 8:06 AM
 34 Chapter 1
• Study the most challenging topic first or last. Classic
studies in psychology revealed that when people were
asked to learn long lists of words, the first words learned
(the primacy effect) and the last words learned (the recency
effect) were the most likely to be recalled. The same prin-
ciples can hold true for learning about behavioral neuro-
science. For example, if you are reading about the cortex,
the thalamus, and the meninges in Chapter 3, and you
already know most of the parts of the meninges, but are
not feeling confident about your understanding of the
cortex and thalamus, plan to study the cortex first, then
the meninges and finally the thalamus information.
• Use mnemonics. Mnemonics are shortcuts for helping
retain new information. For example, you could try
story chaining by inventing a short story to link together
discrepant items; method of loci to use images of physi-
cal locations enabling you to position items along an
imaginary walk; and acrostics to use a word to represent
a list (such as FPOT for the lobes of the cortex: frontal,
parietal, occipital, temporal) (Hattie and Yates, 2014).
• Draw a picture. Trying to learn a new term or concept?
Try drawing a picture of it. Drawing improved memory
for new information compared to copying definitions, im-
proved memory in younger and older learners, and was
effective when people spent only four seconds drawing.
Researchers attribute the improved learning to the involve-
ment of motor activity and active elaboration involved in
drawing (Fernandes et al., 2018; Meade et al., 2018).
How this book is organized:
• The text, animations, interactives, and illustrations are
integrated as closely as possible. In our experience,
Module Review: The Future of Neuroscience: Careers and
Strategies for Learning
Careers in Neuroscience
LO 1.9 Identify careers in behavioral neuroscience.
Researchers in this area work in the fields of general neu-
roscience, behavioral neuroscience, and cognitive neuro-
science. Neurologists are physicians who specialize in the
nervous system. Individuals pursuing careers in neuro-
science typically work in academia or industry and often
pursue graduate education.
Strategies for Learning
LO 1.10  Describe effective learning strategies for
studying behavioral neuroscience.
Active strategies for learning are most effective. Taking
notes, practicing teaching or sharing information with
another person, making sure your study and test-taking
M01_CARL0287_13_GE_C01.indd 34
one of the most frustrating aspects of reading some
books is not knowing when to look at an illustration.
Here, everything is presented to you as you need it.
• Each chapter begins with a case study that profiles a
person’s real-life experience and a list of learning ob-
jectives. The case studies are meant to personalize and
make more relatable the concepts we will discuss in the
chapter. The learning objectives are included to help
you focus on the key ideas included in the chapter.
• You will notice that some words in the text are italicized,
and others are printed in boldface. Italics mean that ei-
ther the word is being stressed for emphasis or it is a
new term. Terms in bold (listed in the glossary of this
book) are key terms that are part of the vocabulary of
the behavioral neuroscientist. You will see many of these
terms used again in later chapters.
• At the end of each section, you will find two differ-
ent types of review activities: module reviews and
thought questions. The module reviews will remind
you of key points from the chapter, and the thought
questions will challenge you to apply what you have
learned to a new context or to expand your thinking
on a relevant topic. Finally, there are chapter review
questions at the end of each chapter to help you assess
your understanding of the concepts.
Now that you have a sense of what the field of behav-
ioral neuroscience entails, welcome to the rest of this book!
The next chapter starts with the structure and functions of
neurons and supporting cells, the most important elements
of the nervous system.
environment share some common features, studying
with as few distractions as possible, spacing out your
study sessions, carefully planning when to study chal-
lenging material, and using mnemonics whenever pos-
sible can enhance your learning.
Thought Question
What is it like to work as a behavioral neuroscientist?
Conduct an online search and locate a job advertisement
for a position in behavioral neuroscience. Read the job
description and qualifications carefully. What qualifica-
tions are required for the job? Why do you think these
experiences or this training is required? What kinds of
responsibilities and activities will the person in this posi-
tion engage in?
25/11/21 8:06 AM

Chapter Review Questions
1. Explain how generalization and reduction relate to
behavioral neuroscience research.
2. Identify the key milestones in the history of
­behavioral neuroscience.
3. Explain the principle of natural selection and its
­application in behavioral neuroscience research.
4. Evaluate the role of laboratory animals in behavioral
neuroscience research.
5. Explain why scientists have to use humans for
­research, considering there are other models.
M01_CARL0287_13_GE_C01.indd 35
Introduction 35
6. Identify important ethical considerations of
­behavioral neuroscience research involving human
participants.
7. Describe the steps you would need to take if you
­decided to become a research group leader in the field
of behavioral neuroscience.
8. Describe career opportunities in neuroscience.
9. Describe how you could apply an effective study
strategy to learning about behavioral neuroscience.
25/11/21 8:06 AM
 Chapter 2
Structure and Functions of
Cells of the Nervous System

Biophoto Associates/Science Source

Neurons are the cells of the nervous system that are specialized for communication.

Chapter Outline
Cells of the Nervous System Communication Between Neurons
The Nervous System: An Overview Structure of Synapses
Neurons Release of Neurotransmitters
Supporting Cells Activation of Receptors
The Blood–Brain Barrier Postsynaptic Potentials
Communication Within a Neuron Effects of Postsynaptic Potentials: Neural
Integration
Neural Communication: An Overview
Termination of Postsynaptic Potentials
Electrical Potentials of Axons
Autoreceptors
The Membrane Potential
Other Types of Synapses
The Action Potential
Nonsynaptic Chemical Communication
Conduction of the Action Potential

