Jorg-Peter Ewert Neuroethology An Introduction to the Neurophysiological Fundamentals of Behavior Translation: Transemantics, Inc. With 171 Figures, Most in Color Springer-Verlag Berlin Heidelberg New York 1980
Professor Dr. rer. nat. J6rg-Peter Ewert Arbeitsgruppe Neuroethologie, FB 19, GHK Universitat des Landes Hessen Heinrich-Plett-Str. 40, D-3500 Kassel, FRG Translation: Transemantics, Inc., Suite 407 1901 Pennsylvania Avenue N.W Washington, D.C. 20006, USA Title of the German edition J6rg-Peter Ewert, Neuro-Ethologie Springer-Verlag Berlin Heidelberg New York 1976 [SBN-I3: 978-3-642-67502-7 e-[sbn-13: 978-3-642-67500-3 00[: [0. [007/978-3-642-67500-3 Library of Congress Cataloging in Publication Data. Ewert, Jorg-Peter, 1938-. Neuroethology. Translation ofneuro-ethologie. Bibliography: p. Includes index. 1. Neuropsychology. 2. Animals, Habits and behavior of. I. Title. QP360.E9313 156'.27923144. 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 1980 Softcover reprint of the hardcover 1 st edition 1980 The use ofregistered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Typesetting: Beltz, Offsetdruck, Hemsbach/Bergstr. 213113130-5432lO
Preface Historically the search for the neural bases of behavior goes back a long way. Neuroethology, which is concerned with the experimental analysis ofthe releasing and control mechanisms of behavior, is a young discipline. Results from this multidisciplinary branch of research, which uses physical, chemical, and mathematical methods, have not yet been extensively treated in textbooks of neurophysiology and ethology. This book is intended as a first attempt to pose major questions of neuroethology and to demonstrate, by means of selected research examples, some of the ways by which these questions are being approached. Inevitably this cannot be a complete and in depth detailed treatment of all of the neurobiology examples, and I realize that such a selection is of a subjective nature. The overall goal ofthe book is to present an introduction. After outlining some of the very basic neurophysiological and ethological concepts (Chaps. 2 and 3), neuroethological questions and methods are demonstrated extensively by means of a particular example (Chap. 4). There are two reasons to choose the visually guided prey-catching and avoidance behavior of the Common Toad: (1) it is a system which I have investigated for about fifteen years and therefore know best, (2) the toad story is one of the most comprehensive neuroethological approaches so far. Thus, it is possible here to outline the major concepts of neuroethology and to pose the basic questions. Chapters 5-8 show that the same questions and concepts are also relevant to other animals and to other systems. Not all of the examples are sufficiently developed because of space limitations. For a more detailed account of some of these examples, references are provided in the figure captions, and reading lists will be found at the end of the book (cf. Special References and Suggested Reading). Numerous, in part, strongly schematized two-color diagrams illustrate the text. Essential summarizing statements are printed on a colored background. The reader can be quickly oriented by reading these paragraphs and studying the illustrations. Information about current neurobiological methods is condensed in an ap-
VI Preface pendix (Chap. 9). The English edition Neuroethology is a translation of the German edition Neuro-Ethologie published in 1976. The English version is more comprehensive, brought up to date and provided in some places with new illustrations. I wish to express my appreciation to Transemantics, Inc. for translating this book. I want to express special thanks to Howard Carl Gerhardt (University of Missouri, Columbia) and Ananda Weerasuriya (National Institutes of Health, Bethesda, Maryland) for careful revision and detailed comments on the manuscript from which I have benefited greatly. I am most grateful to the publisher, Dr. Konrad F. Springer, for the generosity shown in the production of the book. Kassel, Spring 1980 JORG-PETER EWERT
Contents 1 What is Neuroethology? 1.1 Principal Scope 1.2 History 1.3 Problems.. 1 3 11 2 How Does a Stimulus Elicit Muscle Contraction? 2.1 Neural Building Blocks..... 13 2.2 A Behavioral Experiment....... 27 2.3 The "Language" of the Nervous System.. 33 2.4 Communication in the Nervous System.. 37 2.5 The Three Fundamental Arithmetic Methods 41 2.6 Complexity........... 46 2.7 Nerve Impulses Induce Muscle Contractions 51 3 Key-Stimuli and Releasing Mechanisms: Some Fundamental Concepts in Ethology 3.1 Environmental Stimuli and Information Reduction 57 3.2 Innate and Acquired Releasing Mechanisms 58 3.3 Releasing Stimuli 58 3.4 Principles of Stimulus Selection 65 3.5 Motivation and Change of Meaning 67 4 Neurobiological Basis for the Recognition and Localization of Environmental Signals: How Does a Toad Brain Recognize Prey and Enemy? 4.1 The Key-Stimuli "Prey" and "Enemy". 71 4.2 Prey-Catching and Avoidance "Areas" in the Toad Brain 92 4.3 Are There "Prey" and "Enemy Neurons" in the Toad Brain? 99 4.4 A Working Hypothesis on Neuronal Decision- Making 115 4.5 Relevance of the Supposed Gestalt Recognition System 121
VIII Contents 5 Functional Areas in the Visual System of Vertebrates: Comparative Aspects 5.1 Substrates for Stimulus Localization 5.2 Space Constancy and Optokinetic Nystagmus 5.3 Brain Structures for Stimulus Identification. 5.4 Formation and Deterioration of the Visual System: Environmental Effects 6 Examples for Release and Control of Behavior Patterns by Other Sensory Systems 6.1 Olfactory Sense: Scent Coding in Insects 6.2 Vibration Sense: Prey-Catching ofthe Back Swimmer. 6.3 Acoustic Communication in Crickets and Frogs 6.4 Active Exploration of the Environment: Echo- and Electrolocation. 6.5 Attenuation of Self-Stimulation 7 Neuronal Circuits for Fixed Motor Behavior Programs 7.1 Example of a Neuronal Program Circuitry 7.2 Command Neurons 7.3 Command Functions in the Primate Brain 8 Central Representation of Behavior Motivation 8.1 Food Intake: Hunger and Thirst. 8.2 Sexual Behavior 8.3 Aggression: Attack and Defense. 8.4 Learning: Storing, Recalling, Reinforcing 8.5 Social Stress 9 Methodological Appendix 9.1 Prerequisites for Behavior Analysis 9.2 Investigation of Behaviorally Relevant Brain Areas 9.3 Methods for Recording and Measuring Neuronal Activity 9.4 Investigation of Anatomical Connections in the Brain 9.5 Statistical and Systems Theoretical Analysis Special References Suggested Reading Subject Index.. 129 134 135 141 147 153 158 174 199 203 208 211 219 227 235 239 249 256 259 266 284 291 297 311 327