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PROGRESS IN BRAIN RESEARCH VOLUME 100 NEUROSCIENCE: FROM THE MOLECULAR TO THE COGNITIVE EDITED BY FLOYD E. BLOOM Department of Neuropharmacologv, The Scripps Clinic and Research Institute, La Jolla, CA 92307, USA ELSEVIER AMSTERDAM - LONDON -NEW YORK - TOKYO 1994 8 1994 Elsevier Science B.V. 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 the written permission of the Publisher, Elsevier Science B.V., Copyright & Permissions Department, P.O. Box 521, 1000 Ah4 Amsterdam. The Netherlands. No responsibility is assumed by the Publisher for any injury and/or damage to persons or property as a matter of products liability, negligence or otherwise, or from any use or operation of any methods, products, instructions or ideas contained in the material herein. Because of the rapid advances in the medical sciences, the publisher recommends that independent verification of diagnoses and drug dosages should be made. Special regulations for readers in the U.S.A.: This publication has been registered with the Copyright Clearance Center Inc. (CCC) Salem, Massachusetts. Information can be obtained from the CCC about conditions under which photocopies of parts of this publication may be made in the U.S.A. All other copyright questions, including photocopying outside of the USA, should be referred to the Publisher. ISBN 0-444-81678-x (volume) ISSN 0-444-80104-9 (series) Elsevier Science B.V. P.O. Box 21 1 1000 AE Amsterdam The Netherlands Library of Congress Cataloging-in-Publication Data Neuroscience : from the molecular to the cognitive / edited by Floyd E. Bloom. p. CR. -- (Progress in brain research ; v. 100) Includes bibliographical references and index. ISBN 0-444-81678-X (alk. paper). -- ISBN 0-444-80104-9 (series : alk. paper) 1. Neurosciences. I. Bloom, Floyd E. 11. Series. [DNLM: 1. Central Nervous System--physiology. W1 PR667 v.100 1994 / WL 300 N4939 19941 QP355.2.N49 1994 612.8'2--dc20 DNLM/DLC for Library of Congress 94- 12298 CIP Printed on acid-free paper Printed in The Netherlands V List of Contributors H. Akil, Mental Health Research Institute, Department of Psychiatry, University of Michigan Medical School, Ann Arbor, MI 48105, USA D. B&, Department of Neurology, Rudolf Magnus Institute for Neurosciences, Utrecht University, Universiteitsweg 100,3584 CG Utrecht, The Netherlands F.E. Bloom, Department of Neuropharmacology, The Scripps Research Institute, La Jolla, CA, USA H. Cameron, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA T.J. Carew, Department of Psychology, Yale University, POB 1 I-A, Yale Station, New Haven, CT 06520, USA H.M. Chao, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA D.W. Choi, Department of Neurology and Center for the Study of Nervous System Injury, Box 81 1 I, Washington University School of Medicine, 660 S. Euclid Avenue, St. Louis, MO 631 10, USA A.C. Cuello, McGill University, Department of Pharmacology and Therapeutics, McIntyre Medical Building, 3655 Drummond Street, Suite 1325, Montreal, Quebec, Canada H3G 1Y6 N.C. Danbolt, Anatomical Institute, University of Oslo, P.O.B. 1105 Blindem, N-0317 Oslo, Norway I. Divac, Department of Medical Physiology, Panum Institute, University of Copenhagen, Blegdamsvej 3C, DK- 2200 Copenhagen, Denmark N.J. Emptage, Department of Psychology, Yale University, 2 Hillhouse Avenue, Box 11A Yale Station, New Haven, CT 06520, USA S.Y. Felten, Department of Neurobiology and Anatomy, Box 603, University of Rochester School of Medicine and Dentistry, 601 Elmwood Avenue, Rochester, NY. 14642, USA D.L. Felten, Department of Neurobiology and Anatomy, Box 603, University of Rochester School of Medicine and Dentistry, 601 Elmwood Avenue, Rochester, NY. 14642, USA G. Fink, MRC Brain Metabolism Unit, University Department of Pharmacology, 1 George Square, Edinburgh EH8 9JZ, UK L. Giovannelli, Department of Preclinical and Clinical Pharmacology, University of Florence, Viale Morgagni 65, 50 134 Florence, Italy W.H. Gispen, Department of Medical Pharmacology, Rudolf Magnus Institute for Neurosciences, Utrecht University, Universiteitweg 100,3584 CG Utrecht, The Netherlands E. Gould, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA W.T. Greenough, Beckman Institute, Department of Psychology, Neuroscience Program and Department of Cell and Structural