Table Of ContentBased on a symposium, "The Maturation of Brain Mechanisms Related to Sleep Behavior,"
held at Boiling Springs, Pennsylvania, June 21—24, 1970, sponsored by The National In-
stitute of Child Health and Human Development, National Institutes of Health.
Sleep and the
Maturing Nervous System
Edited by
CARMINE D. CLEMENTE
Department of Anatomy
School of Medicine
University of California at Los Angeles
Los Angeles, California
DOMINICK P. PURPURA
Department of Anatomy and the Rose F. Kennedy
Center for Research in Mental Retardation and
Human Development
Albert Einstein College of Medicine
Bronx, New York
FLORENCE E. MAYER
Growth and Development Branch
National Institute of Child Health and Human
Development
Department of Health, Education and Welfare
National Institutes of Health
Bethesda, Maryland
®
ACADEMIC PRESS New York and London 1972
COPYRIGHT © 1972, BY ACADEMIC PRESS, INC.
ALL RIGHTS RESERVED
NO PART OF THIS BOOK MAY BE REPRODUCED IN ANY FORM,
BY PHOTOSTAT, MICROFILM, RETRIEVAL SYSTEM, OR ANY
OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM
THE PUBLISHERS.
ACADEMIC PRESS, INC.
Ill Fifth Avenue, New York, New York 10003
United Kingdom Edition published by
ACADEMIC PRESS, INC. (LONDON) LTD.
24/28 Oval Road, London NW1
LIBRARY OF CONGRESS CATALOG CARD NUMBER: 72-84277
PRINTED IN THE UNITED STATES OF AMERICA
To the memory of Dr. Daniele Jouvet-Mounier (1935-1970). Her research on
the maturation of sleep patterns in different species will remain forever a corner-
stone in the building of our knowledge of sleep-wakefulness behavior. Its im-
portance was amply illustrated by the frequent references to her work through-
out the conference and in the pages that follow.
List of Participants
YOSHIO AKIYAMA, Department of Developmental Neurology, University of
Groningen, The Netherlands
THOMAS F. ANDERS, Department of Psychiatry, Albert Einstein College of
Medicine at Montefiore Hospital, Bronx, New York
FLOYD E. BLOOM, Laboratory of Neuropharmacology, Division of Special
Mental Health Research, National Institute of Mental Health, Saint Elizabeth
Hospital, Washington, D.C.
DAVID BODIAN, Department of Anatomy, Johns Hopkins University School of
Medicine, Baltimore, Maryland
MICHAEL H. CHASE, Departments of Anatomy and Physiology, University of
California, School of Medicine, Los Angeles, California
CARMINE D. CLEMENTE, Department of Anatomy, University of California,
School of Medicine, Los Angeles, California
PAUL D. COLEMAN, Department of Anatomy, University of Rochester School
of Medicine and Dentistry, Rochester, New York
WILLIAM C. DEMENT, Department of Psychiatry, Stanford University, School
of Medicine, Stanford, California
JOHN E. DESMEDT, Brain Research Unit, University of Brussels, Brussels,
Belgium
ROBERT J. ELLINGSON, Nebraska Psychiatric Institute, University of
Nebraska, Medical Center, Omaha, Nebraska
Xlll
XIV List of Participants
WALTER B. ESSMAN, Department of Psychology, Queens College, C.U.N.Y.,
Flushing, New York
HARRY H. GORDON, Rose Fitzgerald Kennedy Center for Research in Mental
Retardation and Human Development, Albert Einstein College of Medicine,
Bronx, New York
WILLIAMINA A. HIMW1CH, Thudichum Psychiatric Research Laboratory,
Galesburg State Research Hospital, Galesburg, Illinois
GABRIELE HINZE, Department of Pediatrics, University of Gottingen, Göt-
tingen, Germany
BARRY J. HOFFER, Laboratory of Neuropharmacology, Division of Special
Mental Health Research, Saint Elizabeth Hospital, Washington, D.C.
