Behavioral Neuroscience. An Introduction

839 Pages·1980·26.143 MB·English

by  Carl W. Cotman and James L McGaugh (Auth.)

With select illustrations created for this book by
Frank Armitage, Ojai, California
And contributions from
Jay Angevine, Jr., University of Arizona College of Medicine, Tucson
Elliott Blass, The Johns Hopkins University, Baltimore
David Cohen, University of Virginia School of Medicine, Charlottesville
Edward Evarts, National Institute of Mental Health, Bethesda
Michael Gabriel, University of Texas at Austin
William Greenough, University of Illinois, Champaign
John Harvey, University of Iowa, Iowa City
Bartley Hoebel, Princeton University, Princeton
Robert Jensen, University of California, Irvine
Leonard Kitzes, University of California, Irvine Medical Center, Irvine
Rodolfo Llinas, New York University Medical Center, New York
Horace Loh, University of California, San Francisco
Andras Pellionisz, New York University Medical Center, New York
Lewis Petrinovich, University of California, Riverside
Jon Sassin, University of California, Irvine Medical Center, Irvine
David Segal, University of California, San Diego
Kenneth Simansky, Cornell Medical School, White Plains, New York
Larry Squire, University of California, San Diego
Arnold Starr, University of California, Irvine Medical Center, Irvine
Maurice B. Sterman, Veterans Administration Hospital, Sepulveda, California
Michael Stryker, University of California, San Francisco
Timothy Teyler, Northeastern Ohio University College of Medicine, Rootstown
Richard Thompson, University of California, Irvine
Roderick van Buskirk, Stanford University, Stanford
E. Leong Way, University of California, San Francisco
Richard Whalen, State University of New York, Stony Brook
Pauline Yahr, University of California, Irvine
Cover illustration from Fig. 5-8D.
Behavioral
Neuroscience
An Introduction
Carl W. Cotman
James L. McGaugh
Department of Psychobiology
University of California
Irvine, California
ACADEMIC PRESS
A Subsidiary of Harcourt Brace Jovanovich, Publishers
New York London Toronto Sydney San Francisco
COPYRIGHT © 1980, BY ACADEMIC PRESS, INC.
ALL RIGHTS RESERVED.
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United Kingdom Edition published by
ACADEMIC PRESS, INC. (LONDON) LTD.
24/28 Oval Road, London NW1 7DX
Library of Congress Cataloging in Publication Data
Cotman, Carl W
Behavioral neuroscience.
Bibliography: p.
1. Neuropsychology. 2. Human behavior.
3. Nervous system. I. McGaugh, James L., joint
author. II. Title. [DNLM: 1. Behavior—Physio-
logy. 2. Nervous system—Physiology. WL102.3 C844b]
QP360.C67 152 79-50214
ISBN 0-12-191650-3
PRINTED IN THE UNITED STATES OF AMERICA
To Ann, Adrian, Danna, Daniel and Cheryl
and
Becky, Doug, Jan and Linda
Preface
Behavioral neuroscience is a new and complex discipline that employs the
tools and language of biology, chemistry, physiology, and psychology in the study
of behavior. Yet for all its diversity, it is among the most exciting of endeavors.
"Behavioral Neuroscience" aims to achieve an understanding of the biological
mechanisms that determine our behavior and that of the animal world in which
we live. This text provides a basic understanding of what is known about the
means by which neurons communicate and about the nervous system which
interprets, integrates, and transmits signals into meaningful and appropriate
behaviors.
This book originated, in part, from an undergraduate course called "Intro-
duction to Psychobiology" that has been taught at the University of California at
Irvine since 1965. At other colleges and universities the course might be called
"Introduction to Physiological Psychology," "Introduction to Biological Psychol-
ogy," "Introduction to Neurosciences," "Introduction to Neurobiology," etc.
This course is attended primarily by students who have some background in
biology and/or psychology and who often are majors in biology, physiology, or
psychology. Many of them plan to attend graduate school or medical school.
