Neuroethology and Behavioral Physiology Roots and Growing Points

Edited by Franz Huber and Hubert Markl

With 183 Figures

Springer-Verlag Berlin Heidelberg New York Tokyo 1983

Professor Dr. Franz Huber Max-Planck-Institut flir Verhaltensphysiologie D-8131 Seewiesen/ Starnberg, F.R. Germany

Professor Dr. Hubert Markl UniversiHit Konstanz FakulHit fur Biologie Postfach 5560 D-7750 Konstanz, F.R. Germany

Cover illustrations

Above: Tracks made by a bat and a moth flying above a sun-lamp (combined filament and ultraviolet mercury lamp). Both enter from upper right. The ripple on both tracks is due to the 120ls flicker of the ultraviolet source. The small tracks were made by small insects flying over the lamp. K. D. Roeder: Animal Behaviour 10,300-304, Plate IV (1962)

Below: Bat captured moth at point where two white streaks intersect. Small streak shows the flight pattern of the moth. Broad streak shows the flight path of the bat. Both streak photographs were made by Frederic Webster ofthe Sensory Systems Laboratories. K. D. Roeder: Scientific American 212 (4), 95 (1965)

ISBN-13: 978-3-642-69273-4 e-ISBN-13: 978-3-642-69271-0 DOl: 10.1007/978-3-642-69271-0

Library of Congress Cataloging in Publication Data. Main entry under title: Neuro­ethology and behavioral physiology. I. Neuropsychology. 2. Animal behavior. I. Huber, Franz, 1925~ . II. Markl, Hubert, 1938~. QP360.N493. 1983.591.1'88.83-17025.

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 'Verwertungsgesellschaft Wort', Munich.

© by Springer-Verlag Berlin Heidelberg 1983. Softcover reprint of the hardcover 1st edition 1983

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.

2131/3130-543210

This book is dedicated to the memory of

Kenneth David Roeder (1908-1979)

Biologist and Humanist, a Founding Father of Neuroethology, