Advances in Vertebrate Neuroethology PDF
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Advances in Vertebrate Neuroethology NATO Advanced Science Institutes Series A series of edited volumes comprising multifaceted studies of contemporary scientific issues by some of the best scientific minds in the world, assembled in cooperation with NA TO Scientific Affairs Division. This series is published by an international board of publishers in conjunction with NATO Scientific Affairs Division A Life Sciences Plenum Publishing Corporation B Physics New York and London C Mathematical and D. Reidel Publishing Company Physical Sciences Dordrecht, Boston, and London 0 Behavioral and Martinus Nijhoff Publishers Social Sciences The Hague, London, and Boston E Applied Sciences F Computer and Springer Verlag Systems Sciences Heidelberg G Ecological Sciences Recent Volumes in Series A: Life Sciences Volume 51-Factors in Formation and Regression of the Atherosclerotic Plaque edited by Gustav R. V. Born, Alberico L. Catapano, and Rodolfo Paoletti Volume 52-Chemical Carcinogenesis edited by Claudio Nicolini Volume 53-Cancer of the Prostate and Kidney edited by M. Pavone-Macaluso and P. H. Smith Volume 54-Leukotrienes and Prostacyclin edited by F. Berti, G. Falco, and G. Vela Volume 55-Durable Resistance in Crops edited by F. Lamberti, J. M. Waller, and N. A. Van der Graaff Volume 56-Advances in Vertebrate Neuroethology edited by Jorg-Peter Ewert, Robert R. Capranica, and David J. Ingle Volume 57-Biochemical and Biological Markers of Neoplastic Transformation edited by Prakash P. Chandra Volume 58-Arterial Pollution: An Integrated View on Atherosclerosis edited by H. Peeters, G. A. Gresham, and R. Paoletti Advances in Vertebrate Neuroethology Edited by Jorg-Peter Ewert University of Kassel Kassel, Federal Republic of Germany Robert R. Capranica Cornell University Ithaca, New York and David J. Ingle Brandeis University Waltham, Massachusetts Plenum Press New York and London Published in cooperation with NATO Scientific Affairs Division Proceedings of a NATO Advanced Study Institute on Advances in Vertebrate Neuroethology, held August 13-24,1981, at the University of Kassel, Kassel, Federal Republic of Germany Library of Congress Cataloging in Publication Data NATO Advanced Study Institute on Advances in Vertebrate Neuroethology (1981: Univer sity of Kassel) Advances in vertebrate neuroethology. (NATO ASI series. Series A, Life sciences; v. 56) "Proceedings of a NATO Advanced Study Institute on Advances in Vertebrate Neuro ethology, held August 13 - 24, 1981, at the University of Kassel, Kassel, Federal Republic of Germany"-T.p. verso. "Published in cooperation with NATO Scientific Affairs Division." Bibliography: p. Includes index. 1. Vertebrates-Behavior-Congresses. 2. Vertebrates-Physiology-Congresses. 3. Neurobiology-Congresses. I. Ewert, Jorg-Peter, 1938 - . II. Capranica, Robert R. III. Ingle, David. IV. North Atlantic Treaty Organization. Scientific Affairs Division. V. Title. VI. Series. [DNLM: 1. Behavior, Animal-Physiology-Congresses. 2. Neurophysiology Congresses. 3. Vertebrates-Physiology-Congresses. QL 751 N279a 1981] QL750.N37 1981 596'.0188 82-22354 ISBN 978-1-4684-4414-8 ISBN 978-1-4684-4412-4 (eBook) DOI 10.1007/978-1-4684-4412-4 © 1983 Plenum Press, New York A Division of Plenum Publishing Corporation 233 Spring Street, New York, N.Y. 10013 Softcover reprint of the hardcover 1s t edition 1983 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 Dedicated to Professor Dr. Dr. h.c. Hansjochem Autrum , a pioneer in the field of Comparative Sensory Physiology, on the occasion of his 75th birthday ACKNOWLEDGMENTS This symposium was sponsored by various organizations, national and international. The Organizing Committee would like to thank them and their representatives: - North Atlantic Treaty Organization, N.A.T.O. Advanced Study Institutes Programme - Bayer Aktiengesellschaft, Leverkusen, F.R. of Germany - Behringwerke Aktiengesellschaft, Marburg, F.R. of Germany - Der Hessische Kultusminister, Wiesbaden, F.R. of Germany - Deutscher Akademischer Austauschdienst DAAD, F.R. of Germany - Deutsche Stiftung fUr Internationale Entwicklung DSE, F.R. of Germany - Hoechst Aktiengesellschaft, Frankfurt a.M., F.R. of Germany - National Science Foundation NSF, U.S.A. - Universitat des Landes Hessen, Gesamthochschule