Biology, University of Illinois, Urbana, IL, USA M.E.R. Hallonet, Institut d’Embryologie Cellulaire et molkculaire, du CNRS et du College de France, 49bis avenue de la belle Gabrielle, 94736 Nogent-sur-Mame Cedex, France R. Hari, Low Temperature Laboratory, Helsinki University of Technology, 02150 Espoo, Finland M.A. Hofman, Graduate School of Neurosciences Amsterdam, Netherlands Institute for Brain Research, Meibergdreef 33, 1105 AZ Amsterdam, The Netherlands B.T. Hyman, Neurology Service, Massachusetts General Hospital and Harvard Medical School, Boston, MA 02114, USA G. Jaim-Etcheveny, Department of Cell Biology and Histology, School of Medicine, University of Buenos Aires, Paraguay 2155, 1121 Buenos Aires, Argentina G.A.R. Johnston, Department of Pharmacology, The University of Sydney, NSW, 2006, Australia J.D. Kocsis, Department of Neurology, Yale University School of Medicine, New Haven, CT 06510; and PVAEPVA Neuroscience Research Center, VA Hospital, West Haven, CT 065 16, USA S.C. Landis, Department of Neurosciences, Case Western Reserve University, School of Medicine, Cleveland, OH 44 106, USA N.M. Le Douarin, Institut d‘Embryologie Cellulaire et mol6culaire, du CNRS et du Colkge de France, 49bis avenue de la belle Gabrielle, 94736 Nogent-sur-Marne Cedex, France VI V. Luine, Department of Psychology, Hunter College, New York 10021, NY, USA A.M. Magarinos, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021 , USA P.J. Magistretti, Institut de Physiologie, Facult6 de MCdecine, Universit6 de Lausanne, CH-1005 Lausanne, Switzerland E.A. Marcus, Department of Biology, Yale University, 2 Hillhouse Avenue, Box 11A Yale Station, New Haven, CT 06520, USA R. Marois, InterdepartmentalN euroscience Program, Yale University, 2 Hillhouse Avenue, Box 11A Yale Station, New Haven, CT 06520, USA B.S. McEwen, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA J. Mendlewicz, Department of Psychiatry, Free University Clinics of Brussels, Erasme Hospital, route de Lennik 808, 1070 Brussels, Belgium P.B. Molinoff, Department of Pharmacology, University of Pennsylvania School of Medicine, Philadelphia, PA 19104-6084, USA O.P. Ottersen, Anatomical Institute, University of Oslo, P.O.B. 1105 Blindem, N-0317 Oslo, Norway C. Pavlides, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA G. Pepeu, Department of Preclinical and Clinical Pharmacology, University of Florence, Viale Morgagni 65, 50134 Florence, Italy 0. Pompeiano, Dipartimento di Fisiologia e Biochimica, Via S. Zen0 31,56127 Pisa, Italy 0. PourquiC, Institut d'Embryologie Cellulaire et molkculaire, du CNRS et du Colltge de France, 49his avenue de la belle Gabrielle, 94736 Nogent-sur-Mame Cedex, France D.B. Pritchett, Department of Pharmacology, University of Pennsylvania School of Medicine, Philadelphia, PA 191 04-6084, USA G. Raisman, Norman and Sadie Lee Research Centre, Laboratory of Neurobiology, National Institute for Medical Research, The Ridgeway, Mill Hill, London NW7 IAA, UK R. Ranney Mix, Department of Anatomy and the Neuroscience Center, Louisiana State University Medical Center, 1901 Perdido Street, New Orleans, LA 701 12, USA G.W. Rebeck, Neurology Service, Massachusetts General Hospital and Harvard Medical School, Boston, MA 02114, USA L.P. Renaud, Neurosciences Unit, Loeb Research Institute, Ottawa Civic Hospital and University of Ottawa, Ottawa, Ontario, Canada KIY 4E9 J.L. Roberts, Dr. Arthur M. Fishberg Research Center for Neurobiology, Mount Sinai School of Medicine, One Gustave Levy Place, New York, NY 10029, USA P. Rudomin, Department of Physiology, Biophysics and Neurosciences, Centro de Investigaci6n y de Estudios Avanzados, MCxico D.F M. Schachner, Department of Neurobiology, Swiss Federal Institute of Technology, 8093 Zurich, Switzerland N.A. Simonian, Neurology Service, Massachusetts General Hospital and Harvard Medical School, Boston, MA 02114, USA R.L. Spencer, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA C.N. Stefanis, Department of Psychiatry, Athens University Medical School, Eginition Hospital, 72-74 Vas Sophias Ave., 115 28 Athens, Greece D.G. Stein, Institute of Animal Behavior, Rutgers, The State University of New Jersey, Newark, NJ 07102, USA J. Storm-Mathisen, Anatomical Institute, University of Oslo, P.O.B. 1105 Blindem, N-0317 Oslo, Norway D.F. Swaab, Graduate School