Brain and Human Behavior

Transcription

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2 Brain and Human Behavior Edited by A. G. Karczmar J. C. Eccles With 162 Figures Springer-Verlag Berlin Heidelberg. New York 1972

3 ALEXANDER G. KARCZMAR, Professor of Pharmacology, Department of Pharmacology and Institute for Mind, Drugs and Behavior, Loyola University of Chicago, Stritch School of Medicine, 2160 South First Ave., Maywood, IL 60153/USA Professor Sir JOHN C. ECCLES, Laboratory of Neurobiology, Department of Physiology, State University of New York at Buffalo, 4234 Ridge Lea Road, Amherst, NY 14226/USA ISBN -13 : e-isbn-13 : DOl: / This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks. Under 54 of the German Copyright Law where copies are made for other than private use, a fee is payable to the publisher, the amount of the fee to be determined by agreement with the publisher. by Springer-Verlag Berlin Heidelberg Library of Congress Catalog Card Number Softcover reprint of the hardcover 1st edition 1972 The use of general descriptive names, trade names, trade marks, etc. in this publication, even if the former are not especially identified, is not be taken as a sign that such names, as understood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Typesetting, printing and binding: Konrad Triltsch, Graphi.cher Betrieb, 87 Wiirzburg, Germany.

4 Foreword This volume is based on the Symposium on "The Brain and Human Behavior," held in October of 1969 as a part of the centennial observance of the Loyola University of Chicago. As President of the University, I was pleased to offer the University's support for the organization of this Symposium and to participate in some of its sessions. The volume which I now have the pleasure to introduce employs the materials of the Symposium as a framework. Its chapters constitute updated and greatly expanded versions of the original presentations, edited and organized so as to constitute an integrated picture of Neurosciences and their epistemological aspects. It seems appropriate for me to describe at this time certain features of this Jesuit University and of its Centennial which are particularly pertinent in the context of the present volume. Loyola University of Chicago opened its classes on September 5, 1870 with a faculty of 4 and a student body of 37. Today, Loyola University is the largest independent University in Illinois and the largest institution of higher learning under Catholic sponsorship in the United States of America. The University comprises twelve schools and colleges, a faculty of more than 1,600 and a student body of 16,545. As an institution of learning, this University is dedicated to knowledge; but perhaps more particularly than others, it is dedicated to the integration of truth and the knowledge of man as such. It is not man simply in his various and single aspectsthe man of technology, the man of organization, or the man functional in terms of his physiological and anatomical units, but rather the whole man who interests us, whom we study and whom we try to the present as an entity to our students. Additionally, as a complex modern urban University, Loyola serves the vast metropolitan Chicago area in a wide variety of ways while, conscious of its particular dedication to the service of man, it strives to put knowledge at the service of the national and international community of scholars. Appropriately then, the theme selected by the faculty for Loyola's centennial observance was "Knowledge in the Service of Man." The concept of "Knowledge in the Service of Man" is, of course, central to the whole history of man's progress. It is knowledge and learning which constitute the basis of the special destiny and progress of mankind. It is knowledge that has provided us with the many benefits that modern man enjoys but regrettably knowledge has also been the basis of many cruel disservices to mankind. The present volume is particularly appropriate for our centennial observance. The knowledge and the learning which, as I stated, so greatly affect man's destiny, are intimately associated with the brain and its functions. It is the special attribute of the human brain to allow us to learn and to accumulate, remember and utilize knowledge.

5 VI Foreword Due to the brain's integrative capacity, the individual can act as a total entity in his functions. And a particular attribute of the brain's activity is that, not only is it outer-directed as man attempts to understand the world and adapt himself to his environment, but it is inner-directed as well, as man, through introspection, endeavors to visualize his fate and his real meaning within this world. The contributors to this volume are among the world's foremost experts in the various phases of brain function and of its correlation to human behavior, whether in the overt or introspective sense. In this volume are crossed the wide ranges of the interdisciplinary areas which are necessary to understand the brain and human behavior as a whole. Just as man is a totality of all his aspects, so is the brain an integrated totality of its neurons. Consequently, the volume ranges widely from the description of the functions of the single neurons which, across a few micra of space, encompass endless complexities of structure and function (leading one of the participants to designate them "the little brains") to the description of the neuronal systems, of the memory, sleep, attention, and perception and, on a still higher level of complexity, to the problems of learning and instinct. The last section of the volume is of particular interest. There, the precise and empirical knowledge described in the preceding sections is the subject of a transcendental scrutiny. The meaning of man isofar as he is intelligible in terms of today's knowledge of his brain and behavior and the special significance of his mind in terms of the problems of choice and of will are given emphasis in this section. This volume then bridges the biological, sociological and philosophical aspects of man, brain and behavior. It expresses well the particular dedication and attitude of this University and it serves as an important summary of the present state of Neurosciences. For this reason, besides constituting for the Loyola University of Chicago community a memento of the centennial observance, the volume should appeal to the widest forum of serious readers. Chicago, 14th September 1971 JAMES F. MAGUIRE, S. J. Chancellor Loyola University of Chicago

