Visuomotor Coordination Amphibians, Comparisons, Models, and Robots Visuomotor Coordination Amphibians, Comparisons, Models, and Robots Edited by Jorg-Peter Ewert University of Kassel Kassel, Federal Republic of Germany and Michael A. Arbib University of Southern California Los Angeles, California Springer Science+Business Media, LLC Library of Congress Cataloging in Publication Data International Workshop on Visuomotor Coordination in Amphibians: Experiments, Comparisons, Models, and Robots (1987: Kassel, Germany) Visuomotor coordination: amphibians, comparisons, and models, and robots I edited by Jorg-Peter Ewert and Michael A. Arbib. p. em. "Proceedings of an International Workshop on Visuomotor Coordination in Amphi bians: Experiments, Comparisons, Models, and Robots, held August 25-27, 1987, in Kassel, Federal Republic of Germany"-T.p. verso. Bibliography: p. Includes index. ISBN 978-1-4899-0899-5 ISBN 978-1-4899-0897-1 (eBook) DOI 10.1007/978-1-4899-0897-1 1. Amphibians-Physiology-Congresses. 2. Vision-Congresses. 3. Sensorimotor integration-Congresses. I. Ewert, Jtirg-Peter, 1938- II. Arbib, Michael A. III. Title. QL669.2.159 1987 89-8467 597.6'041-dc20 CIP Proceedings of an International Workshop on Visuomotor Coordination in Amphibians: Experiments, Comparisons, Models, and Robots, held August 25-27, 1987, in Kassel, Federal Republic of Germany © 1989 Springer Science+Business Media New York Originally published by Plenum Press, New York in 1989 Softcover reprint of the hardcover 1st edition 1989 All rights reserved No part of this book may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, microfilming, recording, or otherwise, without written permission from the Publisher This book is dedicated to the memory of our friends Hans-Wilhelm Borchers, Rolando Lara, and Elena Sandoval killed in a car crash close to Cuernavaca in Mexico on January 19, 1985. We remember their significant contributions to the understanding of the amphibian visual system as we mourn their loss. A sampling of their last papers is included in the present volume. Contributors P. ARAIZA, Instituto de Fisiologia Celular, Universidad Nacional Aut6noma de Mexico, 04510 Mexico, D.F., Mexico MICHAEL A. ARBIB, Departments of Computer Science, Neurobiology, Physiology, Biomedical Engineering, Electrical Engineering, and Psychology, University of Southern California, Los Angeles, CA 90089-0782, USA RONALD C. ARKIN, Department of Information and Computer Science, Georgia Institute of Technology, Atlanta, Georgia 30332, USA ULRICH BAsSLER, Fachbereich Biologie, Universitat Kaiserslautern, D-6750 Kaiserslautern, FR Germany THOMAS W. BENEKE, Abteilung Neuroethologie, Fachbereich Biologie/Chemie, Universitat Kassel, D-3500 Kassel, FR Germany M. B. BERKINBLIT, Institute for Information Transmission Problems, Academy of Sciences and Moscow State University, Moscow 101447, USSR BILL BETTS, Program in Neural, Informational, and Behavioral Sciences, University of Southern California, Los Angeles, CA 90089-0782, USA HANS-WILHELM BORCHERS+, Abteilung Neuroethologie, Fachbereich Biologie/ Chemie, Universitat Kassel, D-3500 Kassel, FR Germany FRANCISCO CERVANTES-PEREZ, Departamento de Neurosciencias, Instituto de Fisiologia Celular, Universidad Nacional Aut6noma de Mexico, 04510 Mexico, D.F., Mexico THOMAS EGGERT, Max-Planck-Institut fiir Verhaltensphysiologie Seewiesen, D- 8130 Seewiesen/Stamberg, FR Germany JORG-PETER EWERT, Abteilung Neuroethologie, Fachbereich Biologie/Chemie, Universitat Kassel, D-3500 Kassel, FR Germany vii viii CONTRIBUTORS A. G. FELDMAN, Institute for Information Transmission Problems, Academy of Sciences and Moscow State University, Moscow 101447, USSR J. FERNANDEZ, Instituto de Fisiologia Celular, Universidad Nacional Aut6noma de Mexico, 04510 Mexico, D.F., Mexico THOMAS FINKENSTADT, Abteilung Neuroethologie, Fachbereich Biologie/ Chemie, Universitat Kassel, D-3500 Kassel, FR Germany EDDA M. FRAMING, Abteilung Neuroethologie, Fachbereich Biologie/Chemie, Universitat Kassel, D-3500 Kasse~ FR Germany 0. I. FUKSON, Institute for Information Transmission ·Problems, Academy of Sciences and Moscow State University, Moscow 101447, USSR FRANCISCO GONZALEZ-LIMA, Department of Anatomy, College of Medicine, Texas A&M University, College Station, TX 77843, USA PAUL GROBSTEIN, Department of Biology, Bryn Mawr College, Bryn Mawr, Pennsylvania 19010, USA EDWARD