Kassel, F.R. of Germany PREFACE This volume presents the proceedings of the NATO Advanced Study Institute on "Advances in Vertebrate Neuroethology" held at the University of Kassel, Federal Republic of Germany in August 1981. During the last decade much progress has been made in understanding the neurophysiological bases of behavior in both vertebrates and invertebrates. The reason for this is that a number of new physiological, anatomical, and histochemical techniques have recently been developed for brain research which can now be combined with ethological methods for the analysis of animal behavior to form a new field of research known as "Neuroethology". The term Neuroethology was originally introduced by S.L.Brown and R.W.Hunsperger (1963) in connection with studies on the activation of agonistic behaviors by electrical brain stimulation in cats. Neuroethology was more closely defined by G.Hoyle (1970) in the context of a review on cellular mechanisms underlying behavior of invertebrates. Since the 6th annual meeting of the Society for Neuroscience held in Toronto in 1976, Neuroethology has become established as a session topic. The NATO Advanced Study Institute "Advances in Vertebrate Neuroethology" was the first large international meeting of neuroethologists working on vertebrates. The Study Institute was attended by 80 participants from 11 NATO member countries and from Poland, Peoples Republic of China, Sri Lanka, and Switzerland. The development of Neuroethology is indeed truly internationaJ with some specialization in different countries. In the U.S.A. and Canada, there is an emphasis on mammals and a stronger tradition of neuroanatomy and neurochemistry. In Europe the majority of workers come from a zoological-ethological tradition and have developed quantitative ethological, single cell recording, and electrical brain stimulation techniques. Thus, exchange of methods and ideas for research was one benefit especially for younger scientists attending this meeting in deciding how to develop their own work. One of the special features of Neuroethology is that it acts as a catalyst in promoting interdisciplinary research among scientists from different fields, such as zoology, medicine, psychology, communication sciences, and engineering. ix x PREFACE Neuroetho1ogy draws its questions from the biology of behavior and strives to explore principles of neurobiological organization in animals with different ecological and behavioral adaptations. Hence, the comparative approach contributes toward the understanding of appropriate processes at the highest plane of integration in the vertebrate nervous system. Some of these questions are: (1) Which processes in sensory systems are responsible for the distinction between behaviorally important and unimportant stimuli, i.e., how are signals recognized? (2) What are the neuronal mechanisms of localization? (3) What are the stimu~us means of acqulrlng, storing, and recalling information? (4) What are the bases for motivation of behavior? (5) How are sensory decisions performed and transformed into efferent motor commands? (6) How is a motor pattern coordinated and controlled? (7) How are ontogenetic developments related to neuronal mechanisms? In pursuing these questions, Invertebrate Neuroetho1ogy and Vertebrate Neuroetho1ogy tend to use different research strategies. Whereas the first approaches the system properties of neuronal networks controlling behavior primarily from the motor side, the latter has focussed its main interest on sensory and motivational systems. Thus, four research lines in Vertebrate Neuroetho1ogy have shown the greatest development: sensory mechanisms subserving localization and identification of behaviorally relevant signals, intraspecific communication, and central control of motivation. These were the session topics of the Study Institute. They were outlined by 23 introductory lectures and accompanied by 30 short seminar papers. Special lectures dealt with the various implications for Vertebrate Neuroethology from the study areas: ethology, neurophysiology, neuroanatomy, and insect neuroethology. Specific topics were treated in depth by panel discussions: (i) sensory processing of key stimuli, (ii) sensorimotor interfacing, and (iii) state dependent responses. Tutorial sessions gave the opportunity to acquaint