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Metabolism of the Nervous System PDF

640 Pages·1957·15.116 MB·English
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METABOLISM OF THE NERVOUS SYSTEM Edited by DEREK RICHTER Neuropsychiatrie Research Centre Whitchurch Hospital, Cardiff PERGAMON PRESS L O N D ON - N EW Y O RK P A R IS · LOS A N G E L ES PERGAMON PRESS LTD. 4 & 5 Fitzroy Square, London W.l. PERGAMON PRESS INC. 122 East 55th Street, New York 22, N.Y. 10638 South Wilton Place, Los Angeles 47, California PERGAMON PRESS S.A.R.L 24 Rue des Écoles, Paris V". Copyright © 1957 Pergamon Press Ltd. Library of Congress Card No. 57-13324 Printed in Great Britain at The Chapel River Press, Andover PREFACE contains the proceedings of the 2nd International Neuro- THIS VOLUME chemical Symposium, held at Aarhus, Denmark, in July 1956. It represents a collective effort by a group of investigators working in different fields of research, but having a common interest in the working of the nervous system in health and in disease. In the arrangement of the Symposium a small group of scientists were first invited to prepare reviews on a series of selected topics deemed to be of outstanding importance in relation to this problem. The reviews were then supplemented by a limited number of short papers designed to bring out recent work of special interest, or to add to the diversity of view- points by reason of the different academic, methodological or geographical backgrounds of their authors. The data derived from these different sources were then brought together at the symposium and submitted to detailed critical discussion. The responsibility for arranging the Symposium fell on the Organizing Committee, and they must take the blame for any shortcomings in the scientific programme. The difficulties of selection and of limitation of numbers in a conference of this kind are very well known, and in this case the problem was not made easier by the success of the previous Neurochemical Symposium in 1954. Some who should have been invited were inevitably omitted: but the whole Committee were unanimously agreed on all the major decisions that had to be made. The First Symposium dealt mainly with the embryology and early develop- ment of the nervous system, and the proceedings were published under the title 4 Biochemistry of the Developing Nervous System ' (Academic Press, 1955). The Second Symposium supplements and extends the previous one by giving a fuller account of the metabolism of adult nervous tissues. As before, the subject has been approached along the broadest possible lines and an attempt has been made to relate metabolism to structure and function, rather than to deal exclusively with the biochemistry of nervous tissues. The meeting proceeded in an atmosphere in which questions could be asked, criticism could be made and differences of opinion could be thrashed out. There must have been few who did not learn something fresh from the meeting. From an editorial point of view, the extent to which the manuscripts originally sub- mitted needed subsequently to be revised was probably an indication of the success of the meeting. We are indebted to Dr. G. B. Ansell for his help with the editorial work. To our Danish hosts in the University of Aarhus we owe a special debt of gratitude, not only for their exceptional kindness and hospitality during the meeting, but also for providing in their fine modern University a perfect setting for the serious work of the conference. It is a pleasure to express our thanks to the Rector of Aarhus University, Professor Willy Munck, to Professor ix Strömgren, to Professor Einarson and to Professor Sch0nheyder