R. GRUBERG, Biology Department, Temple University, Philadelphia, PA 19122, USA UWE ANDER HEIDEN, Fakultat fiir Naturwissenschaften, Universitat Witten/ Herdecke, D-5810 Witten, FR Germany BARRY HORWITZ, Laboratory of Neurosciences, National Institute on Aging, National Institutes of Health, Bldg. 10, RM. 12S-207, Bethesda, MD 20892, USA REINHARD KOY-OBERTHOR, Institut fiir Elektrische lnformationstechnik, Tech nische Universitat Clausthal, D-3392 Clausthal-Zellerfeld, FR Germany PETER R. LAMING, Department of Biology, Queen's University of Belfast, Bel fast BT7 1NN, Northern Ireland, UK ROLANDO LARA+, Centro de Investigaciones en Fisiologia Celular, Universidad Nacional Aut6noma de Mexico, 04510 Mexico D.F., Mexico GYULA LAZAR, Department of Anatomy, University Medical School, Pees, H-7643, Hungary HANSPETERA. MALLOT, Institut fiir Zoologie lli (Biophysik), Johannes Guten berg-Universitat, D-6500 Mainz, FR Germany CONI'RIBUTORS ix CHRISTOPH VONDERMALSBURG, Department of Computer Science and Neuro biology, University of Southern California, Los Angeles, CA 90089-0782, USA GERHARD MANTEUFFEL, Fachbereich Biologie, Universitat Bremen, D-2800 Bremen, FR Germany L. MASSIEU, Instituto de Fisiologia Celular, Universidad Nacional Aut6noma de Mexico, 04510 Mexico, D.P., Mexico NOBUYOSHI MATSUMOTO, Department of Biophysical Engineering, Faculty of Engineering Science, Osaka University, Toyonaka, Osaka 560, Japan HORST MITIELSTAEDT, Max-Planck-Institut fiir Verhaltenspbysiologie See wiesen, D-8130 Seewiesen/Stamberg, FR Germany CHRISTOPH PINKWART, Holderlinstr. 10, D-8500 Niirnberg 20, FR Germany HERBERT J. REITBOECK, Angewandte Physik und Biophysik, Philipps-Universitat Marburg, D-3550 Marburg, FR Germany GERHARD ROTH, Fachbereich 2/Biologie, Universitat Bremen, D-2800 Bremen 33, FRGermany M. E. SANDOVAL+, Instituto de Fisiologia Celular, Universidad Nacional Aut6noma de Mexico, 04510 Mexico, D.F., Mexico EVELYN SCifORG-PFEIFFER, Abteilung Neuroethologie, Fachbereich Biologie/ Chemie, Universitat Kassel, D-3500 Kasse~ FR Germany WOLFGANG W. SCHWIPPERT, Abteilung Neuroethologie, Fachbereich Biologie/ Chemie, Universitat Kassel, D-3500 Kassel, FR Germany WERNER VON SEELEN, lnstitut fiir Zoologie ID (Biophysik), Johannes Gutenberg Universitat, D-6500 Mainz, FR Germany CARME TORRAS, Institut de Cibem~tica (CSIC-UPC), Diagonal647, 08028- Barcelona, Spain ANANDA WEERASURIYA, Department of Physiology, Faculty of Medicine, Uni versity of Colombo, Colombo-8, Sri Lanka Preface Various brain areas of mammals can phyletically be traced back to homologous structures in amphibians. The amphibian brain may thus be regarded as a kind of "microcosm" of the highly complex primate brain, as far as certain homologous structures, sensory functions, and assigned ballistic (pre-planned and pre-pro grammed) motor and behavioral processes are concerned. A variety of fundamental operations that underlie perception, cognition, sensorimotor transformation and its modulation appear to proceed in primate's brain in a way understandable in terms of basic principles which can be investigated more easily by experiments in amphibians. We have learned that progress in the quantitative description and evaluation of these principles can be obtained with guidance from theory. Modeling - supported by simulation - is a process of transforming abstract theory derived from data into testable structures. Where empirical data are lacking or are difficult to obtain because of structural constraints, the modeler makes assumptions and approximations that, by themselves, are a source of hypotheses. If a neural model is then tied to empirical data, it can be used to predict results and hence again to become subject to experimental tests whose resulting data in tum will lead to further improvements of the model. By means of our present models of visuomotor coordination and its modulation by state-dependent inputs, we are just beginning to simulate and analyze how external information is represented within different brain structures and how these structures use these operations to control adaptive behavior. Vzsuomotor Coordination: Amphibians, Comparisons, Models, and Robots presents for a larger audience the proceedings of the third international workshop on this topic, organized by Peter Ewert at the University of Kassel in August '15-27, 1987. The interdisciplinary workshops on visuomotor