36

M02_CARL0287_13_GE_C02.indd 36 08/11/2021 14:18


Learning Objectives
LO 2.1 Contrast features of the central and
peripheral nervous systems.
LO 2.2 Distinguish among the structures of a
neuron.
LO 2.3 Compare supporting cells in the central
and peripheral nervous systems.
LO 2.4 Assess the function of the blood–brain
barrier.
LO 2.5 Explain the process of neural
communication in a reflex.
LO 2.6 Describe membrane potential,
resting potential, hyperpolarization,
depolarization, and the action potential.
LO 2.7 Summarize how diffusion, electrostatic
pressure, and the sodium–potassium
pump help establish membrane
potential.
LO 2.8 Summarize the series of ion movements
during the action potential.
LO 2.9 Describe conduction of the action potential.
Kathryn was getting desperate. She was healthy, ate well, and
stayed active with sports and regular exercise. She went to the
gym almost every day for cardio classes and swimming. But
lately she had been having trouble keeping up with her usual
schedule. At first, she started getting tired toward the end of her
exercise classes. Her arms, particularly, seemed to get heavy.
Then when she started swimming, it was hard to lift her arms
over her head. She did not have any other symptoms, so she
told herself that she needed more sleep.
Over the next few weeks, things only got worse. Her exercise
classes were more and more difficult to complete. Her instructor
became concerned and suggested that Kathryn see her doctor.
She made an appointment, but her doctor found nothing wrong
with her. She was not sick, showed no signs of an infection, and
seemed to be generally healthy. Her doctor asked how things were
going at work. Kathryn explained that she had been experiencing
a particularly stressful month at her job. Kathryn and her physician
agreed that increased stress could be the cause of her problem.
The doctor did not prescribe any medication, but asked Kathryn to
make another appointment if she did not feel better soon.
She did feel better for a while, but then all of a sudden her
symptoms got worse. She quit going to the gym and even had
trouble finishing a day at work. One afternoon she tried to look
up at the clock on the wall and realized that she could hardly
M02_CARL0287_13_GE_C02.indd 37
Structure and Functions of Cells of the Nervous System 37
LO 2.10 Identify the presynaptic structures
involved in synaptic communication.
LO 2.11 Describe the process of neurotransmitter
release.
LO 2.12 Contrast ionotropic and metabotropic
receptors.
LO 2.13 Compare EPSPs and IPSPs in
postsynaptic cells.
LO 2.14 Summarize neural integration of EPSPs
and IPSPs.
LO 2.15 Explain how postsynaptic potentials are
terminated.
LO 2.16 Distinguish autoreceptors from
postsynaptic receptors.
LO 2.17 Identify synapses other than those
involved in neural integration.
LO 2.18 Describe examples of nonsynaptic
communication.
see—her eyelids were drooping, and her head felt incredibly
heavy. Just then, one of her supervisors came over and asked
her to report on the progress she had made on a new project.
As she spoke, she found herself getting weaker and weaker. It
even felt as if breathing took a lot of effort. She managed to fin-
ish the conversation, but immediately afterward she went home.
She called her physician, who arranged for her to go to the
hospital to be seen by a neurologist. The neurologist listened to a
description of Kathryn’s symptoms and examined her briefly. The
neurologist thought she might know what was wrong. She pre-
pared an injection and gave it to Kathryn. She started questioning
Kathryn about her job. Kathryn answered slowly, her voice almost
a whisper. As the questions continued, she realized that it was get-
ting easier and easier to talk. She straightened her back and took
a deep breath. She stood up and raised her arms above her head.
“Look,” she said, her excitement growing. “I can do this again. I’ve
got my strength back! What did you give me? Am I cured?”
The neurologist smiled ruefully. “I wish it were that simple,”
she said. “No, I’m afraid you aren’t cured, but now we know
what is causing your weakness. There is a treatment. The in-
jection I gave you lasts only for a few minutes, but I can give
you some pills that have effects that last much longer.” Indeed,
as she was talking, Kathryn felt herself weakening, and she sat
down again.
08/11/2021 14:18
 38 Chapter 2
All we are capable of doing—perceiving, thinking, learning,
remembering, acting—is made possible by the integrated
activity of the cells of the nervous system. To understand
how the nervous system controls behavior, we need to un-
derstand its parts—the cells that compose it. In Kathryn’s
case, the cells of her nervous system were not functioning
appropriately, causing her symptoms of fatigue. Kathryn
was diagnosed with myasthenia gravis. The term liter-
ally means “grave muscle weakness.” It is an uncommon
disorder, but many experts believe that mild cases go
undiagnosed. Although there are drug treatments for my-
asthenia gravis, unfortunately there is currently no cure.
Developing new treatment options could benefit more
people like Kathryn and improve the quality of their lives
(Binks et al., 2016; Breiner et al., 2016).
Myasthenia gravis is an autoimmune disease. For un-
known reasons the immune system breaks down proteins
in the nervous system that allow cells to receive messages.
Understanding the structure and function of the cells of
the nervous system allowed the neurologist to diagnose
and treat Kathryn. Kathryn’s case highlights many of the
key topics you will learn about in this chapter, including
communication within and between cells of the nervous
system. To learn more about the specific cells involved in
myasthenia gravis, look ahead to the section “Termination
of the Postsynaptic Potentials” on acetylcholine.
Cells of the Nervous
System
Just how many nerve cells, or neurons, are there in the
human brain? The best estimate is around 86 billion neu-
rons. Our brains contain approximately the same num-
ber of non-neuronal cells too (Azevedo et al., 2009; von
Bartheld et al., 2016). The rest of the human nervous sys-
tem contains even more cells. The cells comprise the two
basic divisions of the nervous system: the central nervous
system and the peripheral nervous system.
The Nervous System: An Overview
LO 2.1 Contrast features of the central and peripheral
nervous systems.
The central nervous system (CNS) consists of the brain
and the spinal cord. The peripheral nervous system (PNS)
consists of the nerves and most of the sensory organs. (See
Figure 2.1.)
The CNS communicates with the rest of the body
through nerves attached to the brain and to the spinal cord.
Nerves are bundles of thousands of individual neurons,
all wrapped in a tough, protective membrane. Under a
microscope, nerves look something like electrical cables,
M02_CARL0287_13_GE_C02.indd 38
Figure 2.1 The Central and Peripheral Nervous
Systems
The central nervous system (in pink) includes the brain and spinal
cord. The peripheral nervous system (in blue) includes all of the
nerves that relay information between the central nervous system
and the rest of the body.
Brain
Spinal cord
Nerves
containing many bundles of wires. (See Figure 2.2.) Like
the individual wires in a cable, nerve fibers transmit mes-
sages through the nerve, from a sense organ to the brain or
from the brain to a muscle or gland.
Information, in the form of light, sound, odors, tastes,
or contact with objects, is gathered from the environment
by specialized cells of the PNS called sensory neurons.
Motor behavior requires contracting muscles, which are
controlled by motor neurons in the PNS. Between sensory
neurons and motor neurons are the interneurons—neurons
that lie entirely within the CNS. Local interneurons form
circuits with nearby neurons and analyze small pieces of
information. Relay interneurons connect circuits of local
interneurons in one region of the brain with those in other
regions. Through these connections, circuits of neurons
throughout the brain perform functions essential to tasks
such as perceiving, learning, remembering, deciding, and
controlling complex behaviors. (See Figure 2.3.)
The rest of this module describes the most important
cells of the nervous system—neurons and their supporting
cells—and the blood–brain barrier, which chemically iso-
lates cells in the CNS from the rest of the body.
08/11/2021 14:18
 Structure and Functions of Cells of the Nervous System 39

Neurons come in many shapes and varieties, according

Figure 2.2 Nerves
to the specialized jobs they perform. Most neurons have,
A nerve consists of a membrane sheath encasing bundles of axons.
in one form or another, the following four structures or
regions: (1) cell body, or soma; (2) dendrites; (3) axon; and
Nerve (4) terminal buttons.
Bundle of
axons SOMA The soma (cell body) contains the nucleus and
much of the machinery involved in the life processes of
the cell. (See Figure 2.4.) Soma shape can vary in different
kinds of neurons.

DENDRITES Dendron is the Greek word for tree, and the

Blood
vessel Individual
axons
Figure 2.3 Sensory, Motor, and Interneurons
These three types of neurons relay information between the central
and peripheral nervous systems. In this example, the person sees
the glass of water, and sensory nerves relay the sensory information
toward the central nervous system. The motor output from the
central nervous system allows the person to lift the glass to take a
drink.
branched dendrites of the neuron look a lot like trees. (See
Figure 2.4.) Neurons communicate with one another, and
dendrites receive these messages. Dendrites function much
like antennas to receive messages from other neurons. Just
like an antenna can receive a message over a distance (think
of an antenna that detects radio or Wi-Fi signals), dendrites
receive neural messages that are transmitted across the
synapse (also referred to as the synaptic cleft), a small space
between the terminal buttons (described later) of the send-
ing cell and a portion of the somatic or dendritic membrane
of the receiving cell. Communication at a synapse typically
proceeds in one direction: from the terminal button (on
the presynaptic cell, or sending cell) to the membrane of the
other cell (the postsynaptic cell, or receiving cell). There are
exceptions to this pattern, and as we will see in Chapter 4,
some synapses pass information in both directions.

AXON The axon is a long, thin tube. The outer surface

of the axon carries information from the cell body to the
Sensory neuron
terminal buttons. The axon functions much like an electri-
cal cord carrying an electrical message from an outlet to an
appliance. (See Figure 2.4.) However, the basic message the
axon carries is called an action potential, and it involves
both an electrical and a chemical component. This is an im-
portant concept and will be described in more detail later
in the chapter. An action potential is a brief electrical and
Interneuron
chemical event that starts at the end of the axon near the
cell body at a point called the axon hillock, and travels to the
ends of the terminal buttons. In any given axon an action
Motor neuron
potential is always exactly the same size and duration. If an
action potential reaches a point where the axon branches, it
splits but does not diminish in size. Each branch receives a
full-strength action potential.
Like dendrites, axons and their branches come in dif-
ferent shapes. Axons can be extremely long relative to their
diameter and the size of the soma. For example, the longest
axon in a human stretches from the foot to the base of the
brain. Because some distant parts of the neuron may need
Neurons items that can be produced only in the soma, there must be a
system that can transport these items rapidly and efficiently
LO 2.2 Distinguish among the structures of a neuron.
inside the axon (like a subway system). This process is axo-
The neuron is the basic information-processing and plasmic transport. Action potentials occur along the surface
information-transmitting unit of the nervous system. of the axon, but axoplasmic transport moves substances

M02_CARL0287_13_GE_C02.indd 39 08/11/2021 14:18

 40 Chapter 2

Figure 2.4 Parts of a Neuron

Dendrites

Terminal
buttons

Soma
(cell body) Myelin sheath

Axon (inside
myelin sheath) Direction of
messages

along “tracks” that run inside the length of the axon. (See the item being transported down the axon. The kinesin mol-
Figure 2.5.) This form of transport is accomplished by mol- ecule then walks down a microtubule, carrying the cargo to
ecules of a protein called kinesin. In the cell body, kinesin its destination (Yildiz et al., 2004). Energy for this process
molecules, which resemble a pair of legs and feet, attach to is supplied by adenosine triphosphate (ATP) molecules

Figure 2.5 Axoplasmic Transport

This figure shows how kinesin molecules transport cargo along the cytoskeleton from the soma to the
terminal button. Another protein, dynein, carries cargo from the terminal buttons to the soma.

Terminal
buttons

Soma
(cell body) Axon (inside
myelin sheath)

Soma

Vesicle
Axon Dynein

Terminal
Axon button

Microtubules
Kinesin

Microtubules
Microtubules
Vesicle

M02_CARL0287_13_GE_C02.indd 40 08/11/2021 14:18


produced by the mitochondria. Another protein, dynein, car-
ries substances from the terminal buttons to the soma, a pro-
cess known as retrograde axoplasmic transport. Anterograde
axoplasmic transport is remarkably fast, moving contents at
up to 500 millimeters (mm) per day. Retrograde axoplasmic
transport is about half as fast as anterograde transport.
MYELIN SHEATH The axon is often covered by a myelin
sheath. Myelin is a fatty substance that provides insulation
for the electrical message carried along the axon membrane,
much like insulation on a wire isolates an electrical current.
(See Figures 2.4 and 2.5). Bundles of myelinated axons ap-
pear as white matter in the brain and in nerves. In the brain,
these bundles of myelinated axons are sometimes referred to
as tracts. When looking at brain tissue that has been removed
from the skull and sectioned, the bundles appear white be-
cause of the fat content in the myelin. Specialized neuroim-
aging techniques can be used to visualize the myelin-dense
white matter tracts in the brains of living organisms. These
techniques often add color overlays to the tracts to help re-
searchers better identify them in magnetic resonance imaging
(MRI) images (See Figure 2.6.)
Figure 2.6 Image of Tracts in the Brain
White matter fibers overlaid on an MRI scan and a female head. The
fibers transmit action potentials between brain regions and between
the brain and the spinal cord.
SCIENCE PHOTO LIBRARY/Agefotostock
M02_CARL0287_13_GE_C02.indd 41
Structure and Functions of Cells of the Nervous System 41
TERMINAL BUTTONS Most axons divide and branch
many times. At the ends of the branches are little knobs
called terminal buttons. Terminal buttons have a very im-
portant function: When an action potential traveling down
the axon reaches them, they secrete a chemical called a neu-
rotransmitter. This neurotransmitter (there are many differ-
ent ones) either excites or inhibits the receiving cell and helps
determine whether an action potential occurs in the receiv-
ing cell’s axon. The release of neurotransmitters from the
terminal buttons is similar to being asked to open an oven
while something delicious is cooking. The axon sends a mes-
sage to the terminal buttons (check the oven) and a chemi-
cal message is released to diffuse into the synapse, relaying
a message from the presynaptic cell to the postsynaptic cell
(the smell of the food from the open oven diffuses across a
kitchen, alerting another person of the meal to come). Details
of this process will be described later in this chapter.
An individual neuron receives information from the ter-
minal buttons of axons of other neurons—and the terminal
buttons of its axons form synapses with other neurons. A neu-
ron may receive information from dozens or even hundreds
of other neurons, each of which can form a large number of
synaptic connections with it. Figure 2.7 illustrates the nature
of these connections. As you can see, terminal buttons can
form synapses on the membrane of the dendrites or the soma
(or even another axon, as you’ll see at the end of this chapter).
OTHER CELL STRUCTURES Figure 2.8 illustrates the inter-
nal structure of a typical neuron. Let’s start from the outside
of the neuron and work our way in. Much like your skin, the
cell membrane defines the boundary of the neuron. It con-
sists of a double layer of lipid (fatlike) molecules. Embedded
in the membrane are a variety of protein molecules that have
special functions. Some proteins detect substances outside the
cell (such as hormones) and pass information about the pres-
ence of these substances to the interior of the cell. Other pro-
teins control access to the interior of the cell, allowing some
substances to enter but preventing others from entering. Still
other proteins act as transporters, using energy to transport
certain molecules into or out of the cell. Because the proteins
that are found in the membrane of the neuron are especially
important for transmitting information, their characteristics
will be discussed in more detail later in this chapter.
The interior of the neuron contains a framework of pro-
tein strands. Much like the bones of your skeletal system,
this framework, called the cytoskeleton, gives the neuron
its shape. The cytoskeleton is made of three kinds of protein
strands, linked to each other to form a cohesive mass. The
thickest of these strands, microtubules, are bundles of 13 pro-
tein filaments arranged around a hollow core. In addition to
providing structural support to the neuron, the microtubules
form the “tracks” for axoplasmic transport. (See Figure 2.5.)
Cytoplasm is a semiliquid, jellylike substance that
fills the space surrounded by the membrane, including the
08/11/2021 14:18
 42 Chapter 2

Figure 2.7 Overview of Structure and Synaptic Connections Between Neurons

The arrows represent the direction information is traveling.

Synapse on soma

Soma
Myelin
sheath

Synapse on Axon Terminal

dendrite button

soma. It contains small, specialized structures, just as the
human body contains specialized organs. The generic term
for these structures is organelle, or “little organ.” Some im-
portant organelles are described next.
Deep inside the cell is the nucleus. The nucleus is enclosed
by the nuclear membrane. The nucleus contains the chromo-
somes. Chromosomes consist of long strands of deoxyribo-
nucleic acid (DNA). The chromosomes have an important
function: They function like cookbooks and contain the reci-
pes for making proteins. Portions of the chromosomes, called
genes, contain the individual recipes for individual proteins.
Genes are responsible for initiating the process
of ­protein synthesis in the cell. An overview of protein
synthesis is described next, followed by additional
­
details in the subsequent paragraphs. When they are
­
active, the genes help produce another complex mol-
­
ecule, m­ essenger r­ ibonucleic acid (mRNA). The mRNA
copies the ­information stored by the gene. The mRNA
then leaves the nucleus with the copied information and
attaches to r­ibosomes in the soma. The ribosomes use
the copied information from the mRNA to synthesize
proteins for the call. (See Figure 2.9.)

Figure 2.8 Internal Structures of a Neuron

Dendritic
spines

Nucleus

Cytoplasm Dendrite

Membrane

Microtubules
Terminal
buttons

Myelin
Mitochondria
sheath

M02_CARL0287_13_GE_C02.indd 42 08/11/2021 14:18


Figure 2.9 Protein Synthesis
When a gene is active, a copy of the information is made onto a molecule of messenger RNA (transcription). The mRNA leaves the nucleus and
attaches to a ribosome, where the protein is produced (translation).
Nucleus
Transcription
Neuron
Newly synthesized
protein
Protein synthesis a two-step process. In the first
step of the process, transcription, information from DNA
(which cannot leave the nucleus) is transcribed into a
portable form: mRNA. mRNA takes this information to
the ribosomes for the second step of the process: transla-
tion. During translation, the ribosomes use the informa-
tion from the mRNA to create proteins from sequences of
amino acids.
To help you remember the process of protein produc-
tion, compare it to making a cake from a top-secret recipe.
Imagine that the recipe for the cake is found in a rare cook-
book in a library, and you cannot remove the cookbook
from the library. You can go into the library and take a
picture of the recipe with the camera on your cell phone.
Now you have the information in a new, more portable
form. Next, you bring the picture of the recipe home with
you to your kitchen. There, you use the recipe informa-
tion to combine raw ingredients like flour, eggs, and milk
into the cake batter that you will bake. In this example, the
cookbook locked in the library is like the DNA stored in
the nucleus. The process of photographing the cookbook
and removing the recipe information from the library rep-
resents transcription of information from DNA in the nu-
cleus to a new, more portable form of information, mRNA.
Taking the photo home and using the information it con-
tains to assemble raw materials into a final product rep-
resents translation as the mRNA leaves the nucleus and
M02_CARL0287_13_GE_C02.indd 43
Structure and Functions of Cells of the Nervous System 43
Detail of Nucleus
DNA
Gene mRNA
Membrane of nucleus
Translation
mRNA
Ribosome
takes information to the ribosomes, which the ribosomes
then use to create proteins.
Proteins are important to cell functions. In addition to
providing structure, proteins serve as enzymes, which di-
rect the chemical processes of a cell by controlling chemical
reactions. Enzymes are the cell’s construction and demo-
lition crews: They join particular molecules together or
split them apart. In this way, enzymes determine what
gets made from the raw materials in the cell and determine
which molecules remain intact.
Found in the cytoplasm, mitochondria use nutri-
ents such as glucose to provide the cell with energy to
perform its functions. Mitochondria produce a chemical
called ­adenosine triphosphate (ATP), which can be used
throughout the cell as an energy source. Mitochondria
perform a vital role in the economy of the cell. Many of
the biochemical steps involved in extracting energy by
breaking down nutrients take place within mitochondria,
and are controlled by enzymes located there. Most cell bi-
ologists believe that many eons ago, mitochondria were
free-living organisms that came to “infect” larger cells.
Because the mitochondria could extract energy more ef-
ficiently than the cells they infected, the mitochondria
became useful to the cells and eventually became a per-
manent part of them. Because of their role in generat-
ing usable energy, mitochondria can be considered the
“power plants” of neurons.
08/11/2021 14:18
 44 Chapter 2
Supporting Cells
LO 2.3 Compare supporting cells in the central and
peripheral nervous systems.
Neurons constitute only about half the volume of the CNS.
The rest of the CNS contains of a variety of supporting
cells. Neurons have a very high rate of metabolism but
have no means of storing nutrients, and they must con-
stantly be supplied with nutrients and oxygen or they will
quickly die. Because of this, the cells that support and pro-
tect neurons are critical to our existence.
SUPPORTING CELLS OF THE CENTRAL NERVOUS
­SYSTEM The most important supporting cells of the CNS
are the neuroglia, or “nerve glue.” Glia (also called glial cells)
do much more than just hold the nervous system together.
Glial cells have a wide variety of important functions in the
nervous system. Neurons lead a very sheltered existence.
They are buffered physically and chemically from the rest of
the body by the glial cells. Glial cells surround neurons and
hold them in place, regulating their supply of nutrients and
some of the chemicals they need to exchange messages with
other neurons. Glial cells also insulate neurons from one an-
other so that neural messages do not get scrambled; destroy
and remove pathogens or dead neurons; are involved in
growth, repair, and development of the nervous system; and
can be involved in synaptic communication.
Here we will focus on introducing three important
types of glial cells: astrocytes, oligodendrocytes, and
Figure 2.10 Structure and Location of Astrocytes
The processes of astrocytes surround capillaries and neurons of the central nervous system. Astrocytes
regulate chemicals in the synapses (upper panel) and the chemical composition of the fluid surrounding
neurons (lower panel).
Astrocyte
Neuron
M02_CARL0287_13_GE_C02.indd 44
microglia. Additional types of glia have been recently dis-
covered but the full extent of their function in the nervous
system is not yet known (Dimou and Götz, 2014).
Astrocytes Astrocyte means “star cell,” a name that refers
to the shape of these cells. Astrocytes provide physical sup-
port to neurons and clean up debris within the brain. (See
Figures 2.10 and 2.13.) They produce some of the chemi-
cals that neurons need to fulfill their functions. They help
to control the chemical composition of the fluid surround-
ing neurons by actively taking up or releasing substances
whose concentrations must be kept within critical levels. The
somatic and dendritic membranes of neurons are largely sur-
rounded by astrocytes, and astrocytes are involved in provid-
ing nourishment to neurons. In addition, astrocytes function
as “neuron glue” and help hold neurons in place. These cells
also surround and isolate synapses, limiting the dispersion of
neurotransmitters that are released by the terminal b ­ uttons.
When cells in the central nervous system die, certain
kinds of astrocytes clear away the debris. These cells are
able to travel around the CNS. When these astrocytes con-
tact a piece of debris from a dead neuron, they engulf and
digest it. This process is called phagocytosis. If there is a
lot of injured tissue to be cleaned up, astrocytes will divide
and produce enough new cells to do the job. Once the dead
tissue has been broken down, a framework of astrocytes
will be left to fill in the vacant area, and a specialized kind
of astrocyte will form scar tissue, walling off the area.
Energy
Blood
vessel Lactate
Lactate
Glucose
Glucose
Glycogen
Lactate (storage)
08/11/2021 14:18
Another random document with
no related content on Scribd:
XII.