for the im- portant contributions they made to the success of the meeting. To Dr. Mogens Schou and his Staff we are also greatly indebted for their help in dealing with the administrative arrangements for the Symposium. Cardiff DEREK RICHTER November 1956 ACKNOWLEDGEMENTS IT ISA pleasure to thank the private and governmental agencies who sponsored the attendance of delegates to the Symposium. These bodies include the Rockefeller Foundation and the Division of Behavioural Sciences of the Ford Foundation (grants administered by Columbia University, New York); the Mental Health Research Fund; the Wellcome Trust; the Academy of Sciences of the U.S.S.R. ; the French Foreign Office (Relations Culturelles); the University of Paris; Hessischer Minister für Erziehung und Volksbildung; Auswärtiges Amt der Bundesrepublik Deutschland; Stiftung zur Erforschung der Spinalen Kinderlähmung und Multiplen Sklerose; Norges Almenvidenskapelige Forskningsfond; Rask-0rsted Fondet; Aarhus Oliefabriks Fond; the Welsh Regional Hospitals Board. We are also indebted to the following for permission to reproduce published material: The Royal Society; the Journal of Physiology (Cambridge University Press); the American Journal of Physiology; the Long Island Biological Association. ÷ Strömgren, to Professor Einarson and to Professor Sch0nheyder for the im- portant contributions they made to the success of the meeting. To Dr. Mogens Schou and his Staff we are also greatly indebted for their help in dealing with the administrative arrangements for the Symposium. Cardiff DEREK RICHTER November 1956 ACKNOWLEDGEMENTS IT ISA pleasure to thank the private and governmental agencies who sponsored the attendance of delegates to the Symposium. These bodies include the Rockefeller Foundation and the Division of Behavioural Sciences of the Ford Foundation (grants administered by Columbia University, New York); the Mental Health Research Fund; the Wellcome Trust; the Academy of Sciences of the U.S.S.R. ; the French Foreign Office (Relations Culturelles); the University of Paris; Hessischer Minister für Erziehung und Volksbildung; Auswärtiges Amt der Bundesrepublik Deutschland; Stiftung zur Erforschung der Spinalen Kinderlähmung und Multiplen Sklerose; Norges Almenvidenskapelige Forskningsfond; Rask-0rsted Fondet; Aarhus Oliefabriks Fond; the Welsh Regional Hospitals Board. We are also indebted to the following for permission to reproduce published material: The Royal Society; the Journal of Physiology (Cambridge University Press); the American Journal of Physiology; the Long Island Biological Association. ÷ Second International Neurochemical Symposium ORGANIZING COMMUTEE L. EINARSON J. FOLCH-PI D. RICHTER J. ELKES G. W. HARRIS F. SCHONHEYDER L. FLEXNER S. S. KETY H. WAELSCH PARTICIPANTS L. BAKAY D. A. CLOUET Mass. General Hospital, Neuropsychiatrie Research Centre, Boston, Mass. (U.S.A.) Whitchurch Hospital, Cardiff (U.K.) H. BAUER University of Hamjburg, R. V. COXON Hamburg (Germany) University of Oxford, Oxford (U.K.) S. BJÖRKERUD University of Gothenburg, J. CROSSLAND Gothenburg (Sweden) Physiology Department, St. Andrews (U.K.) P. B. BRADLEY University of Birmingham, J. N. CUMINGS Birmingham (U.K.) National Hospital, London (U.K.) G. BRANTE Centrallasarettet, S. DARLING Aarhus University, Eskilstuna (Sweden) Aarhus (Denmark) S.-O. BRATTGÂRD University of Gothenburg, G. B. DAVID Neuropsychiatrie Research Centre, Gothenburg (Sweden) Whitchurch Hospital, J. B. BRIERLEY Cardiff (U.K.) Institute of Psychiatry, London (U.K.) P. DlEZEL University of Heidelberg, Heidelberg (Germany) F. BRINK, JR. Rockefeller Inst, for Medical Research, G. W. F. EDGAR New York (U.S.A.) University of Amsterdam, Amsterdam (Holland) F. BUCHTHAL University of Copenhagen, J.