NATHANIEL KLEITMAN, 222 Washington Avenue, Santa Monica, California
DONALD B. LINDSLEY, Departments of Psychology, Physiology, and Psy-
chiatry, and Brain Research Institute, University of California, Los Angeles,
California
DENNIS J. McGINTY, Neuropsychology Research, Veterans Administration
Hospital, Sepulveda, California
DAVID R. METCALF, Department of Psychiatry, University of Colorado
Medical Center, Denver, Colorado
PETER J. MORGANE, Laboratory of Neurophysiology, Worchester Foundation
for Experimental Biology, Shrewsbury, Massachusetts
EDWARD M. ORNITZ, Department of Psychiatry, University of California,
School of Medicine, Los Angeles, California
ARTHUR H. PARMELEE, JR., Department of Pediatrics, University of Cal-
ifornia, School of Medicine, Los Angeles, California
H. F. R. PRECHTL, Department of Developmental Neurology, University of
Groningen, The Netherlands
JAMES W. PRESCOTT, National Institute of Child Health and Human Devel-
opment, Bethesda, Maryland
DOMINICK P. PURPURA, Department of Anatomy and the Rose F. Kennedy
Center for Research in Mental Retardation and Human Development, Albert
Einstein College of Medicine, Bronx, New York
OLGA PETRE-QUADENS, Department of Developmental Neurology, Born-
Bunge Research Foundation, Berchem-Antwerp, Belgium
OSCAR RESNICK, Worchester Foundation for Experimental Biology, Shrews-
bury, Massachusetts
List of Participants xv
EUGENE ROBERTS, Division of Neurosciences, City of Hope National Medical
Center, Duarte, California
HOWARD ROFFWARG, Department of Psychiatry, Albert Einstein College of
Medicine at Montefiore Hospital, Bronx, New York
GUENTER H. ROSE, Laboratories of Developmental Psychobiology, University
of Nebraska Medical School, Omaha, Nebraska
MORTIMER G. ROSEN, Department of Obstetrics-Gynecology, University of
Rochester School of Medicine and Dentistry, Rochester, New York
ARONOLD B. SCHEIBEL, Departments of Anatomy and Psychiatry, University
of California, School of Medicine, Los Angeles, California
MADGE E. SCHEIBEL, Departments of Anatomy and Psychiatry, University of
California, School of Medicine, Los Angeles, California
GERLIND SCHREMPF, Department of Pediatrics, University of Göttingen, Göt-
tingen, Germany
F. J. SCHULTE, Department of Pediatrics, University of Göttingen, Göttingen,
Germany
ROBERT J. SHOFER, Department of Anatomy and the Rose F. Kennedy
Center for Research in Mental Retardation and Human Development, Albert
Einstein College of Medicine, Bronx, New York
GEORGE R. SIGG1NS, Laboratory of Neuropharmacology, Division of Special
Mental Health Research, National Institute of Mental Health, Saint Elizabeth
Hospital, Washington, D.C.
M. B. STERMAN, Neuropsychology Research, Veterans Administration Hos-
pital, Sepulveda, California
EVELYN STERN, Department of Pediatrics, University of California, Center for
Health Sciences, Los Angeles, California
ELLIOT D. WEITZMAN, Department of Neurology, Albert Einstein College of
Medicine, Bronx, New York
DONALD J. WOODWARD, Department of Physiology, School of Medicine and
Dentistry, University of Rochester, Rochester, New York
Preface
It is now well established that both genetic and environmental factors play
important roles in determining the morphological and electrophysiological
maturation of the brain. What is not known, however, is how such maturational
features contribute to the progressive elaboration of complex neural interactions
that characterize the development of sleep-wakefulness activities in the maturing
organism. There can be little doubt that this problem is central to the issue of
the development of complex behaviors in general. This follows from the fact
that such behaviors are expressed upon a background of variable wakefulness
which requires the integrative activity of many different neuronal organizations.
But equally impressive is the fact that transitions from sleep to wakefulness and
vice versa bring into operation perhaps more complex neural and biochemical
mechanisms. Thus the understanding of how maturational processes are related
to the development of sleep-wakefulness behavior should provide information
essential for the adequate appreciation of the biological basis of behavior in the
broadest sense.
The conference, held at the Allenberry Inn in Boiling Springs, Pennsylvania
from June 21-24, 1970, was organized to explore the extent to which basic data
on brain maturation and data on the ontogeny of sleep-wakefulness behavior
might be discussed from the standpoint of their mutual relevancy. Additional
aspects of the conference focused on the developmental implications of con-
ditions, experimental or otherwise, which might lead to the establishment of
normal and abnormal sleep behavior in maturing organisms. The program was
divided into five sessions: (1) The Maturation of Neural Elements, (2) Neuro-
chemical Factors in Maturation of Sleep Behavior, (3) Development of EEG and
Activity Patterns in Relation to Sleep, (4) Development of Reflex Patterns in
xvn
XV111 Preface
Sleep, and (5) Developmental Aspects of Normal and Abnormal Sleep Behavior.
The proceedings were summarized by Dr. Donald B. Lindsley.
The editors of this symposium volume were joined in the planning stages of
the conference by Dr. Robert J. Ellingson, Dr. Donald B. Lindsley, and Dr.
Merrill S. Read. Grateful acknowledgment is also made to Miss Wanda Burnett
for editorial assistance in the preparation of the final manuscript and to the
Growth and Development Program of the National Institute of Child Health and
Human Development whose enthusiastic support and sponsorship of the con-
ference made this volume possible.
Carmine D. Clemente
Dominick P. Purpura
Florence E. Mayer
1
Principles of Synaptogenesis and Their
Application to Ontogenetic Studies
of Mammalian Cerebral Cortex*
Dominick P. Purpura and Robert J. Shofer
Newborn altricial mammals exhibit relatively little awake behavior (crying,
crawling, sucking, etc.) and quiet sleep in contrast to the large proportion of
time they spend in "active" or rapid eye movements sleep (REMS). Maturation
of sleep-wakefulness cycles results in an increasing capacity for wakefulness, as
Kleitman has correctly insisted, and a reorganization of temporospatial patterns
of various sleep states. Although the neurobiological processes underlying these
developmental events are poorly understood, maturation of sleep-wakefulness
behavior is evidently as much a consequence of the reorganization of early
established patterns of activity as it is the elaboration of new modes of operation
in developing neuronal subsystems. Considering the prominence of the physio-
logical mechanisms subserving active sleep in the altricial neonate, it is difficult
♦Supported in part by grants from NINDS (NS-07512) and The Given Foundation.
3