TAIDE- JA AVIOELÄMÄÄ.

Se valtava suosio, joka tuli Ida Aalbergin Kööpenhaminasta käsin

tekemien vierailunäytäntöjen osaksi Suomessa sekä Helsingissä
että, kesällä 1886, maaseudulla, antoi näyttelijättärelle kokemuksen,
jonka voisi kuvata vanhalla sananlaskulla: »Oma maa mansikka muu
maa mustikka.»

Noiden vierailujen aikana helsinkiläinen arvostelu taaskin totesi,

että näyttelijättären taide oli käynyt entistä hillitymmäksi ja että
entiset suuret luomat yhäkin olivat kehittyneet. Helmikuussa 1887
hän Camillena Musset'n »Ei lempi leikin vuoksi» näytelmässä antoi
todistuksen lyyrillisen lausuntansa sulosta ja tehosta.

Kesällä 1887 Ida Aalberg Viipurin maanviljelysnäyttelyjen aikana

esiintyi vanhojen loisto-osiensa ohella uudessa tehtävässä, Rebekka
Westinä Ibsenin »Rosmersholmissa.» Se ei koitunut menestykseksi,
sillä Ida Aalberg suhtautui kielteisesti koko osaan. Sulhaselleen hän
Viipurista kirjoitti:
 »Me harjoitamme nykyään Rosmersholmia, minä olen niin
onneton täytyessäni näytellä Rebekkaa, en häntä ollenkaan käsitä,
koko kappale on niin pessimistillinen, että se tulee koomilliseksi.

»Varsin suuri menestys oli Ida Aalbergilla »Orleansin neitsyessä»

syksyllä 1887. J'eanne D'Arc'in osan suoritus teki illasta yhden niitä
Arkadia-teatterin kuuluisia »lämpimän tunnelmin iltoja», joita
suomalaisuudenharrastajat ovat muistelleet ylpeydellä ja ilolla.

Mutta muut uudet tehtävät näytäntökautena 1887— 1888 eivät

tuottaneet Ida Aalbergille vähintäkään tyydytystä. Svava Björnsonin
»Hansikkaassa» ei saavuttanut mainittavaa menestystä, ei liioin
Hjördis Ibsenin »Helgelannin sankareissa.» Tähän aikaan ei Kaarlo
Bergbomilla tunnu olleen yritteliäisyyttä eikä voimaa viedä häntä
uusiin voittoihin. Gustaf von Numers ihaili Ida Aalbergin
näyttelemistä »Erik Puke» draamassa, mutta siinäkään menestys ei
ollut niin ehdoton kuin tavallisesti. Sanomalehtiarvostelu oli joko
kuivaa, kielteistä tai typerää.

Syksyllä 1888 Ida Aalberg käy uudestaan ulkomailla. Hän sai

sangen edullisen vierailutarjouksen Bergenin teatterista, jossa tähän
aikaan oli johtajana Henrik Jäger, mies, joka 1885 oli Kristianiassa
julistanut Ida Aalbergin ylistystä. Bergenin lehdet ovat paljon
kaunopuheisempia kuin kotoiset suomalaiset.

»Bergens Aftenblad»kirjoitti »Nukkekodin» johdosta m.m.:

»Sanottakoon kaikkein ensiksi: rva Aalbergin esitys Noorasta

»Nukkekodissa» oli suuri voitto. Esiinhuutoja seurasi joka näytöksen
jälkeen. Se olikin esitys, joka ansaitsee mitä suurinta huomiota. Se
osoitti rva Aalbergin lahjain monipuolisuutta ja etevyyttä, jolla hän
ryhtyy taiteelliseen tehtäväänsä ja väsymättä muodostaa kuvan piirre
piirteeltä —. Rva Aalberg ei alleviivaa, ei osota sormellansa. Kaikki
tulee aivan itsestänsä, on luonnollista, on suurta.

Voisi tuoda esiin piirteen toisensa perästä rva Aalbergin

esityksestä, joka on suuren taiteilijan työtä. — Lapsellisuus,
huolettomuus ensimmäisen näytöksen alussa esitettiin suurella
reippaudella. Mutta vaimon, joka on ollut kahdeksan vuotta
naimisissa, ei tarvitse kuitenkaan näyttää aivan niin nuorelta.
Nuoruus kyllä havaitaan, vaikka Noora näyttäisikin hiukan
vanhemmalta. — Hauska oli kaikissa tapauksissa makeiskohtaus,
joka esitettiin hyvin leikillisesti. Huudahdus »tuhat tulimmaista» tuli
sekä lausunnan että liikkeen vallattomuuden johdosta todelliseksi
loistokohdaksi. Kohtaus lasten kanssa oli, kuten tavallisesti,
hurmaava. — Ja sitten alkaa rva Aalberg tuoda esiin roolin vakavaa
puolta kohdattuaan Krogstadin. Kaikille vakaville mielenilmauksille
on hänellä sanat, katseet ja liikkeet. Varsinkin oli joulukuusen
koristaminen ja keskustelu Helmerin kanssa Krogstadista
unohtumaton. Nooran kuva joulukuuseen pusertuneena, kun hän
tuijottavin silmin näyttää mielessään kuvittelevan kuilua, johon on
vaipunut, on yksinään kappale taidetta. — Toisessa näytöksessä
näytteli rva Aalberg suurella innostuksella. Siinä oli harvinaista
ylevyyttä ja lyriikkaa.

Rva Aalberg pysyi kaikin puolin todellisuuden rajoissa.» — — —

»Bergensposten» kirjoitti »Erään naisen rangaistuksen» johdosta:

»Nimi — sillä tällä näyttelijällä on nimi — ei ollut yleisöllemme

tuntematon.

Mutta se, mikä hänessä on läpeensä suomalaista, joka tulee ilmi

jo hänen nimessään ja sen soinnussa, se pujahtaa alituisesti esille
hänen lausumistavassaan ikäänkuin sanoen: »Minä en ole
ruotsalainen, tanskalainen enkä norjalainen» — nämä jäljet
huomataan selvästi, vaikkapa taiteilija vuosikausia olisi puhunut
ruotsia Ruotsissa, tanskaa Tanskassa ja, kuten nyt, ruotsia Norjassa.

Heti, kun hän esiintyy ja puhuu, tuntuu ikäänkuin hän toisi

mukanaan jotain tuntematonta, jotain vierasta.

Muuten sen, mikä tässä on vierasta ja tuntematonta, ei tarvitse

pelätä olevansa koditonta.

Valtiossa saa kansalaisoikeudet laillisella tavalla. Niin ja niin monta

vuotta maassa, sanotaan.

Taiteen laki murtaa muodollisuudet valon ja salaman nopeudella.

Ja monta minuuttia ei kestänyt, ennenkuin Ida Aalberg-Kivekäs sai

kansalaisoikeudet Bergenin näyttämöllä. —

Yhtä uljas kuin miellyttävä ulkonainen olemus, jalo ja hieno

plastiikka — toisinaan ehkä liiaksi à la tragédienne — ovat ne hyvät
ominaisuudet, jotka raivaavat hänelle tietä. Mutta hänen kaikkein
parhaat enkelinsä eivät ole silmin nähtävissä. Voi ainoastaan
aavistaa niiden olemassaolon, kuulla niiden siipien suhinaa läpi
voimakkaiden aistimusten, läpi sydänsävelten vuon, läpi sen
intohimojen yhtenäisyyden, joka astuu eteemme moisesta
näyttelemisestä, näytteleminen on liian heikko sana, siis: tuosta
elävästä elämästä näyttämöllä.

Sellainen Mathilde vaikuttaa »Rangaistuksen» toisessa

näytöksessä niin, ettei sitä voi koskaan unhoittaa. Puhutaan
intohimojen musiikista, mutta tämä naisellisuuden intohimoisuus
kaivautuu syvemmälle sydämeen kuin sävelet.

Kaikki, mikä on rikkiruhjottua, juurineen ylös kitkettyä, avutonta,

tuskallista, parantumatonta aina itseuhrautuvaisuuden epätoivoon
asti, kaikki esitettiin niin kauhean todellisena kuin olisi se ollut itse
elämää.»