6 Preface At the outset I would stress the transcendent importance of the scientific study of the brain to which this volume is dedicated. By the use of traditional methods of science we have come to recognize that it is the most complexly organized structure in the natural world. At the simplest level of analysis we can consider the brain as being composed of many thousands of millions of neurones or nerve cells that communicate with each other by impulse discharges and by transmission across highly specialized chemical synapses. The main transmitting lines of neuronal communication are now well defined and there is much detailed knowledge of the synaptic mechanisms in some simple standard types of circuit that have been subjected to systematic analysis. However, in the case of the cerebral cortex we have as yet only fleeting glimpses of neuronal operation. One of the most challenging problems dealt with in this volume concerns the manner in which memory storage is brought about, and the manner of its retrieval. Converging on these problems there are already the theories and experiments of a wide range of neurobiologists in such a variety of disciplines as electronmicroscopy, neurochemistry, neuropharmacology, behavioral science, neurophysiology, neurogenesis and neurocommunications. I regard the progress as so encouraging that we may hope for a good understanding in the near future, though not of course a complete solution. At one level of comprehension and investigation we can regard the brain as a neuronal machine of almost infinite complexity. Even from our present fragmentary knowledge we can appreciate how it can give us the immense variety and subtlety of performance throughout life, and develop the immense range of storage and retrieval that we experience in memory. However, there is another aspect of brain function that is beyond any scientific comprehension at the present time, and that is its relationship to conscious experience. This fundamental problem lies rather marginal to the scope of a volume on brain and behavior, yet it has been referred to in many papers, particularly those concerned with philosophical problems. I agree with the philosophers at the symposium that conscious experiences are not reducible to physics, and that we do not yet have any satisfactory approach to the brain-mind problem. This problem seems to become ever more intractable the more we advance in the scientific study of the brain. Dr. KARCZMAR has prepared a remarkable introduction to this important symposium, but it is much more than a summary. It is a critical appraisal and an attempt at synthesis of the diverse contributions. Buffalo, 4th October 1971 JOHN C. ECCLES

7 Acknowledgment The Editors gratefully acknowledge the financial support of Loyola University of Chicago which made the Symposium on "Brain and Human Behavior" possible. Most of the contributions to the present volume while related to or based on, the symposial presentations, were subsequently expanded, updated and edited to meet the aims of this particular book; some of the contributions were especially prepared for this volume.

8 Contents Introduction. What we know now, will know in the future, and possibly cannot ever know in neurosciences: A. G. KARCZMAR 1 I. Molecular and Synaptic Organization Molecular Organization of Synapses for Chemical Transmission in the Central Nervous System: E. DE ROBERTIS. With 7 Figures. 22 Discussion. 35 Possible Synaptic Mechanisms Subserving Learning: J. C. ECCLES. With 12 Figures. 39 Synaptic Modulations: A. G. KARCZMAR, S. NISHI and L. C. BLABER. With 9 Figures 63 II. Biochemical Mechanisms and Pharmacological Approaches Some Brain Protein Changes Reflecting Neuronal Plasticity at Learning: H. HYDEN. With 8 Figures. 94 Discussion. 110 Norepinephrine in Central Nervous System and its Correlations with Behavior: S. S. KETY. With 1 Figure. 115 Discussion. 128 Some Monoaminergic Mechanisms Controlling Sleep and Waking: M. jouvet. With 15 Figures. 131 Discussion. 161 Ill. Neurophysiological Correlates Data Structure and Algorithms in the Primate Somatosensory Cortex: G. WERNER, B. L. WHITSEL and L. M. PETRUCELLI. With 16 Figures 164 Cortical Visual Areas and Their Interaction: M. MISHKIN. With 18 Figures 187 Neurophysiological and Psychophysical Correlates in Vision Research: R. JUNG. With 13 Figures. 209 Integrative Properties of Parastriate Neurons: F. MORRELL. With 12 Figures 259 Neural Correlates of Learning and Memory: E. R. JOHN. With 7Rgu~ ~1 Electrophysiological Correlates of Positive Reinforcement: Post Reinforcement Synchronization, Modulation of Sensory Input, and Steady Potentials: T. j. MARCZYNSKI. With 7 Figures. 303

9 x Contents IV. Psychological Aspects Nature, Nurture and the Psychological Approach to Learning: D. BOVET. With 9 Figures Phylogeny of Memory Development in Vertebrates: I. S. BERITASH- VILI. With 5 Figures Discussion Time, Space, Motivation, Memory, and Decision: W. T. LIBERSON. With 23 Figures The Ontogeny of Behavior and the Concept ofinstinct: S.A.BARNETT 377 Operational Structures of the Intelligence and Organic Controls: J. PIAGET V. Epistemological Aspects In Defense of Teleology: R. GRANIT The Mentality of Man's Brain: S. TOULMIN 409 What Difference Does Mind Make?: E. McMuLLIN. 423 Conditions for a Mechanistic Theory of Behaviour: CH. TAYLOR 449 Index 467

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