of Neurosciences Amsterdam, Netherlands Institute for Brain Research, Meibergdreef 33, 1105 AZ Amsterdam, The Netherlands D.A. Utzschneider, Department of Neurology, Yale University School of Medicine, New Haven, CT 06510; and PVAEPVA Neuroscience Research Center, VA Hospital, West Haven, CT 065 16, USA J. Verhaagen, Department of Medical Pharmacology, Rudolf Magnus Institute for Neurosciences, Utrecht University, Universiteitweg 100,3584 CG Utrecht, The Netherlands X. Wang, Neuroscience Program, University of Illinois, Urbana, IL, USA Y. Watanabe, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA S.J. Watson, Mental Health Research Institute, Department of Psychiatry, University of Michigan Medical School, Ann Arbor, MI 48105, USA VII S.G. Waxman, Department of Neurology, LCI 708 Yale School of Medicine 333 Cedar Street New Haven, CT 065 10, USA I.J. Weiler, Beckman Institute and Department of Psychology, University of Illinois, Urbana, IL, USA K. Williams, Department of Pharmacology, University of Pennsylvania School of Medicine, Philadelphia, PA 19104-6084, USA C. Woo:ley, Laboratory of Neuroendocrinology, Rockefeller University, 1230 York Avenue, New York, NY 10021, USA W.G. Young, Department of Neuropharmacology, The Scripps Research Institute, La Jolla, CA, USA J. Zhong, Department of Pharmacology, University of Pennsylvania School of Medicine, Philadelphia, PA 19104- 6084, USA M.J. Zigmond, Department of Neuroscience, University of Pittsburgh, 570 Crawford Hall, Pittsburgh, PA 15260, USA IX Preface All editors like to regard their volumes as unique. Reflecting back through the series of the prior 99 volumes in this series, that premise is strongly supported. However, in all modesty, we would have to assert that the present volume carries at least two features that distinguish it fiom its predecessor volumes in the series: (1) it is a commemorative milestone issue; and (2) unlike most of the prior volumes, it does not provide an overview of a specific scientific meeting. Instead, for appropriate and creative reasons on the part of the Publisher, Elsevier Science, Amsterdam, it was determined to have a special number in the series to commemo- rate the one hundredth volume. The question then evolved as to exactly what form this com- memorative effort would take and how it would be organized. After due consideration of a variety of ideas by the editorial advisory board to the Progress in Brain Research series, the idea was put forward that the volume should reflect the views of the international body of active neuroscientists, to report on contemporary top- ics of their choice. Accordingly, authors were invited to write on any topic, given that their choice represented the topic most near and dear to their own efforts over a significant period of the recent past, and to which they would likely continue to be devoted in the future. The authors were also urged where possible to reflect on the evolution of their selected topics and their likely future course. In that sense this volume reflects not “just” a single scientific meeting, but rather an overview sample of the problems and methodologies that epitomize brain research broadly at this special moment in the maturation of the field. The editorial advisory board were polled for their recommendations of the scientists to be invited, and the inevitably required selec- tion of countries. The collection of essays in this volume therefore reflect these choices, and in addition, another inevitable reflection of the hectic environment of scientific publications, many of our invitees who accepted, were in the end, simply unable to provide their chapters within the deadline. Given this background of its origination, it seems remarkable that the chapters comprising this volume assorted themselves so readily into five or six easily established categories of topics: developmental brain research, molecular brain research, integrative brain research, neuroplasticity, and neuro-psychiatric conditions. These topics do indeed seem to reflect well on the major streams of effort of our field when measured against the indices of the leading journals. We are indebted to the authors for their diligent efforts to provide their chapters rapidly. We offer here a volume which reports on a sample of recognized leaders in the