coordination in frogs and toads have a successful tradition. The first meeting was organized by Michael Arbib at the University of Massachusetts at Amherst in 1981 and the second by Rolando Lara at the University of Mexico in 1982. Meanwhile a wealth of new experimental data on visuomotor functions in target-oriented behaviors of amphibians - in terms of ethology, physiology, anatomy, and pharmacology - has accumulated. Our third international workshop proceeds in the tradition of furthering the dialog between experimentalists and theorists. Because there are no boundaries between artificial and natural sensorimotor systems in cybernetics, cross-disciplinary studies are particularly fruitful. Implicit in xi xii PREFACE the goals of our interdisciplinary research is another branch touching on interests of applied science. The study of biological information processing systems holds promise of indicating elegant solutions to practical problems in robots. Natural pattern recognition systems, for example, successfully deal with different invariance conditions in visual sensorimotor tasks. Comparable properties have proven difficult to implement in optoelectronic devices or machines (e.g., position/orientation/velocity invariance in the recognition of work-pieces on assembly lines). Computer simulations represent operating algorithms derived from the basic properties of the model which can be implemented in robotic systems. Hence, with the aid of models constructed from data on biological mechanisms of visuomotor coordination a first step in the development and the testing of algorithms for performance and tuning of robotic devices may be taken. Moreover, the interplay of empirical and theoretical analyses will actually increase the pace toward our understanding of the properties of information processing carried out by central nervous systems. PART I opens the workshop proceedings with a general discussion on experimentation and modeling in neuroscience, transcribed from tape recordings at the meeting. Main topics of general interest concern the dialog between model and experiment, the structure/function problem, the modulation of sensorimotor function, and methodological interpretational problems. PART II offers two overviews of the theme of this volume. The neuroethological approach (Ewert) develops the neurophysiological equivalent of the ethological Lorenzian concept of an "innate releasing schema" (also called releasing mechanism after Tinbergen) as a "sensorimotor code" constituted by different combinations of neurons, each of these expressing information processing in a functional unit of interacting cells and specified to encode significant coincidences of stimulus cues that refer to an object, e.g. a visual releaser. The corresponding command releasing systems result from parallel distributed processing in a neural macro-network that evaluates visual input under various aspects, such as features, background, space, and past experience. The systems theoretical treatment (Arbib) uses perceptual and motor schemas to provide models of amphibian visuomotor coordination bridging behavior and neural circuits, and allowing approaches to perceptual robotics, both in the control of dexterous robot hands and the design of machine vision systems. PART m reports new results on the cellular architecture and connectivity of the anuran's optic tectum and pretectum (Lazar) and the response properties of labeled neurons (Matsumoto). We are faced here with an enormous richness in cytological features of local circuits and of connecting and projecting neurons whose bifurcating axons might provide a substrate for parallel processing. Response properties raise the question of isomorphisms between the geometry of dendritic trees, axonal projections, and the function of neurons. Application of the pulse-triggered averaging technique provides insight into retino-tectal synaptic connectivity. Biochemical investigations are becoming of increasing importance to understand interneuronal communication and its control. Using isolated nerve endings or brain slices from the optic tectum, various neurochemical criteria - sodium dependent uptake and calcium dependent release - are applied to identify amino acid and amine neurotransmitters (Sandoval, Massieu, Araiza & Fernandez). A model of retino-tectal information
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