Marsaa »Ruhtinas Zilahissa» norjalainen arvostelu kiittelee

parhaansa mukaan, sanoen, että esitys merkitsee Ida Aalbergin
kohoamista aikakautensa parhaiden näyttelijäin tasalle. Ofelia-
suoritus saa hyvän arvioinnin ja uudesta osasta, Agneksesta Ibsenin
»Brandissa» »Bergens Tidende» kirjoitti:

»Rouva Aalberg oli kaunis Agneksena. Hän oli kaunis, näimmepä

hänet joko poloisena äitinä, joka kaipauksen ja tuskan vallassa
tuijottaa hautausmaalle, missä hänen pieni aarteensa makaa lumen
alla, tai hirvittävän, voimakastahtoisen lainorjan alistuvana vaimona,
joka hylkää kaiken seuratakseen miestään tämän raskaalla tiellä
Jumalaa kohti, tiellä, joka päättyy Jehovan näkemiseen ja
kuolemaan; aina oli rva Aalbergin näytteleminen yhtä suurta. ‒ ‒ ‒
Tämän naisen uhrautuva rakkaus ikäänkuin levitti sovittavaa valoaan
yksin onnettoman fanaatikonkin ylitse. —

»Bergens Aftenblad» totesi »Brandin» johdosta, että

näyttämöllinen suoritus tuskin on eduksi kappaleelle, joka tällöin
vaikuttaa liian julmalta ja liian voimakkaalta.

Myöskin Camillesta »Ei lempi leikin vuoksi» kappaleessa Ida

Aalberg
Bergenissä sai tunnustuksen sanoja.
 Kirjeenvaihto Bergen-vierailujen ajalta todistaa, että Lauri Kivekäs
ja Ida Aalberg, vaikka olivat olleet vuoden naimisissa, yhä elivät
kuherruskuukausitunnelmassa.

Ida Aalberg kirjoittaa miehelleen Bergenistä m.m.: Ida Aalberg —

18

»Eikös kulta ole iloinen, että Misse on ensimmäinen, joka sanoo

sinulle hyvää huomenta, suutelee sinua, suutelee sinua oikein
raivokkaasti ja pyytää, että Kisse on iloinen ja heittää kaikki
hämärät ajatukset pois. Tee se, kulta, ei sinulla ole oikeutta
sanoihisi »elä kokonaan unohda minua!» Minä ajattelen sinua joka
hetki, kulta, ja toivon, että sinulla olis hyvä ja asiasi kävisi hyvin.
Missen sielu on Kissen luona.» —

»Kultani!

Kaikki pitävät sinun kuvastasi eikähän se ole ihme, siinä niinkuin

sinussa itsessäsikin on niin paljon jaloutta niin paljon lujuutta. Kun
katselee sitä, tulee mieleni niin haikeaksi ja surulliseksi sillä minä
tahtoisin koskea sitä, puhua sille, saada sitä puhumaan, kuulla
taasen sinun rakasta ääntäsi ja sydämmesi lämmintä tykytystä.
Mutta nyt sinulla on niin monta ympärilläsi, jotka sinusta pitävät ja
sinua ihmettelevät ja nielevät joka sanan huuliltasi, että minä
luultavasti joudun unohduksiin, et suinkaan sinä minua enää
kaipaakaan. Mutta jos niin on, jos se ei ole sen enempää, sen
syvempää, sitten minä en enään koskaan tahdo tulla kotiin, en
koskaan näyttää kasvojani, sillä minä en voisi sitä kärsiä ja kantaa.
Minä itse en tasaa tunteitani enkä myöskään kärsi sitä sinussa.
Minä tahdon yksin omata sinun elikkä ei ollenkaan. Oh! Tällä
hetkellä minä olen mustasukkainen seinille, joidenka sisällä sinä
oleskelet. Kyllä minä rakastan sinua hulluuteen asti. Olen kovin
väsynyt Menen levolle suudellen tuhansia kertoja sinun kuvaasi.»
—

»Sinä armas. Kun tänään tulin teatteriin näin pöydälläni sinun

kirjeesi. Herra Jumala minä tulin niin iloiseksi, etten uskaltanut
ottaa sitä käteeni. En uskaltanut lukea sitä, sillä silloin en olisi
uskaltanut näytellä Matilden ääretöntä sydämenhaikeutta. Koko
illan ollessani näyttämöllä ajattelin vaan tuota suljettua kirjettä ja
sen sisältöä, katsos minä olin kuin hurja, sillä minulla on ollut niin
ikävä, niin hirveästi ikävä. Kuinka minä iloitsin nähdessäni että
sinulla on ollut niin paljon työtä ja ettäs löydät työssä lohdutusta ja
tyyneyttä. Työ on suurin siunaus sen olen minä usein, usein
yksinään ollessani huomannut ja tuntenut. Ole, kultani, vaan
varovainen rahojen kanssa!» —

Jo 1888 Lauri Kivekäs on onneton vaimonsa viipymisestä

vieraassa maassa. Ida Aalberg kirjoittaa hänelle:

»Kultani, toivoni, iloni!

Coriolanus'en äiti Volumnia sanoo heikolle miniälleen Virginialle:

»Jos poikani olisi mieheni, niin iloitsisin enemmän poissaolosta,
jolloin hän saavuttaisi kunniaa, kuin hänen syleilyistään, niin helliä
todistuksia hänen rakkaudestaan kuin ne antaisivatkin. Hän ei olisi
paljoa parempi taulua, joka riippuu seinällä, ellei kunnianhimo
antaisi hänelle elämää. Jos minulla olisi tusina poikia niin tahtoisin
kernaammin, että yksitoista niistä kuolisi kunnialla isänmaan
puolesta kuin että yksi tuhlaisi elämänsä hekumassa ja
laiskuudessa.» Noita sanoja olen aina ihaillut, ne olen vereeni
imenyt. Niin pitäisi joka vaimon ajatella miehestään, joka miehen
ajatella vaimostaan, silloin elämä tulisi rikkaaksi, ihmiset
onnellisemmiksi ja näköalat laveammiksi. Niin tahdon minä aina
ajatella ja niin Kissekin ajattelee.» ‒ ‒ ‒

Toisessa kirjeessä:

»Elä nyt Lauri kultani ole ikävissäsi, sinulla on parempaa

tehtävää kun antaa ikävän saada valtaa itsesi yli. Me teemme
molemmat työtä toistemme hyväksi ja kun sitten taas tulemme
toistemme luo — voi! voi!!! sitä riemua.» —

Näihin aikoihin Lauri Kivekäs yhdessä nuoremman veljensä

kanssa osti Helsingistä Kaivopuistosta erään palaneen talon rauniot
ja ilmoitti vaimolleen, että lähiaikoina saataisiin oma koti omaan
rakennukseen. Tämä tiedonanto sai Ida Aalbergin suuren riemun
valtaan. Hän kirjoitti miehelleen:

»Sinä oma kultani! Voi kuinka liikuttavaa, että sinä noin

lämpimästi ajattelet meidän yhteistä tulevaisuutta, julmasti
rasitettuna työssäsi kuitenkin annat itsellesi aikaa toimia ja sääliä
niin energillisesti, että todellakin pääset niin loistavaan resultaattiin.
Kuinka olin iloinen, kuinka onnellinen kun luin kirjettäsi, tuhatta
vertaa onnellisempi, ettäs teit tuon kaikki minun poissa ollessani, se
todistaa sinun syvää rakkauttasi, sinun lämmintä huolenpitoasi.
Hyvät henget ovat liehuneet ympärilläsi ja suositelleet sinua
kaupassa, joka minusta nähden on jotakin niin aavistamatonta, että
tuskin voin uskoa sitä. Lauri kulta, ajatteles mikä onni! 8000
tuhannella markalla! mikä hinta se nyt on, kun sillä saamme aivan
oman kodin ja pienen, pienen ryytimaan, eikö totta? Voi armaani
kuinka olen iloinen, tahtoisin lentää luoksesi, lentää syliisi ja
suudella sinua ja sanoa kuinka olen riemuissani. Minä muistan
paikan aivan hyvin. — Eiköhän ole tarpeen, että ryhdytään
piirustuspuuhiin nyt heti, muista, että menee pitkä aika ennenkuin
ne tulee valmiiksi ja jos aletaan korjaamaan kiviseiniä niin ei ole
aikaa odottaa. Me itse otamme asuntomme ylikerrassa
puukerroksessa. Huoneet siellä pitää tulla korkeiksi. Jos en muista
väärin on huoneet pienet, mutta emmekö voi ylhäällä ottaa jonkun
seinän pois ja sen kautta saada esimerkiksi vähän suurempi
ruokasali, se on kaunista kun ruokasali on suuri. Mitä salonkiin
tulee niin se luullakseni onkin koko suuri, jos ei niin voimme jättää
väliseinään, rinnalla olevaan pieneen huoneeseen, suuren
kaarevan aukon, sillä lailla täällä paljon käytetään, niin se on kuin
yksi suuri ja toki kaksi eri huonetta. ‒ ‒ ‒ Ole praktillinen, elä
maksa enemmän kuin täytyy. Jos tahdot halpoja muuraria niin
kirjoita isälle, hän mielellään ottaa ne toimittaakseen. Kirjoitan tästä
kaikesta sillä arvaan, että joka päivä maksaa. Tulenhan, kultani,
kohta itse, mutta sinun täytyy varmaan jo sitä ennen ryhtyä toimiin.
Kuinka olen iloinen! Herranen aika kuinka Kisseni on kiltti! ‒ ‒ ‒