neuroscien- tific community at a significant instant in the history of this renowned series and in the evo- lution of the field. It is likely, given the high momentum of progress in this field, that many X of these topics, and some wholly unexpected by our authors, will establish new directions for the next one hundred volumes. F.E. Bloom The Scripps Research Institute, La Jolla, CA N. Spiteri Elsevier Science, Amsterdam XI Then and Now Dominick P. Pwpura President, IBRO Despite the harsh political cliate that characterized the Cold War in 1961, scientists dedi- cated to the understanding of brain mechanisms overcame communication barriers to attend a “Colloquium on Brain Mechanisms” sponsored by the International Brain Research Or- ganization. The central theme of the Colloquium was “Sensorimotor Integration”. The formal papers and extensive discussions appeared as Volume One of Progress in Brain Research in 1963. Elsevier’s prescience in recognizing the emergence of brain research as a major growth industry in the life sciences may have been as important as the reports of the Collo- quium. Not surprisingly, much of what was discussed during the 1961 Pisa meeting was but a prologue to current problems. Alfted Fessard provided the Colloquium’s last words: “In conclusion, it seems that in the general case of a multineuronal assembly engaged in a spe- cific sensori-motor operation, the question of a special mechanism destined to confer upon it in isolation the quality of an ‘entity’ does not really exist. It can only be called ‘integrated’ in relation to all other congruent assemblies, at all levels of the neuraxis, which participate in the particular operation”. Three decades later “integration” remains the central goal of brain research, its Holy Grail - now defined as vertical integration fiom “the Molecular to Cognition”, the theme of the present milestone volume of Progress in Brain Research. In the intervening years since the first volume, no aspect of the universe of discourse referred to now as “neuroscience” has escaped close pursuit in the hundred volumes of Progress. Examples abound here from con- siderations of the molecular subunit structure of transmembrane channels to complex cogni- tive processes. In keeping with the international spirit of the inaugural volume, the Editor has succeeded in encouraging sixty-six neuroscientists ftom fourteen countries to examine the canon of brain research and this they have accomplished with due respect for the constraints of space. Remarkable is it also that the Editor was able to elicit from many contributors their views and speculations and even some of their most prophetic visions. Milestones give pause for review to examine the present and map the future course of discovery. It was not intended that this commemorative issue of Progress in Brain Research would be a comprehensive repository of the most recent advances in neuroscience. Rather it may be viewed as an affirmation of the coming of age of brain research as a scholarly endeavor that now permits objective inquiry into problems that have perplexed humankind since the dawn of human consciousness. To speak about “brain mechanisms” 30 years ago was to infer cau- sality without process. Today even the most complex brain mechanisms are explicable in terms of cellular, molecular and genetic events: Reductionism as champion not challenger of Integration. None of the participants of the 1961 Colloquium ventured into the forbidden realm of pathobiology, unable as they were to grasp at fundamentals of molecular patho- genesis. Today, as this volume attests, neuroscience is at the threshold of comprehending the essential mechanisms that give rise to Alzheimer’s disease, epilepsy, stroke, multiple sclero- sis and serious mental disorders. Translating understanding to application, though arduous, augers well for the future of brain research and the human condition. The Editor is to be further congratulated for alerting us in his own contribution to the ineluctable problem of data overload on the “infobahn”. How neuroscience deals with the appalling mass of “facts” being disgorged daily from thousands of laboratories throughout the world will be as critical to the success of the neuroscience enterprise as any number of new conceptual advances. The world community of neuroscientists