-E. EDSTRÖM Copenhagen (Denmark) University of Gothenburg, Gothenburg (Sweden) G. L. CANTONI National Institutes of Mental Health, L. EINARSON Bethesda, Md. (U.S.A.) Aarhus University, Aarhus (Denmark) H. I. CHINN Office of Naval Research, J. ELKES London Branch, University of Birmingham, New York (U.S.A.; Birmingham (U.K.) xii SECOND INTERNATIONAL NEUROCHEMICAL SYMPOSIUM K. A. C. ELLIOTT H. HARTELIUS McGill University, Karolinska Institutet, Montreal (Canada) Stockholm (Sweden) A. ENGSTRÖM E. JACOBSEN Karolinska Institutet, Medicinalco Ltd., Stockholm (Sweden) Copenhagen (Denmark) U. S. v. EULER Karolinska Institutet, H. JATZKEWITZ Stockholm (Sweden) Max-Planck-Institut, Munich (Germany) W. FELDBERG National Institute for Medical Research, S. S. KETY London (U.K.) National Institute of Mental Health, J. B. FINEAN Bethesda, Md. (U.S.A.) University of Birmingham, R. D. KEYNES Birmingham (U.K.) University of Cambridge, L. B. FLEXNER Cambridge (U.K.) University of Pennyslvania, Philadelphia, Pa. (U.S.A.) E. KLENK University of Cologne, J. FOLCH-PI Cologne (Germany) Harvard Medical School and McLean Hospital, V. KLINGMÜLLER Waverly, Mass. (U.S.A.) University of Hamburg, A. GEIGER Hamburg (Germany) University of Illinois, Chicago, 111. (U.S.A.) V. KOEFOED-JOHNSEN University of Copenhagen, R. GEIGER Copenhagen (Denmark) University of Illinois, Chicago, 111. (U.S.A.) E. KROGH R. W. GERARD Aarhus University, University of Michigan, Aarhus (Denmark) Ann Arbor, Mich. (U.S.A.) M. G. LARRABEE L. GJESSING Johns Hopkins University, Dikemark, Baltimore (U.S.A.) Oslo (Norway) N. A. LASSEN R. GJESSING Copenhagen University, Dikemark, Copenhagen (Denmark) Oslo (Norway) S. GLUECKSON-WAELSCH P. R. LEWIS Albert Einstein College of Medicine, University of Cambridge, New York (U.S.A.) Cambridge (U.K.) D. E. GOLDMAN F. LIPMANN U.S. Embassy, Harvard University and Mass. London (U.K.) General Hospital, P. GREENGARD Boston, Mass. (U.S.A.) National Institute for Medical Research, London (U.K.) R. LIVINGSTON National Institute of Mental Health, Mrs. P. GREENGARD Bethesda, Md. (U.S.A.) The Institute of Psychiatry, London (U.K.) K. A. LORENTZEN R. G. GRENELL University of Aarhus and Risskov University of Maryland, Mental Hospital, Baltimore, Md. (U.S.A.) Risskov (Denmark) SECOND INTERNATIONAL NEUROCHEMICAL SYMPOSIUM XHl A. LOWENTHAL Ο. Ε. PRATT Institut Bunge, Institute of Psychiatry, Berchem-Anvers (Belgium) London (U.K.) Ο. H. LOWRY G. QUADBECK Washington University, Max-Planck-Institut, St. Louis, Mo. (U.S.A.) Heidelberg (Germany) C. E. LUMSDEN J. H. QUASTEL Leeds University, McGill University, Leeds (U.K.) Montreal (Canada) F. LYNEN D. RICHTER University of Munich, Neuropsychiatrie Research Centre, Munich (Germany) Whitchurch Hospital, Cardiff (U.K.) J. MAGNES Hebrew University, R. ROSSITER Jerusalem (Israel) University of Western Ontario, London, Ontario (Canada) P. MANDEL University of Strassburg, F. O. SCHMITT Strassburg (France) Mass. Institute of Technology, Cambridge, Mass. (U.S.A.) N. MATUSSEK University of Munich, M. SCHNEIDER Munich (Germany) University of Cologne, Cologne (Germany) H. MCILWAIN Institute of Psychiatry, F. SCHONHEYDER London (U.K.) Aarhus University, Aarhus (Denmark) W. H. MCMENEMEY M aida Vale Hospital, M. SCHOU London (U.K.) Aarhus University and Risskov Mental Hospital, C. J. MUNCH-PETERSEN Risskov (Denmark) Aarhus University, Aarhus (Denmark) N. SCHÜMMELFEDER University of Bonn, S. NAKA Bonn (Germany) Osaka City University, Osaka (Japan) F. SEITELBERGER University of Vienna, Α. V. PALLADIN Vienna (Austria) Academy of Sciences of the Ukrainian S.S.R., V. S. SHAPOT