participants with modern research techniques relevant to Neuroethology, such as telemetric brain stimulation, telemetric single cell recording, film analysis, and neuroanatomical techniques. Hence, this conference presented a spectrum of methods ranging from a straight-forward quantitative analysis of sensory response mechanisms to the new eclecticism of techniques and experimental design for studies of motivation and individual experience. Historically, the search for the neural bases of behavior goes back a long way. Neuroethology did not emerge from a certain school, but from an eclectic confluence of many streams rooted in a wide assortment of traditions, such as ethology - influenced, e.g., by Craig, Heinroth, James, Lorenz, MacDougall, Pavlov, Skinner, Thorndike, Tinbergen, v.UexkUll ,comparative physiology influenced, e.g., by Autrum, Bullock, Doty, Galambos, Gray, Griffin, Hess, v.Holst, Huber, Jordan, Lettvin, Lissmann, Loeb, Magoun, Pringle, Prosser, Roeder, Sechenov, Segaar, Sherrington, PREFACE xi Sperry, Weiss -, comparative neuroanatomy - influenced, e.g., by Caja1, Edinger, Glees, Go1gi, Herrick, Nauta, Retzius -, as well as clinical neurology - influenced, e.g., by Bartholow, Ferrier, Fritsch, Hitzig, Jackson, Lashley, Nordenskio1d, Teuber. The editors dedicate this book to Hansjochem Autrum, a pioneer in the field of comparative physiology, on the occasion of his 75th birthday. Fifty years ago, when Hansjochem Autrum tackled the problem of receptor sensitivity by means of e1ectrophysio10gica1 measurements, he had to build his own amplifiers and even a suitable oscilloscope. At that time there were no commercial DC amplifiers available. Upon Autrum's advice, his friends physicists of the "Forschungsansta1t der Reichspost" - developed within 9 months the first DC-amplifier which was battery powered by a 200V unit consisting of many small Ni-Fe batteries each of them yielding just 1.5V. In 1952 Hansjochem Autrum began an investigation of photoreceptor properties and, therefore, he applied to the Deutsche Forschungsgemeinschaft for an "optical bench". The referees by a hair's breadth almost rejected his application since they could not imagine that this kind of equipment can be used in a zoological department. Today almost everybody knows that experimental biology and technology are bound together in a kind of dualistic manner. The most recent past has shown how progress within a field of research, whose goal is to explore the neural processes underlying behavior, goes hand in hand with technical developments, especially in the field of electronics and chemical tracers. I wish to thank the co-directors of the NATO Advanced Study Institute, Richard J. Andrew, Robert R. Capranica, and David J. Ingle, for their constructive help in the development of the scientific program, and my staff members Hans-Wilhelm Borchers, Harald Burghagen, Evelyn SchUrg-Pfeiffer, and Ananda Weerasuriya for their assistence in the local organization of the meeting. I am most grateful to Ursula Reichert, Inge Samm1er, and Christa Uthof for type setting of the camera-ready manuscripts with the aid of our word processor and for their help in the lay-out. The willing collaboration of all participants has contributed greatly toward a successful meeting and a fast publication of its proceedings. During the last plenary session of the Study Institute an "International Society for Neuroetho10gy" was founded. We are looking forward to seeing the development of this young Society, and I wish our new chairman Mark Konishi good luck for the organization of the next Symposium. Summer, 1982 Jorg-Peter Ewert Kassel Federal Republic of Germany CONTENTS Sensory processing of key stimuli ....................... 3 R.R. Capranica Sensorimotor interfacing 7 H. Scheich State dependent responses 15 R.J. Andrew SECTION I - IMPLICATIONS FOR NEUROETHOLOGY Some ethological implications for neuroetho1ogy: The ontogeny of birdsong ................................ 21 P. Marler Implications for neuroetho1ogy from comparative neurophys i 01 ogy ......................................... 53 T. H• Bu 11 oc k Neuroanatomica1 implications for neurot 1010gy .......... 77 S.O.E. Ebbesson Implications of insect neuroetho1ogy f studies on vertebrates ..................... . 91 F. Huber xiii
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