Toissa päivänä kirjoitin yhden pienen kirjeen, jossa olen kovasti

pahoillani. Asia oli se, että minulla oli niin kovasti ikävä, aamusta
iltaan ja illasta aamuun en tehnyt muuta kuin ikävöin kirjettä sinulta,
ja kun sitä ei kuulunut tulin kovin onnettomaksi, ja ajattelin, että
sinä viihdyt liian hyvin toisten seurassa ja sitten tulin niin
mustasukkaiseksi tuota ajatellessani, että en tietänyt mitä sanoin,
olin aivan mieltä vailla. Mutta elä ole suuttunut Misseen, se tapahtui
rakkaudesta sinuun. Kulta, kulta, kuinka olen riemuissani, että
kohta pääsen pois luoksesi. — Tiedätkö, täällä on eräs konsuli
Gran, rikas, mahtava perhe ja heidän poikansa tahtoo tulla
näyttelijäksi, suomalaiseksi näyttelijäksi. Vanhemmat ovat
epätoivossa. Hänen äitinsä kävi luonani tänään, kaunis, intelligentti
nainen, minä annoin hänelle Bergbomin adressin ja paljastin
meidän surkeat teatteriolot.» ‒ ‒ ‒
 Näinä vuosina Kaarlo Bergbom tuntuu teatterinjohtajana eläneen
jonkinlaista suuren heikkouden aikaa. Näyttelijät kapinoivat häntä
vastaan ja nerokas iva, jolla hän suhtautuu näiden hommiin, ei
muuta sitä tosiasiaa, että suomalaiselta teatterilta hetkellisesti
puuttui toiminnan ryhtiä ja tarmoa. Norjasta palattuaan Ida Aalberg
heti on suuren tyytymättömyyden vallassa. Hän esiintyy ensin
Suomalaisen teatterin vierailunäytännöissä Turussa, mistä kirjoittaa
miehelleen m.m.:

»En pidä ollenkaan Bergbomeista, kadun niin että tulin. He ovat

kylmiä ihmisiä. He antavat arvoa ihmiselle vain silloin, jos tämä
pystyy auttamaan pyrkimystä tehdä tohtorin nimi
kuolemattomaksi.» (Viimeinen lause on ruotsinkielinen).

Näyttelijätovereihin hän pian purkaa halveksumistaan ja vihaansa.

Jo Norjasta saavuttuaan Ida Aalberg ihmettelee naisnäyttelijäin
puolelta tulevaa kateellisuutta, ja keväällä 1889 hän miehelleen
kirjoittamassaan kirjeessä nimittää koko seuruetta eläinkokoelmaksi.
Näytäntökausi 1888—1889 voi yhtä vähän kuin edellinenkään
tyydyttää häntä. Tosin hän silloin saavutti taiteilijauransa suurimman
voiton ingénue-alalla, Cypriennenä »Erotaan pois» komediassa,
mutta muut uudet tehtävät, Desdemona »Othellossa» ja Ellida
»Meren tyttäressä» kulkivat vähin äänin ohitse.

»Erotaan pois»[22] kappaleen esityksestä »Valvoja» m.m. sanoo

Ida
Aalbergin osalle:

»Vaikea on sanoin kuvata naiivista totisuutta, vallatonta iloisuutta

ja todellista tunteellisuutta, sekä näiden pääsäveleiden välillä
väikkyviä puolisäveleitä, joita näyttelijätär toi esiin vaihtelevan
osansa eri kohdissa. Vilkkauteen nähden oli hän kyllä ranskalainen,
mutta itse luonteen tulkitsemisessa oli, niinkuin muutoin jo toinen
kielikin tekee luonnolliseksi, pohjoismainen väritys.»

»Finland» toteaa, että »Erotaan pois» saavutti myrskyisen

menestyksen.
Sitten lehti jatkaa:

»Rouva Ida Aalberg Cypriennenä näyttäytyi aivan uudessa

valossa. Tosin rouva Aalbergilla ennenkin on ollut samantapaisia
osia, mutta niin täydellisen hienona ja nuorekkaan naivina, niin
mestarillisen yksinkertaisena tuskin milloinkaan olemme häntä
nähneet. Tässä ei ollut edellisten osien erinomaisia detaljeja, vaan
tästä muodostui uusi ja erikoisesti muovailtu tyyppi ‒ ‒ kaikki tuo
kokemattomuus, joka avioerosta tahtoo löytää uneksittujen
ihanteiden toteutumisen, tuo naivi tyytyväisyys, jolla hän kuvaa
pieniä juoniaan miehen selän takana, ja lopuksi tuo tiedoton rakkaus,
joka yhdistää hänet tuohon samaan mieheen, tulkittiin niin raikkain,
luonnollisin piirtein, kuin vain todellinen taiteilija voi löytää.»

Keväällä 1889 Ida Aalberg suuren, mutta verraten epämääräisen,

tyytymättömyyden vallassa erosi Suomalaisen teatterin vakinaisesta
palveluksesta. Kului pari vuosikymmentä, ennenkuin hän siihen
virallisesti uudelleen liittyi. Tämä »erotaan pois» merkitsi hänelle
uutta eurooppalaistumisyritystä.

*****

Kesällä 1889 Ida Aalberg ja Lauri Kivekäs oleskelivat Ranskassa,

Aix les Bains’issä, missä hoitivat terveyttään. Tällä matkalla Ida
Aalberg ensimmäistä kertaa elämässään pääsi myöskin Italiaan.
Miehensä jo matkustettua Suomeen hän yhä jäi Alppien
eteläpuolelle ja aviopuolisoiden toisilleen lähettämistä kirjeistä
näkee, että heidän välillään vallitsi suuri hellyys. Ida Aalberg kirjoitti
päivittäin pitkiä kirjeitä, joissa hän puhuu kaikesta, mitä hänelle
tapahtuu, kertoo suuresta ikävästään ja kaipauksestaan. Hänen
Italianmatkansa päätarkoituksena oli tutustuminen Eleonora Dusen
taiteeseen, mutta sattui niin, että Duse oli ulkomailla, eikä Ida
Aalberg saanutkaan tätä nähdä. Firenzessä ja Torinossa hän mitä
suurimmalla mielenkiinnolla seurasi erään vanhemman italialaisen
näyttelijättären, Marinin, esityksiä ja ihaili tavattomasti tätä varsinkin
»Odettessa». Michelangelon taide ja elämänhistoria — Ida Aalbergin
mielilukemista olivat tähän aikaan suurten henkilöiden elämäkerrat
— tekivät häneen syvän vaikutuksen.

Ida Aalbergin persoonallinen vastenmielisyys Suomalaista

teatteria ja kotimaan oloja kohtaan saavat hänen kirjeissään sangen
selvän ilmauksen. Hän vastaa erittäin kopeasti Kaarlo Bergbomin
nöyrään pyyntöön, että hän tulisi vierailemaan Helsinkiin ja Pietariin.
Hänellä oli toisia suunnitelmia, joista hän ilmoitti vain miehelleen.
Hän tahtoi taaskin päästä väljemmille vesille, hän tahtoi päästä
näyttämölle Saksassa. Palatessaan syksyn lopulla Italiasta
Suomeen hän poikkesi Berliniin. Hän saapui kaupunkiin ypöyksin,
eikä hänellä ollut siellä ketään tuttavaa. Onnellinen sattuma oli
johtanut samaan pensionaattiin, johon hän asettui asumaan, myös
erään venäläisen kenraali Weljaminovin rouvan. Tämä oli
aikaisemmin ihaillut Ida Aalbergia näyttelijänä ja oli hurmaantunut
saadessaan nyt persoonallisesti tutustua taiteilijaan. Rouva
Weljaminovin avustuksella sai Ida Aalberg pian paljon suosijoita,
jotka koettivat hankkia hänelle tilaisuuden esiintymiseen
berliniläisellä näyttämöllä. Se näytti aluksi varsin toivottomalta.
Saksan teatterioloissa oli ehtinyt tapahtua suuri muutos,
vierailunäytännöt olivat tulleet huonoon huutoon ja niitä vastustettiin
periaatteen vuoksi. Ainoan mahdollisuuden tuntui antavan vasta
perustettu parisilaismallinen yhdistys »Freie Bühne», jonka
yhdistyksen tarkoituksena oli tilapäisin voimin esittää uudenaikaista
ja radikaalia ohjelmistoa. Kysymys Ida Aalbergin esiintymisestä
lykättiin kuitenkin kevätkaudeksi, ja hän tuli viettämään jouluaan
kotimaahan.

Joulun jälkeen Ida Aalberg vieraili lyhyen aikaa Suomalaisessa

teatterissa, ja tarkoitti tämä esiintyminen lähinnä valmistumista
Berlinin varalle. Hän näytteli nimiosaa Emile Zolan ultrarealistisessa
»Thérèse Raquin» dramatiseerauksessa, jonka luuli tulevan esille
Freie
Bühnessä.

