The Neural Control of Sleep and Waking

213 Pages·2002·1.995 MB·English

by  Jerome Siegel (auth.)

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The Neural Control of
Sleep and Waking
Springer
New York
Berlin
Heidelberg
Hong Kong
London
Milan
Paris
Tokyo
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The Neural
Control of Sleep
and Waking
Jerome Siegel
With a Foreword by Jerome M. Siegel
With 34 Illustrations
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Jerome Siegel
Departments of Psychology and Biological Sciences
University of Delaware
Newark, DE 19716
USA
[email protected]
Library of Congress Cataloging-in-Publication Data
Siegel, Jerome H., 1932–
The neural control of sleep and waking / Jerome Siegel.
p. cm.
Includes bibliographical references and index.
ISBN 0-387-95536-4 (hardcover: alk. paper)
ISBN 0-387-95492-9 (softcover: alk. paper)
1. Sleep—Research—History. 2. Sleep–wake cycle—Research—History. I. Title.
QP425 .S585 2002
612.8'21—dc21 2002067019
ISBN 0-387-95536-4 (hardcover) Printed on acid-free paper.
ISBN 0-387-95492-9 (softcover)
© 2002 Springer-Verlag New York, Inc.
All rights reserved. This work may not be translated or copied in whole or in part without the
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Printed in the United States of America.
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Springer-Verlag New York Berlin Heidelberg
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I dedicate this book to my wife, Priscilla Wishnick Siegel.
To her I owe a special debt of gratitude for her enormous
patience and encouragement over the four years it took
to write the book. I dedicate the book also to our three
children, Peter, Bennett, and Becky, who periodically
remind me that they spent considerable time during their
formative years visiting my lab and observing cat
surgeries, brain recordings, brain histology, and other
aspects of my early research on sleep and waking. I hope
this book will put those experiences into a larger and
meaningful perspective.
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Foreword
My first contact with “the other” Jerome Siegel came in 1973, when I moved
to Los Angeles to do postdoctoral work at UCLA. My thesis work had been
listed in a nationally available posting without any address. The Brain Infor-
mation Service, thinking they knew where I was, listed “the other” Jerome
Siegel’s Delaware address for reprint requests. I soon received a letter from
Jerry along with the requests he had received and we have remained in con-
tact ever since. I am occasionally reminded of my namesake when I meet a
new colleague who is impressed that someone “so young” published a paper
in Science in 1965 (one year out of high school, if it had been me). I entered
the field in the early 1970s just as he left. My interests in REM sleep and
brainstem mechanisms have been eerily similar to his (and he also did post-
doctoral work at UCLA), so our research contributions can be distinguished
easily only by my use of my middle initial (which has occasionally been omit-
ted from my publications).
So, my namesake and I both have an interest in seeing to it that no one
“brings shame to the name.” The current work certainly fulfills that dictum.
This is a very unusual book, both in its scope and in its approach to the ma-
terial. It takes us back to the roots of sleep research going from Galvni’s work
in 1791, Berger’s observations of the electroencephalogram, von Economo’s
prescient observations growing out of the encephalitis lethargicia epidemic at
the end of World War I, Kleitman’s scholarly work and codiscovery with
Aserinsky of REM sleep, and many other important historical developments.
He discusses this work clearly and compellingly, presenting not just dates and
names, but rather the motivations and findings of these pioneers. The author
extracts the enduring scientific and technical significance of this early work
in a way that only a sleep scientist could do. The historic context of the early
sleep work is explained. Several of the pioneers of sleep research were caught
up in the turmoil of World War II. The Nazi-driven deaths of Beck and Berger
are rarely presented in histories of the science of brain wave recording. The
interruptions caused by the war and the rapid technological advances that fol-
lowed it are explained.
The book does not talk down to the reader, but rather provides necessary
vii
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viii FOREWORD
background material so that all can follow the main points of the narrative.
When we reach the contemporary issues of sleep science, the work is pre-
sented clearly in a way that is accessible to an interested undergraduate. At
the same time, frequent “I didn’t know that” insights are regularly dished up
for experienced sleep researchers such as myself.
The author presents our current understanding of the sleep disorders that
afflict more than 10% of the population. The mechanisms, as well as accepted
treatments of these disorders are explained in a way that will be useful to
those suffering from sleep disruption. Recent advances in our understanding
of sleep mechanisms and the 24-hour (circadian) rhythms are presented along
with a clear review of current controversies in the field. The recent advances
in our understanding of narcolepsy are presented.
The relatively short length of the book was achieved not by omission of
key findings. Rather it was accomplished by the author’s hard work in clari-
fying and understanding issues that were occasionally lost on the original au-
thors and presenting them in a very approachable manner. New students of
sleep research as well as longtime practitioners will be enlightened by this
outstanding book.
Jerome M. Siegel
Professor of Psychiatry and Biobehavioral Sciences
Center for Sleep Research
University of California, Los Angeles
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Preface
“Thy best of rest is sleep.”
William Shakespeare, Measure for Measure, Act 3, Scene 1
Shakespeare succinctly tells us an important truth. After a day’s activity, a
night spent in bed resting but not sleeping does not refresh us. The next day
we are tired and sleepy and often out of sorts. A more serious consequence
is that we also are prone to accidents. Daytime sleepiness is a major cause of
highway and workplace accidents.
I must admit at the very beginning of the book that how sleep has its ben-
eficial effects and why we need to sleep are not known. Sleep workers have
speculated about those issues, and some of their ideas are described in Chap-
ter 13, Theories of Sleep and Waking. Much of the speculation has been in-
formed by the knowledge gained from the biological research on sleep and
waking gathered during the past 50 years. We still do not know why we sleep,
but we know a fair amount about how we sleep and awaken; that is, we have
learned much about the neural mechanisms that control sleep and waking.
Most of this book is devoted to describing that work.
The modern era of biological research on sleep and waking began at the
turn of the twentieth century. This book surveys that research and briefly in-
cludes nineteenth century work upon which the early twentieth century de-