in IBRO now consists of more than 30 000 neuroscientists in over 70 countries. Considering the intellectual power of this workforce, it will surely not be another 30 years before the appearance of the 200th vol- ume of Progress in Brain Research. Surely, the need for rapid communication will only in- tensify as new technologies drive new advances and vice versa. Progress in Brain Research will continue to inform us about the next revolution in neuroscience, the understanding of how the brain works in health and disease. But whether the new canon will be conveyed on paper or via the “infobahn” remains to be seen. Elsevier was midwife to the Progress series and hand maiden to brain research. Now we celebrate the marriage of neuroscience and in- formatics. The offspring of the consummation will further test Elsevier’s prescience. F. Bloom (Editor) Progress in Brain Research, Vol. 100 0 1994 Elsevier Science B.V.A ll rights reserved 3 CHAPTER 1 Cell migrations and establishment of neuronal connections in the developing brain: a study using the quail-chick chimera system Nicole M. Le Douarin, Marc E.R. Hallonet and Olivier Pourquie Institut d’EmbryologieC ellulaire et molkculaire, du CNRS et du Coll2ge de France, 49bis avenue de la belle Gabrielle, 94736 Nogent-sur-MarneC edex, France Introduction development of the brain at the rostra1 end of the neu- ral tube (Gans and Northcutt, 1983). The process of In vertebrates, the first morphological sign of emer- cephalization involves the activity of a number of gence of the nervous system is the appearance of the genes (Simeone et al., 1992) as well as an intricate neural plate, the medio-dorsal line of which is in inti- network of intercellular signalling. An important re- mate contact with a mesodermal structure, the noto- quirement in attempts to decipher how the brain is chord. Laterally, the neural epithelium is separated built is to be able to follow embryonic cells within the from the presumptive superficial ectoderm by a tran- neural primordium while the complexity arises, i.e. sitional zone, the neural fold. The next important step from the early stages of the neural plate up to comple- in neurogenesis is the transformation of the initially tion of neurogenesis. This involves the construction of flat neural anlage into a tubular structure. The dorso- fate maps and the tracing of neuroepithelial cells along medial part of the neural plate thus becomes medio- their developmental history. By using the quail-chick ventral and its lateral ridges reach the medio-dorsal chimera system we have undertaken a study in which line where they fuse to form the neural crest and the movements and fate of the neuroepithelial territories roof plate. During these morphogenetic events, the were followed in the embryo in ovo during the entire floor plate acquires important inductive properties un- period of development. We have thus been able to sort der the influence of the notochord (Yamada et a]., out what is the respective contribution of the alar and 1991) and the neural tube becomes divided into six basal plates of the mes- and rhombencephalon to compartments along the dorsoventral axis: ventrally structures like the cerebellum and various brain stem the floor plate in contact with the notochord, the roof nuclei (Hallonet et al., 1990; Tan and Le Douarin, plate dorsally, and laterally the alar and basal plates I 99 1 ; Hallonet and Le Douarin, 1993). Moreover, we corresponding respectively to the dorsal and ventral have isolated an immunoglobulin-like (Ig-like) cell quarters of the neural tube. Although these territories surface glycoprotein (called BEN) whose expression have long been recognized and named, it is only re- is developmentally regulated in definite subsets of cently that their developmental significance is being neurons (Pourquik et al., 1990, 1992a,b). On the basis really investigated. of fate mapping results at the mid- and hindbrain lev- One of the characteristics of vertebrates among the els and combinations of the quail-chick chimerism chordates is the emergence of the neural crest and the with anti-BEN immunoreactivity, we could show that

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Most books are stored in the elastic cloud where traffic is expensive. For this reason, we have a limit on daily download.