Berliniin 1890 tehty matka muodostui Ida Aalbergille mitä

kovimmaksi koetukseksi. Hänen miehelleen kirjoittamissaan kirjeissä
esiintyy tahtoihminen Ida Aalberg erinomaisen elävänä. Hän
taistelee sankarillisesti vieraan kielen tuottamia vaikeuksia vastaan,
hän taistelee raivoisasti muita vaikeuksia vastaan.

Aluksi kirjeiden sävy on toivehikas. Kieliopinnoistaan Ida Aalberg

kertoo miehelleen seuraavan jutun:

»Sinä Missen ainoa kulta, tulen juuri opettajani luota, olen vielä
aivan ilon vallassa siitä mitä siellä sain kokea ja nähdä. Katsos asia
on se, kuten kerroin olen ollut hirmuisen ahkera, luopunut ihan
omasta itsestäni ja ollut vaan — koulutyttö. Lukenut a.b.c. aamusta
iltaan, vieläpä yölläkin, aivan kuivan kuivaa mekaanillista työtä jota
pränttäsin itseeni saadakseni kieleni saksalaiseksi. Tunnissani olen
ollut yhtä kuiva ja mekanillinen ynnä suljettu. Olin usein tuskissani
tuntui kun en pääsisi hiuskarvaa edemmäksi. Viime kerralla sanoi
vihdoin opettajani että nyt olemme niin pitkällä että voimme jotain
helppoa proosaa alkaa lukemaan, jotain »Kindergeschichte»
[lastentarinaa] minä vihdoin protesteerasin, sanoin että hän aivan
henkisesti murhaa minut ja pyysin että saisin itse jotain valita, —
johon hän vihdoin suostui. — Otin siis Romeo ja Julian. — Tulin
sinne sitten tänään kirjoineni ja ne nähtyään, hänen suunsa meni
hymyyn, kummalliseen hiljaiseen hymyyn ja muuhun puuttumatta
hän sanoi alkakaamme — ja me aloimme. — En tahdo nyt kertoa
hänen kummastustaan ja ihmetystään sen enempää sanon vaan
että preussilainen kunniansa on kovin arka, en luule että hän vielä
koskaan on oppilastansa tambuuriin saattanut, vielä vähemmän
auttanut kappaa hänen päälleen, — tänään hän sen teki, vieläpä
kumarsi ylpeän selkänsä ja haki galoschini, — ja siitä minä olen
ylpein! Työni menee eteenpäin ja minä tunnen itseni onnelliseksi,
ensi kertaa, olen iloinen, oikein noin sisällisesti, siitä saakka kun
kultani luota lähdin.» —

Eräässä toisessa kirjeessään hän kertoo elämästään ja olostaan

Berlinissä, missä oli asettunut asumaan v. Sanckenin pensionaattiin:

»Elä kulta murehdi terveyttäni, toivon, että se yhä ja yhä menee

eteenpäin, että piankin olen terve kokonaan. Koetan hoitaa itseäni
paraimman mukaan ja neiti v. Sancken tekee minkä voi
hemmoitellakseen minua. Hän on vanhaa preussiläistä aatelia,
harmaahapsinen hieno, hento nainen joka nyt tuhlaa minuun
kaiken tukahdetun lempensä. Minulle keitetään eri ruokaa usein,
annetaan parempaa olutta (»Maltzbier») kuin muille ihmisille y.m.
Kenraalskalta [rouva Weljaminov], joka ennen usein kävi tohtorinna
Gompertzin luona joka myös asuu täällä hän oli kuullut minusta, ja
se kai vaikutti sen että hän heti koroitti minut oikealle kädelleen
pöydässä. Naapurini toisella puolella on eräs tohtori Preuss,
hienosta perheestä lähtenyt hieno mies, Zolan ja Flaubertin ja
Ibsenin ihailija, »freisinnig» [vapaamielinen] politikissaan ja vapaa
ennakkoluuloista, hän on amanuensi suuressa kirjastossa. Neiti v.
Sancken minä ja hän olimme katsomassa Ibsenin »Hærmendena
paa Helgeland» minä olin huonolla tuulella. Hjördis näytteli paljon
paremmin kuin Misse sitä rollia. En kuitenkaan virkkanut mitään
muille. Neiti v. Sanckenin toisella puolella istuu Tohtori Weitnitz,
rikas ja riippumaton, kuluttaa aikansa matkoilla, suuri taiteen
ystävä, Jerusalemista kirjoittanut, kuulen ma, hyvin intresantin
kirjan. Hänellä kuuluu olevan komea koti mutta syö täällä päivällistä
seuran tähden. Hänen rinnallaan 3 neitiä Bauisch, puolalaista
sukua, meidän kaikkein kauhistus, he ovat oikein »todellisia naisia»
tietämättömiä ja haluttomia, elävät aivan työtöntä elämää, jotka
kaikista ponnistuksista ja rahoista huolimatta vielä 30, 33—40
vuotiaina ovat yksin elämässä. Heidän vieressään yksi Berlinin
paraista maalareista Lindemann. Minun naapurini vieressä eräs
professori Lontoosta sekä sitten joukko engl. ja amerikalaisia
nuoria neitoja jotka ovat täällä joko kieltä tai soittoa oppimassa.
Pöydän toisessa päässä toht. Gompertz rouvineen sekä eräs
baroni Fuchs, joka on diplomati. Ja siinä näet koko seuramme
koossa. Useimmiten kuitenkaan ei synny yleistä keskustelua, seura
on liian suuri, 24 henkeä, ja sitäpaitsi kovin erilaisia. Tulevaisuuden
ihmisiä on vaan tohtorinna Gompertz, (mies on politikissa
»liberaali» tiedät siis minmoinen muussa) mistress Dingee,
Weitnitz, Preuss ja Misse. Itsestään seuraa että me useimmin
puhumme toistemme kanssa kun muiden. — — Tänä iltana Misse
menee, jos jaksan, olen nim. maannut koko aamupäivän,
kuulemaan Schlenterin luentoa tulevaisuuden dramasta. Hän on
paras kritikeri täällä. En kuitenkaan vielä tunne häntä
persoonallisesti.» —

Vähitellen kirjeiden sävy alkaa kuitenkin käydä toiseksi.

Toukokuun alku 1890 on hyvin vaikeata aikaa Ida Aalbergille, kuten
nähdään esim. seuraavasta kirjeestä:

»En minä kulta ole sinua unohtanut, mutta minä olen ollut niin
suruissani, niin kovin suruissani, etten ole jaksanut kirjoittaa. Koko
elämä on tuntunut kuin yksi ainoa suuri tuska. Minun terveyteni on
varmaan iäksi päivin mennyt. Ja mitä on elämä ilman terveyttä ja
työtä. Minä en enää niinkuin muinoin kestää pitkiä ponnistuksia.
Vieraan kielen omaaminen ei ole helppoa ja harva sen on perille
vienyt. Minä kuitenkin mielelläni sitä tahtoisin (ja olenkin suuresti
edistynyt), sillä minä en tahdo kauvan enää näytellä, muutaman
vuoden vaan, ja kuitenkin tahtoisin silläaikaa koota jotain kokoon, ei
suurta summaa mutta kuitenkin muutaman kymmenen tuhatta ja
semmoista voi vaan suurissa maissa. Siis olen ollut kovin ahkera,
hyvin vähän käynyt ulkona valmistuakseni niin pian kuin
mahdollista, lukenut yöt ja päivät ja vähä vähällä aina maannut pari
päivää kipeänä. Minun kunnianhimoni ei sallinut että puhuisin
huonoa kieltä, sitä paitsi ajattelin, että minulla aina tulevaisuudessa
on siitä suuri hyöty ja tie avoinna suurille näyttämöille. Sitten kun
olin varma itseni kanssa ja ilmoitin itseni valmiiksi, sitten alkoivat
kaikki nuo intrigit, joita on ollut niin paljon ja usein naurettavia.
Kerron yhden niin saat aavistuksen kaikesta. Tohtori Blumenthal
joka oli kuullut minusta (hän on Lessing teaatterin direktööri) tulee
luokseni (hän oli jo ennen tuttu) ja kysyy jos en tahtoisi näytellä
Noraa. Minä tahdoin tietysti. Hän pyysi että vissinä päivänä kävisin
luonansa. Tein sen, B. oli erinomaisen ystävällinen — tulin myös
esitellyksi rouvalleen, ja kaikki oli miten voin arvata hyvin. Kun
sitten hänen vanha rouvansa (hän on vanhempi miestä) saa sen
päähänsä tulla — mustasukkaiseksi! Ajattele kuinka hassua! ja
mitä kaikkea heidän välillään lienee tapahtunut en tiedä, kaikessa
tapauksessa B. kovin geneerattuna tulee luokseni ja selittää että se
ei nyt voi tapahtua. Uh! sitä todellista naisellisuutta! Ja senlaista
harmia on paljon ollut, ja minua kovin rasittanut. Kun sitten vielä
vuoteella ollessani (koko pääsiäisenhän olin kipeä) otin vastaan
sen suuren rollin »Freie Bühneltä» en tuntenut rajaa työlleni. Päivät
eivät riittäneet, luin kaiket yöt ja päivällä harjoitin teatterissa monta
tuntia peräkanaa. Sillä se »Aufgabe» [tehtävä oli pääosa A.
Fietgerin draamassa »Jumalan armosta»] ei ollut helppo! —
Makasin pari päivää kipeänä ja aloitin taasen, minä tahdoin jaksaa
ja niin kokosin viimeiset voimani ja tein työtä samalla vauhdilla
kunnes hurjalla surulla aloin tuntea että en enään jaksanut 2 viime
näytöstä harjoittaa yhtä haavaa. Niin väsymys vähitellen minut
musersi kunnes prof. Krause absolutisti kielsi. Kun minä en
vieläkään totellut aikoi hän kirjoittaa sinulle ja pyytää sinua
kieltämään, — mutta sitä minä en sallinut. Hän sanoi että sillä
tavalla »reichten Sie sich zu Grunde, und ist keinen arzt mehr
möglich Ihnen wieder Lebenskräfte zu geben.» Minä pysähdyin,
lähetin rollin Schlentherille ja kerroin mitä lääkäri sanoi. Ja siihen
raukesi sitten se unelma. Oh! se on niin kurjaa. En ymmärrä kuinka
pääsisin entisiin voimiini jälleen. Ja kuitenkin minun täytyy, minun
täytyy, minä tahdon päästä eteenpäin, minun täytyy tulla terveeksi!
— Oh! kuinka minä kadun että olen niin paljon antanut Suom.
teaatterille! Mitä minulla nyt siitä on? Kehnoutta, kateutta ja ilkeyttä,
se on palkinto. Krause (sama joka myös viimeksi kun ei enään
mikään auttanut oli kutsuttu keisari Fredrik vainajan luo) hän on niin
viisas, niin hieno niin erinomainen lääkäri. Hän pitää paljon
Missestä ja on kovasti intreseerattu harrastuksistani täällä. Hoitaa
minua erinomaisesti. — Toki ei hän anna mitään lääkkeitä, muuta
kuin rautavettä jota juon 2 suurta putelia päivässä, sitäpaitsi juon
kannun maitoa päivässä, yks' suuri lasi joka tunti. 2 kert. päivässä
kylmä kylpy ja sitten päälliseksi käyn joka päivä luonaan, jolloin hän
elektriseeraa koko ruumiin ja kaulani, tiedätkö, sisäpuolelta sondien
kautta jotka suun kautta viedään sisään. Krause sanoo, ettei hän
vielä koskaan ole hoitanut naista, joka ylellisten rasitusten kautta
niin kokonaan olisi tärvellyt fysiikinsä kuin minä. Krause tahtoo, että
jättäisin näyttelemisen syksyyn. Ja kai sen teenkin niin kovin
alentavalta kun se tuntuukin, kun olin niin tyhmä, että edeltäkäsin
ilmaisin aikomukseni. Katson kuitenkin vielä vähän ympärilleni ja
sitten päätän. — Niin, siinä on nyt kaikki suuret unelmat yhtenä
rauniona! — En minä ole niin erinomainen kun sinä luulet, eikä
minun energiani niin suuri kuin luulet. Enhän edes tätä nyt ole
voinut läpi viedä. ‒ ‒ ‒

Oma Pirjo raiskasi.»

Edelleen hän tietää miehelleen kertoa:

»Ajatteles mitä vaikeuksia!!! Kerroin että minun piti ylös

harjoitukseen. Sen teinkin ja otin tuoksi 'profviksi' Romeo ja Julian.
Kaikki olivat ihastuksissaan ja huusivat »Sie müssen deutsch
lernen und deutsche schauspielerin werden» — Seuraavana
päivänä tuli rouva Kainz ja teatterin direktööri luokseni, hän oli
hyvin hämillään eikä tietänyt miten alkaa — arvasin heti että nyt oli
jotain tapahtunut — ja kertoi että hänen molemmat primadonnansa
eivät suvaitse että hän laskee minut esille sillä näyttämöllä, sillä
silloin ne rikkovat kontrahtinsa.» — — ‒

»Voi kuinka olen ollut onneton ja nöyryytetty, mutta voihan sitä

välistä kumartaa maahan saakka, sitä korkeammin voidakseen
hypätä kerta pystyyn. Kun näen tätä keskinkertaisuutta täällä
taiteen alalla, täällä ei löydy ainoatakaan intressanttia
naisnäyttelijää — ja tunnen mitä minä voisin luoda jos olisin
voimissani ja kerta saatuani kielen haltuuni, niin olen aivan
menehtyä.» —
 »Kulta, nyt olen juossut rappusia ylös ja alas 10 päivää, ollut
onneton, valvonut yöt ja tuntenut itseni aivan hyljätyksi ja
mitättömäksi mutta nyt olen päässyt niin pitkälle että kävin tänään
Kainzin luona ja näyttelin Julian hänen salongissaan muutamien
auktoriteettien edessä. — Ja kun olin lopettanut huusi Kainz »Sie
werden in Berlin spielen!» [Te näyttelette Berlinissä], vaikka se on
niin vaikeata päästä esille että Sonnenthal Wienin suuri näyttelijä
— ei pääse esille. Mutta Kainz sanoi »so jung und so gross, so was
haben wir nie gesehen!» [»Niin nuori ja niin suuri, senvertaista
emme ole milloinkaan nähneet»] hän löi käsiään yhteen ja sanoi
»und so spricht man von Goreva, das ist ja kein vergleich!» [Ja
sitten puhutaan Gorevasta, eihän heitä voi verratakaan.] — Goreva
Venäjän ensim. näyttelijätär joka 85 teki fiascon täällä ja jonka
vuoksi 3 teatteria on kieltänyt minun pyyntöni. — Kainz tahtoo että
näyttelen 3 roolia ja jään tänne 14 päiväksi. Jo ehkä huomena on
suuri harjoitus Residenz teatterissa.» —

Josef Kainz oli vähäistä aikaisemmin rikkonut välikirjansa erään

teatterin kanssa ja oli tähän aikaan hovi- ja kaupunkiteattereiden
pannaan julistama. Senvuoksi hän esiintyi näyttämöllä, jolla ei oltu
totuttu näkemään suurta draamaa. Kainz oli itse juutalainen, ja
Berlinin vaikutusvaltainen ja suurilukuinen juutalaisväestö osoitti
näissä Ostend-teatterin näytännöissä mielenosoituksellisesti
suosiotaan hänelle.

Ida Aalberg esiintyi Kainzin vastanäyttelijänä kahdessa

kappaleessa,
»Romeossa ja Juliassa» ja »Kavaluudessa ja rakkaudessa».

»Berliner Tageblatt» kirjoitti hänen Juliansa johdosta:

 »Oli vaikea tuntea Ostend-teatteria eilen. Missä vähän aikaa sitten
pyöveli D. Krantz heilutti kirvestään, missä helposti tyydytetty
porvariyleisö nautti ylen romanttisista hirmunäytelmistä, sinne oli
eilen kokoontunut kirjallisesti sivistynyt, hemmoteltu Berlin.
Semmoista ihmispaljoutta, joka eilen täytti Ostend-teatterin, saanee
tuskin pitkiin aikoihin nähdä. Ei ollut tyhjää paikkaa, ja orkesteri oli
otettu hätävaraksi. Ja näistä tiheistä joukoista, jotka jännityksellä
istuivat esiripun tällä puolen, nähtiin, ettei mikään tavallinen
tapahtuma ollut saanut heitä tähän kaukaiseen rakennukseen.
Salissa vallitsi omituinen mielentila, joka tavallisesti syntyy, kun
odotetaan jotain erinomaista, jokapäiväisyydestä eroavaa. Ja
todellakin mieltäkiinnittävä oli se koe, joka tapahtui tuolla ylhäällä
näyttämöllä. Suomalainen taiteilija, neiti Ida Aalberg, joka
kotimaassaan ja pohjoisissa naapurimaissa jo oli saavuttanut suuria
voittoja, aikoi astua ensimmäiset askeleensa voittaakseen
saksalaisena taiteilijana kunniaa ja laakereita. On aina uhkayritys
jättää varma taiteilija-asema ja tehdä hyppäys, josta vielä ei tiedä
viekö se ehkä pimeään syvyyteen, josta ei ole mitään paluuta. Siihen
vaaditaan rohkeutta, suurta rohkeutta, mutta sen lisäksi myöskin
suurta itseluottamusta. Ja se ei olekaan pettänyt nti Ida Aalbergia.
Hänen Juliansa, jonka hän eilen esitti Shakespearen »Romeossa ja
Juliassa», oli suurta kiitosta ansaitseva. Neiti Aalbergilla on hyvin
harvinainen lahja, nimittäin luonto, ja se merkitsee hyvin paljon jo
itsessäänkin, vaikkei siihen yhtyisikään, kuten tällä taiteilijalla,
vilpitön tunne ja todellinen intohimo. Hänellä on tavattoman vilkas
temperamentti ja tunteen syvyys, joka synnyttää säveleitä, mitkä
sydämestä tulevat ja sydämiin menevät. Parveke-kohtauksen hän
esitti kerrassaan hurmaavasti. Tässä hänellä oli apuna hienon
naisen sulava luontevuus. Luonnollinen intohimo, joka ei ollut
teatterimaista esiintymistä, teki valtavan vaikutuksen. Siinä ei ollut