velopments were built. The early work, until World War II at midcentury, oc-
curred primarily in Europe. During the first half of the twentieth century,
developments in technology and brain research laid the groundwork for the
science of sleep and waking. The benefits of this groundwork were reaped
during the second half-century. After the war, spurred by the Cold War chal-
lenge of the Soviet Union, the United States committed significant resources
to research that led to the burgeoning of technological and scientific advances,
including many in brain research. Although U.S. science was dominant dur-
ing the postwar period, research became increasingly international as the coun-
tries of Europe and Asia recovered from the devastation of the war.
ix
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x PREFACE
This book does not provide an exhaustive review of sleep–waking research
in the twentieth century. Rather, it describes the major advances in the field,
as seen through the eyes of this observer. Some readers may take exception
to what I emphasize and what I leave out. This apology has been stated be-
fore, most elegantly, by Sir Michael Foster, the founder and first head of the
Physiology Department at Cambridge University. In the preface to his book
Lectures on the History of Physiology During the Sixteenth, Seventeenth and Eigh-
teenth Centuries, published in 1901, Foster writes: “I do not pretend to have
given a complete history of physiology even within the period to which I have
limited myself. I have chosen certain themes which seemed to me important
and striking, and I have striven to develop these, leaving untold a great deal
which might be told concerning other themes” (Foster, 1901). From the per-
spective of a century later, there are obvious gaps in Foster’s history. I expect
it will not take anything like a century to point out gaps in my presentation
here.
I have tried to make the research on sleep and waking accessible to readers
who have a background no more extensive than an introductory-level course
in biology or psychology in which the basic facts about the nervous system are
covered. The book has been written for those who would like a survey of the
scientific developments, including the current state of knowledge, in this inter-
esting area of research. The book is also suitable as a text for courses in psy-
chology, biology, or physiology and in the allied health fields. By selecting what
I consider to be the major findings in this field, I have kept the book short. My
goal has been to provide readers with a brief and manageable overview of the
scientific work on sleep and waking. For those interested in more detail and
depth, the references provided in the bibliography would be useful.
As an aid to readers not familiar with the technical terms used in this field,
a glossary is provided at the end of the book. A list of abbreviations used in
this field is also presented. My experience with students who have had an in-
troductory exposure to the nervous system is that they remember the basics
about information transfer in the nervous system, such as facts relating to
spike action potentials, synaptic connections, and neurotransmitters. How-
ever, students are somewhat rusty when it comes to the relative locations of
the major structures within the nervous system. I have, therefore, included a
short chapter at the beginning of the book that reviews anatomical terms and
the basic anatomical organization of the brain—paying more attention to struc-
tures that are relevant to sleep and waking.
The subject matter of a book on sleep and waking could be presented in
a variety of ways. As is often the case in science, research developments are
built upon previous findings and conceptualizations, either as direct follow-
ups or as reactions to earlier work. I have, therefore, chosen a chronological
approach as a logical and meaningful framework for this material. In addi-
tion, the scientific facts and principles are not presented in a social vacuum.
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Preface xi
The reader will learn to appreciate that people from far-flung places like
Bologna, Italy; Jena, Germany; and Krakow, Poland, made important contri-
butions to an area of current interest. This adds a humanizing dimension to
the scientific story.
With respect to research on sleep and waking, the twentieth century di-
vides neatly in half. Research during the first half-century provided the foun-
dation for the major developments that occurred after World War II. How-
ever, as can be seen from the list of the contents of the book, the amount of
space devoted to each of the two half-centuries is uneven. Part I, The First
Half-Century, on historical developments, is considerably shorter than Part II,
The Second Half-Century, in which the major advances made after World War
II are described. The early groundwork for the postwar advances was certainly
important; indeed, the advances in research in the second half-century could
not have occurred without the instrumentation developed in the first 50 years.
Nevertheless, however important these and the other first half-century con-
tributions were, they do not approach the huge amount of work and the ad-
vances in knowledge that followed World War II.
Similarly, in Part II more space is devoted to research on sleep and dream-
ing (Chapters 6–12) than to the waking part of the story (Chapters 4 and 5).
The first major development that ushered in the postwar, modern era of re-
search in this field was published in 1949. This landmark paper was on the
brain mechanisms controlling arousal. However, by 1960, research interest
had shifted to sleep, where, at the close of the twentieth century and the be-
ginning of the twenty-first century, it remains. Consequently, the chapters on
sleeping and dreaming occupy the major share of the book and describe the
current state of knowledge in the field.
The last section of the book, Part III, describes the functions and disorders
of sleep and waking. Chapter 13 presents a number of theories on the func-
tions of sleep. This chapter is divided into three sections: theories related to
the ontogenetic and phylogenetic development of sleep and waking, theories
based on the effects of sleep deprivation, and finally, an area of intense cur-
rent interest, the role of sleep in the process of memory storage. Some of the
findings in sleep and waking research have led to increased understanding
about the disorders of sleep and waking. These research findings, plus an
awareness of the societal costs of sleep and waking disorders, have led to a
new specialization in clinical medicine, that of sleep medicine. Workers
in this field diagnose, treat, and do research on disorders of sleep and wak-
ing. The final chapter of the book describes some of those disorders.
My own background in this area is based on work I had done in the 1960s
and 1970s when I published on the neurophysiology and behavioral aspects
of sleep and waking. While my research in subsequent years turned to other
topics, I have maintained a keen interest in the area of sleep and waking and
have kept up with related developments. In my teaching over the years, I have