B. Kramer Electro- communication in Teleost Fishes Behavior and Experiments Springer-Verlag Zoophysiology Volume 29 Editors S. D. Bradshaw W. Burggren H.C. Heller S.Ishii H.Langer G. Neuweiler D. J. Randall Zoophysiology Volumes already published in the series: Volume 1: P.J.Bentley Volume 16: P. Bouverot Endocrines and Osmoregulation Adaption to Altitude-Hypoxia (1971) in Vertebrates (1985) Volume 2: L. Irving Volume 17: R.J.F. Smith Arctic Life of Birds and Mammals The Control of Fish Migration (1985) Including Man (1972) Volume 18: E. Gwinner Volume 3: A.E. Needham Circannual Rhythms (1986) The Significance of Zoochromes (1974) Volume 19: J.C. Rtiegg Volume 4/5: A. C. Neville Calcium in Muscle Activation (1986) Biology of the Arthropod Cuticle Volume 20: J.-R. Truchot (1975) Comparative Aspects of Extracellular Volume 6: K. Schmidt-Koenig Acid-Base Balance (1987) Migration and Homing in Animals Volume 21: A. Epple and J.E. Br inn (1975) The Comparative Physiology of the Volume 7: E. Curio Pancreatic Islets (1987) The Ethology of Predation (1976) Volume 22: W.H. Dantzler Volume 8: W. Leuthold Comparative Physiology of the African Ungulates (1977) Vertebrate Kidney (1988) Volume 9: E. B. Edney Volume 23: G.L. Kooyman Water Balance in Land Arthropods Diverse Divers (1989) (1977) Volume 24: S. S. Guraya Volume 10: H.-U. Thiele Ovarian Follicles in Reptiles and Carabid Beetles in Their Environments Birds (1989) (1977) Volume 25: G.D. Pollak and Volume 11: M.H.A. Keenleyside J.H. Casseday Diversity and Adaptation in The Neural Basis of Echolocation in Fish Behaviour (1979) Bats (1989) Volume 12: E. Skadhauge Volume 26: G.A. Manley Osmoregulation in Birds (1981) Peripheral Hearing Mechanisms in Reptiles and Birds (1989) Volume 13: S. Nilsson Autonomic Nerve Function in the Volume 27: U. M. Norberg Vertebrates (1983) Vertebrate Flight (1990) Volume 14: A.D. Hasler Volume 28: M. Nikinmaa Olfactory Imprinting and Homing Vertebrate Red Blood Cells (1990) in Salmon (1983) Volume 29: B. Kramer Volume 15: T. Mann Electrocommunication in Teleost Spermatophores (1984) Fishes (1990) B. Kramer Electrocommunication in Teleost Fishes Behavior and Experiments With 140 Figures Springer-Verlag Berlin Heidelberg NewYork London Paris Tokyo Hong Kong Barcelona Professor Dr. BERND KRAMER Zoological Institute of the University 8400 Regensburg, FRG ISBN 3-540-51927-0 Springer-Verlag Berlin Heidelberg New York ISBN 0-387-51927-0 Springer-Verlag New York Berlin Heidelberg Library of Congress Cataloging-in-Publication Data. Kramer, Bernd. Electrocom- munication in teleost fishes: behavior and experiments/B. Kramer, p. cm. - (Zoo- physiol; v. 29) Includes bibliographical references and indexes. ISBN 3-540-51927-0 (Springer-Verlag Berlin Heidelberg New York: alk. paper). - ISBN 0-387-51927-0 (Springer-Verlag New York Berlin Heidelberg: alk. paper) 1. Fishes - Physiology. 2. Animal communication. 3. Electroreceptors. 4. Fishes - Behavior. I. Title. II. Series. QL639.1.K74 1990 597'.50459-dc20 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, re-use of illustrations, recitation, broadcasting, reproduction on microfilms or in other ways, and storage in data banks. Duplication of this publication or parts thereof is only permitted under the provisions of the German Copyright Law of September 9, 1965, in its current version, and a copyright fee must alway be paid. Violations fall under the prosecution act of the German Copyright Law. © Springer-Verlag Berlin Heidelberg 1990 Printed in Germany The use of registered 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, printing and binding: Brühlsche Universitätsdruckerei, Giessen 2131/3145-543210 - Printed on acid-free paper Fisches Nachtgesang A fish's nightsong Fisches Nachtgesang: Das tiefste deutsche Gedicht A fish's nightsong: The deepest German poem Christian Morgenstern, Gesammelte Werke: Galgenlieder. Piper, München, Zürich 1965 Preface The aim of writing a book is to collect the currently available data on a chosen matter in order to realize a synthetic work. In our days, however, scientific methods evolve and develop at such a fast pace that because of the amount of new data appearing during the 1 to 2 years necessary for its preparation, the book may become obsolete before its publication. Hence, much courage is necessary to make such an effort and to write a book mainly presenting the state of actual knowledge. Electric fish biology has only a short history although the striking power of strong electric fish was already known by the Ancient Egyptians. It began in 1951 with Lissmann's discovery of the weak electric signals emitted by African tropical fish. During this 40-year period, despite a large number of scientific and popularized publications, only a small number of reviews and books have appeared on electric fish biology. Most of these dealt with anatomical, physiological and biophysical aspects of the electric organs and with problems concerning the peripheral and central nervous mechanisms of electrical signal emission and reception. They seldom, or only partially, touched on behavioural or ethological aspects. It is therefore my pleasure to welcome the present book on Electrocommunication in teleost fishes. Bernd Kramer's monograph, subtitled A fish's nightsong: the deepest German poem (C. Morgenstern) treats a particular sensory modality, the electric sense, used for communication by certain families of a class of vertebrates living in a conductive medium. It contains four chapters in which the different aspects of electrocommunication in relation to the different genera and species are discussed in a very systematic manner. In the first chapter a concise index of taxonomy and biogeography of (gymnotiform, mormyriform and siluriform) electroreceptive fish families is proposed, a use- ful key for comparative ethological research especially for gymnotiforms, the taxonomy of which means still a real burden for those working in this field. In the second and third chapters the author describes the characteristics (forms and patterns) of the emitted electrical signals, the structure of the specific sense organs and explains the modes of electric signal reception, the integration mechanisms of electrosensory information, as well as the input- output links (electroreception - electric emission and its neuronal command system). In the fourth chapter, the bulk of this monograph, the principles of communicating with electric signalling are systematically expounded and a comprehensive review is presented of many experimental data concerning the specific electric organ discharge patterns occurring during non-reproduc- tive social behaviour or courtship and spawning. A large part of this chapter is devoted to an analysis of the electrocommunicating system by experimental VII manipulation in order to understand the meaning of the electric signal pat- terns emitted in response to stereotyped exogenous electric stimuli in a given experimental context. I like this book because its encyclopedic character provides a very com- plete and clear picture of the current state of our knowledge on the subject. This well conceived book has the advantage over a multiauthor publication in that it covers this subject uniformly. The systematic presentation of neuroethological aspects from a comparative point of view will be useful in further investigations. It is a real pleasure to read the many minute historical notes distributed throughout the chapters evoking the time of pioneer scien- tists who worked in this particular field of sensory physiology. The nicely shown contrast between old and modern-day methods traces and explains the rapid widening of our understanding of neuroethological problems in electrocommunication. I sign this preface in friendship to Bernd Kramer and in remembrance of many pleasant hours spent together in our laboratories in Gif and in Regensburg. Gif-sur-Yvette, March 1990 T. SZABO VIII Acknowledgement To write a book while engaged in teaching, research and administration at a university is an extremely difficult job. First of all, I would like to thank my family for their understanding. Thomas Szabo stimulated my interest in communication in electric fish and gave me the opportunity to work in his laboratory for three most rewarding years. I wish to thank him and all colleagues from the College and Gif laboratory for their hospitality and support; last, but not least, also Dr. Helge Szabo. I am grateful for Hubert Markl's unstinting support of this research during my stay in his group at the University of Konstanz, and for an excellent and stimulating research environment. Hubert Markl also suggested that this book be written. I wish to thank all colleagues who collaborated with or joined our group for their precious contributions and their enthusiasm. It is especially for the younger among them that this book was written. D. Burkhardt provided the photograph of an Egyptian mural showing a mormyrid, represented on the book cover. B. Otto's help with the figures is highly appreciated. The Deutsche Forschungsgemeinschaft supported this research from the very beginning; without this invaluable support neither this book nor the work it is based on could have been achieved. I wish to express my gratitude