"Behavioral Neuroscience" is organized so that it starts with a broad overview
presented in Chapter 1. In Chapter 2 we describe the general operation and
organization of the nervous system and introduce some of the major types of
neurons in the context of their systems. The presentation then moves directly to
the cellular level and focuses on neurons, their basic characteristics (Chapter 3),
and how they communicate (Chapters 4 and 5). We explore the action at the
neuronal membrane and across the synapse. Synapses are plastic, modifiable. We
discuss these processes and the basic integrative properties of defined groups of
neurons (Chapter 6). Based on the principles and concepts derived we discuss
learning in model systems (Chapter 7) and follow this directly with a discussion of
complex learning and memory (Chapter 8). We define the problem, present the
xiii
data, and derive the central conclusions. We proceed with a description and
analysis of the systems which process sensory stimuli (Chapters 9 and 10) and
generate movement (Chapter 11). We describe the operation and upkeep of the
body in relation to behavior (Chapter 12) and the performance of certain be-
haviors necessary for the care of the body, for example, thirst and hunger in
Chapter 13 and sleep and activity rhythms in Chapter 14. What factors control
these behaviors? We describe the development of the neural circuitry and its
plasticity throughout life (Chapter 15) as it is modified in accordance with a
lifetime plan. In Chapter 16 we describe the development of behavior, the
ever-so-complex product of all that "wiring" during development. Behavior is
sometimes abnormal, brought on by mental disorders, drug abuse, and al-
coholism, and in Chapter 17 we analyze these from a behavioral and neuroscien-
tific perspective. The final chapter (18) focuses on man, on us, and asks the
question, "Why am I what I am?" Certain known aspects of man's nervous system
give some of the answers—but in reality these answers raise even more questions.
Overall then, in the course of this text, we explore behavior from the perspective
of basic neuroscience.
The coverage of behavioral neuroscience is fairly complete. Thus it is un-
likely that any quarter or semester course at an undergraduate level will cover the
entire book. Most faculty will wish to select from it, depending on their own
particular orientation. Those wishing a more biological approach to behavioral
neuroscience may prefer to concentrate on the earlier chapters in the text; those
wishing a more psychological approach will draw more heavily on the later
chapters, more or less along the outline suggested in the instructor's manual.
As an aid to the creation of this text, we have enlisted the help of many
experts who have assisted in the preparation of various segments of this book. The
contributions of these specialists have enabled us to produce a text which is
uniformly accurate and up to date. We are indeed grateful to all the following
experts who have given so generously of their time and effort to provide this
information: Dr. J. Angevine, Jr. (neuroanatomy, Chapter 2, and cortical lobes,
Chapter 18); Drs. J. Harvey and K. Simansky (neurotransmitter circuitry, Chapter
6); Drs. R. Llinas and A. Pellionisz (original figures on dendritic integration,
Chapter 6); Dr. R. Thompson (model systems and elementary learning, Chapter
7); Dr. M. Gabriel (neurophysiological correlates of learning, Chapter 8); Dr. L.
Squire (human learning and memory, Chapter 8); Dr. L. Kitzes (auditory recep-
tors, Chapter 9); Dr. A. Starr (clinical disorders of hearing, Chapter 9); Dr. M.
Stryker (visual system and plasticity, Chapter 10); Dr. E. Evarts (motor system,
Chapter 11); Drs. R. Whalen and J. Sassin (neuroendocrinology, Chapter 12);
Drs. T. Teyler, R. Jensen and D. Cohen (autonomie nervous system, Chapter
12); Dr. E. Blass (thirst, Chapter 13); Dr. B. Hoebel (contribution on hunger,
Chapter 13); Drs. M. Sterman and R. Jensen (sleep, Chapter 14); Dr. R. van
XIV PREFACE
Buskirk (biological rhythms, Chapter 14); Dr. W. Greenough (environmental
influences on brain structure, Chapter 15); Dr. P. Yahr (contribution on genetic
influences on behavior, Chapter 16); Dr. R. Whalen (hormones and behavior,
Chapter 16); Dr. L. Petrinovich (material on language development in birds and
man, Chapter 16); Dr. D. Segal (schizophrenia and affective disorders, Chapter
17); Dr. E. Way (drug abuse, Chapter 17); and Dr. H. Loh (endorphins and
enkephalins, Chapter 6 and 17).
We acknowledge the aid of these experts; however, we bear the responsibility
for the material as it appears in final form. We have made every effort to provide
essential introductory background for each section. When concepts are intro-
duced that depend on basic principles derived from chemistry, biology, or psy-
chology, we have provided the appropriate background. Our goal was to see that
despite varied backgrounds, students will find this text easy to read. We have also,
insofar as possible, written sections to be self-contained so that the instructor can
select only parts of the text and still achieve continuity. The list of key terms at the
end of each chapter will aid in reading and retaining information contained in the
chapter. These plus the outline at the beginning of each chapter and the sum-
mary at the end should serve as a helpful study guide.
We are also pleased to acknowledge especially Dr. Robert Jensen who aided
us extensively, particularly with Chapters 8, 10, 12, 13 and 14, and Dr. Jay
Angevine, Jr., who provided many helpful editorial and content suggestions on
numerous chapters. Dr. Jensen played a vital and stimulating role and we are
infinitely appreciative of his diligent help and good cheer. Dr. Angevine spent a
sabbatical in Dr. Cotman's laboratory; his inspiration and fellowship are very
much appreciated. We are also grateful to Dr. W. Greenough and Dr. S. Gerling
who read the entire text and provided many helpful critical comments. Finally,
we thank Dr. S. Erulkar, Dr. H. Killackey, Dr. J. Swett, Dr. P. Landfield, Dr. M.
