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Sensory Biology of Aquatic Animals Conference Participants Sensory Biology of Aquatic Animals Conference participants. From left to right: Front, row 1: W. S. M. Hagedorn, J. Wyneken, M. Salmon, C. McCormick, J. Song, Wilcox, W. Saidel, W. Plassman, B. Fritsch, H. Zakon, M. Braford, M. Kreithen, M. Wullimann, J. Case, J. Gray, M. Powers, J. Levine, P. Borroni, B. U. Budelmann, C. Platt, H. Bleckmann, C. Hawr­ D. Hoekstra, T. Finger, P. Hamilton, D. Woodward, A. Kalmijn, wyshyn, J. Sivak. Middle, row 2: T. Waterman, B. Sokolowski, T. Cronin, T. Bullock, M. Laverack, H. Munz, J. Douglass, S. W. Heiligenberg, M. Swain, S. Coombs, E. Denton, R. G. Northcutt, Holderman. Last, row 4: C. Hopkins, W. Stachnik, T. Ream, P. W. Tavolga, J. Atema, R. Fay, A. Popper, J. Patton, P. Gomer, Gilbert, J. Kendall, B. Ache, A. Elepfandt, R. Barlow, R. Brill, P. Moller, J. Caprio, J. Crawford, J. Blaxter, R. Voigt, J. Webb, R. Fernald, R. Eaton, B. Zahuranec, W. Carr, J. Janssen, J. Lythgoe, R. Gleeson, C. Derby, I. Assip. Back, row 3: P. Rogers, M. Cox, J. R. Strickler, E. Hartwig, K. Wiese. lelle Atema Richard R. Fay Arthur N. Popper William N. Tavolga Editors Sensory Biology of Aquatic Animals Springer-Verlag New York Berlin Heidelberg London Paris Tokyo JELLE ATEMA, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts 02543, USA Richard R. Fay, Parmly Hearing Institute, Loyola University, Chicago, Illinois 60626, USA ARTHUR N. POPPER, Department of Zoology, University of Maryland, College Park, MD 20742, USA WILLIAM N. TAVOLGA, Mote Marine Laboratory, Sarasota, Florida 33577, USA The cover illustration is a reproduction of Figure 13.3, p. 343 of this volume Library of Congress Cataloging-in-Publication Data Sensory biology of aquatic animals. Papers based on presentations given at an International Conference on the Sensory Biology of Aquatic Animals held, June 24-28, 1985, at the Mote Marine Laboratory in Sarasota, Fla. Bibliography: p. Includes indexes. 1. Aquatic animals--Physiology-Congresses. 2. Senses and sensation-Congresses. I. Atema, Je1le. II. International Conference on the Sensory Biology of Aquatic Animals (1985 : Sarasota, Fla.) QL120.S46 1987 591.92 87-9632 © 1988 by Springer-Verlag New York Inc. Softcover reprint of the hardcover 1st edition 1988 All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher (Springer-Verlag, 175 Fifth Avenue, New York 10010, U.S.A.), except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. The use of general descriptive names, trade names, trademarks, etc. In this publication, even if the former are not especially identified, is not to 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. Typeset by Arcata Graphics/Kingsport, Kingsport, Tennessee. 9 8 7 6 5 4 3 2 1 ISBN-13: 978-1-4612-8317-1 e-ISBN-13: 978-1-4612-3714-3 DOl: 10.1007/978-1-4612-3714-3 This volume is dedicated to Sven Dijkgraaf Karl von Frisch C. Judson Herrick Erich von Holst George H. Parker Jacob von Uexkull J. Z. Young Each of these investigators delved deeply and broadly into the sensory biology of aquatic animals. Their contributions to the study of sensory systems of aquatic animals have directly and indirectly influenced all of the work reported in this volume. Preface This volume constitutes a series of invited chapters based on presentations given at an International Conference on the Sensory Biology of Aquatic Animals held June 24-28, 1985 at the Mote Marine Laboratory in Sarasota, Florida. The immediate purpose of the conference was to spark an exchange of ideas, concepts, and techniques among investigators concerned with the different sensory modalities employed by a wide variety of animal species in extracting information from the aquatic environment. By necessity, most investigators of sensory biology are specialists in one sensory system: different stimulus modalities require different methods of stimulus control and, generally, different animal models. Yet, it is clear that all sensory systems have principles in common, such as stimulus filtering by peripheral structures, tuning of receptor cells, signal-to-noise ratios, adaption and disadaptation, and effective dynamic range. Other features, such as hormonal and efferent neural control, circadian reorganization, and receptor recycling are known in some and not in other senses. The conference afforded an increased awareness of new discoveries in other sensory systems that has effectively inspired a fresh look by the various participants at their own area of specialization to see whether or not similar principles apply. This inspiration was found not only in theoretical issues, but equally in techniques and methods of approach. The myopy of sensory specialization was broken in one unexpected way by showing limitations of individual sense organs and their integration within each organism. For instance, studying vision, one generally chooses a visual animal as a model. Subsequent intense focus on that animal's visual