Medical Institute, Kiev (U.S.S.R.) Vitebsk (U.S.S.R.) J. PATZIG I. C. SKOU Max-Planck-Institut, Aarhus University, Marburg (Germany) Aarhus (Denmark) E. PETTE University of Hamburg, G. H. SLOANE-STANLEY Hamburg (Germany) Institute of Psychiatry, London (U.K.) A. POPE Harvard Medical School and McLean G. SMTTS Hospital University of Amsterdam, Waverly, Mass. (U.S.A.) Amsterdam (Holland) J. POSTERNAK L. SOKOLOFF University of Geneva, National Institutes of Mental Health, Geneva (Switzerland) Bethesda, Md. (U.S.A.) xiv SECOND INTERNATIONAL NEUROCHEMICAL SYMPOSIUM W. M. SPERRY G. E. VLADIMIROV N.Y. State Psychiatric Institute and Pavlov Institute of Physiology, Columbia University, Academy of Science, New York (U.S.A.) Leningrad (U.S.S.R.) H. B. STEINBACH M. VOGT University of Copenhagen, University of Edinburgh, Copenhagen (Denmark) Edinburgh (U.K.) H. J. STRECKER G. VRAA-JENSEN N.Y. State Psychiatric Institute and Copenhagen Municipal Hospital, Columbia University, Copenhagen (Denmark) New York (U.S.A.) R. VRBA E. STRÖMGREN Dept. of Industrial Hygiene, Aarhus University and Risskov Mental Prague (Czechoslovakia) Hospital, Risskov (Denmark) H. WAELSCH N.Y. State Psychiatric Institute and L. SVENNERHOLM Columbia University, University of Gothenburg, New York (U.S.A.) Gothenburg (Sweden) J. D. WEILL R. H. S. THOMPSON University of Strassburg, Guy's Hospital, Strassburg (France) London (U.K.) H. WEIL-MALHERBE J. R. THUILLIER Runwell Hospital, University of Paris, Wickford (U.K.) Paris (France) G. B. WEST D. C. TOSTESON School of Pharmacy, University of Copenhagen, London (U.K.) Copenhagen (Denmark) G. WILKE S. UDENFRIEND Max-Planck-Institut, National Institutes of Health, Glessen (Germany) Bethesda, Md. (U.S.A.) S. YAMASAKI H. H. USSING Kyushu University, University of Copenhagen, Fukoaka (Japan) Copenhagen (Denmark) K. ZERAHN P. H. VERMEHREN Aarhus University, Copenhagen (Denmark) Aarhus (Denmark) SECTION 1 Molecular Structure and Morphology ELECTRON MICROSCOPY OF NERVOUS TISSUE H. FERNÂNDEZ-MORÂN Department of Nerve Ultrastructure, Instituto Venezolano de Neurologia, Caracas, Venezuela L Introduction II. Material and Methods 1. Fixation 2. Embedding Techniques 3. Ultrathin Sectioning Techniques 4. Dissociation and Replica Techniques III. Observations with the Electron Microscope 1. The Fine Structure of Neurons 2. The Synapses 3. Fine Structure of the Glial Cells IV. The Submicroscopic Organization of Nerve Fibers 1. The Fine Structure of the Myelin Sheath 2. Effects of Prolonged Ultracentrifugation on the Myelin Sheath 3. Experiments on the Application of Nuclear Magnetic Resonance Techniques V. Discussion I. INTRODUCTION THE INTRICATE nature of its known functions, and the great variety of the processes taking place in rapid yet co-ordinated sequence within the nervous system, all apparently require a morphological substrate more complex and more highly differentiated than the patterns of nervous tissue structure revealed hitherto by the light microscope. A detailed knowledge of the fine structure of nervous tissue extending to the molecular level is therefore becoming increasingly necessary. In particular, the phenomena studied by the neurophysiologists and neurochemists are inextricably linked to specific localization and highly ordered spatial arrangement in the submicroscopic domain. Much valuable information on the submicroscopic organization of fresh and even living tissue has already been derived from polarized light analysis (63-65) and X-ray diffraction studies (66-70, 25-30). But it is largely due to the successful application of the high resolving power of the electron 1

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