Nieto, Dr. S. Hoff, Dr. E. Zaidel, J. Ryan, and D. Smith for their discussion,
assistance, and critical comments on various chapters.
We were very fortunate indeed to obtain the services of Frank Armitage for
the preparation of conceptual sketches and "story boards." Each drawing was
developed in close consultation with Dr. Cotman to assure conceptual accuracy.
Through this process, Mr. Armitage has been able to provide animated repre-
sentations of many key concepts. The line drawings were done by Maureen
Killackey except for those in Chapter 11 which were done by Christine Bondante.
We are also pleased to acknowledge Julene Mueller and Linda O'Shea for their
dedicated and invaluable secretarial, editorial, and managerial assistance and
Christine Gentry for assistance in photography and reference verification. We
also gratefully acknowledge the photographic contributions of M. Weisbrodt
(photos in Chapters 11 and 17). Dr. Cotman is grateful to his wife for her
exceeding patience and support and her helpful comments on various sections of
PREFACE XV
the text. Dr. McGaugh appreciates the encouragement, understanding and good
cheer of his wife. And, finally, thanks are due to the many at Academic Press who
have played important roles in the project.
This book is written for students. It is our hope that those reading it will enjoy
it and share some of the excitement and enthusiasm we and the contributors have
for behavioral neuroscience. We welcome any comments that the users of this
book may have. Please write to either of us at the Department of Psychobiology,
University of California, Irvine, California. We enjoyed writing the book and
hope that you find it both interesting reading and exciting learning.
Carl W. Cotman
James L. McGaugh
XVI PREFACE
1
What Does the Nervous
System Do?
I. Introduction
II. Functions of the Nervous System
A. The Nervous System Organizes and Directs Motor
Responses
B. The Nervous System Monitors Its Outside World
C. Learning and Remembering
D. Thinking and Personality
E. Much of the Nervous System Is Devoted to the Care
and Upkeep of the Body
F. Development and Maturation
G. Behavioral Abnormalities
III. What the Nervous System Does
References
I. Introduction
. . . We think of ourselves and others as engaged from moment to moment in doing
this or that. That is a convenience of speech. Each of us at any moment of the
waking day is a whole bundle of acts simultaneously proceeding. . . . In no case
does any other of all the doings of the moment disturb the one focal doing. We are
each therefore at any moment a pattern of active doing; a single pattern of pieces all
subordinate to one keypiece. No other part of the pattern is allowed to disturb the
keypiece of the pattern. Should it do so then the pattern changes and the disturbing
piece becomes usually the keypiece of a new pattern which supplants the previous.
The keypiece is the crown of the unified doing of the moment (Sherrington, 1946,
pp. 172-173).
1
Our responses are the products of the nervous system acting through the
body. We walk, talk, touch, look, listen, even laugh, in so many ways that our
total responses can only be summed up as behavior—behavior designed for the
moment. Our nervous system provides an immense repertoire of specific be-
havior, together with built-in contingency plans. It is at one and the same time
reporter, editor, producer, file clerk and delivery boy for our moment by moment
events. It collects critical newsworthy information, analyzes it, modifies it in
relation to our interests, keeps a record of all transactions and delivers a report to
our muscles and glands so we may act. In our behavior we see reliability, precision
and versatility—programs of living designed specifically for us. Yet our reactions
to many things are identical, universals of response shared so commonly among
all men and women that we are able to analyze the biological basis of behavior
from them.
In this text we shall concern ourselves with the biological bases of behavior,
the "hows," "whys," and "so whats." How does the nervous system administer that
miraculous performance called behavior? What is the role of the subcomponents
of behavior in relationship to its overall plan? These questions are challenging
ones, and at present we really do not have all the answers. But in behavioral
neuroscience we are attaining more and more exciting findings. The prospects for
the future are very bright indeed. Neurobiology is still a very young field, and
much is happening every day. In this text we shall take you into the nervous
system and from within try to show how it pilots our behaviors. However, prior to
embarking on this journey and seeking out the ways the nervous system controls
behavior, we would do well to put our mission into perspective. Just what does the
nervous system do?
II. Functions of the Nervous System
A. The Nervous System Organizes and Directs Motor
Responses
We can start at the end—ourselves moving. Much behavior is purposeful
movement, and the nervous system is the executor of this movement. As Lord
Adrian (1955), a revered neurophysiologist and one of the founding fathers of the
study of the brain, once wrote, "The chief function of the nervous system is to send
messages to the muscles which will make the body move effectively as a whole."
Such effective, unitary movement has extraordinarily diverse components. Em-
bodied in it are some of the simplest and some of the most complex behavioral
2 1. WHAT DOES THE NERVOUS SYSTEM DO?