performance easily leads to ignoring the fact that in the real world this animal uses other senses as well. Thus, the demands the animal places on vision are tempered in the context of other sensory information. The behavioral demands on visual detection range and acuity for instance may not be as severe as one might think, if it were known that acoustic or chemical senses serve to locate and identify the source of interest for that animal at greater distances, whereas vision is used mainly at shorter range. This multisensory conference thus allowed the different senses to show not only their individual strengths, but also viii Preface their weaknesses and limitations, and their inherent interrelatedness as they appear in the aquatic environment. It is our hope that these issues will not be lost in book form. The long-range purpose of the conference was to create a book that could serve as a conceptual framework for further investigations of the aquatic "Umwelt" and the intricate, often unexpected, methods and processes used by animals to utilize the information contained in the acquatic stimulus world. This book and its organization reflect the central premise that sensory systems are constrained by the stimulus environ­ ment (Section I), by the behavioral requirements of the organism (Section II), and by phylogeny (various chapters in Section III). In addition, there is the tacit assumption of economy in sensory design. A major and unique effort was made to stimulate the development of conceptual models of the aquatic stimulus environment per se. This, we believe, is of paramount importance since only a detailed and comprehensive understanding of the natural distribution of stimuli and their noise backgrounds will allow the design of the proper experiments which, in tum, will lead to an understanding of sensory function in a natural context. This focus primarily serves to probe specific aquatic problems, but it also bears on terrestrial sensory problems by providing the aquatic contrast. The aquatic environment is a different medium than the terrestrial environment in nearly all aspects of sensory biology. Some differences are profound, such as the possibility of electroreception in the conductive aquatic medium. Other differences are less funda­ mental, such as the propagation speed of sound and the properties of water as a filter for light and a carrier for chemical stimuli. The sensory constraints imposed by the stimulus environment may emerge in clearest form when comparing aquatic and terrestrial solutions to similar sensory problems. Although a number of chapters refer explicitly to terrestrial animals and environments for contrast, a systematic compar­ ison falls beyond the scope of this volume. Intimately related to the direct constraints and opportunities presented by the sensory "Umwelt" are indirect sensory constraints imposed by animal behavior, the motoric "Umwelt." An animal's behavioral requirements invite, to various degrees, the devel­ opment of sensory capabilities. The reverse may be true as well: extended sensory horizons may allow the expansion of motoric virtuosity. Thus, to understand the physiology of receptor cells and organs it is instructive to know the animal's behavioral capabilities. In the aquatic environment these capabilities are influenced to a large degree by the physical density and viscosity of water, regardless of whether the animal moves through the medium or moves water through or along its body. An understanding of the constraints of behavior is a rich source of inspiration in the study of sensory function. Ultimately all sensory processing stands in the service of making behavioral decisions. These are often decisions of immediate life or death, and always decisions affecting competitive fitness. The central question concerns what an animal must know about its environment in order to survive and compete successfully. Behavioral ecology can show the knowledge that an animal must have had about its environment in order to have performed a certain behavior, or the information required to have followed a certain behavioral strategy. The acquisition of this knowledge is at least partially a matter of sensory information. Memory and expectation are also essential ingredients of this knowledge, and both memory and expectation require constant updating by sensory information. Preface ix In contrast to the acquisition of information, we recognize stimulus acquisition behavior, i.e., those behavioral actions and strategies that increase the animal's proba­ bilities of interfacing with the proper sensory stimuli. These actions include search patterns, choice of location, current generation, acoustic and visual scanning, flicking and sniffing, probing, and any other behavior that appears designed specifically or primarily to serve the function of encountering further sensory cues that, in tum, may provide information about the source of the stimulus.
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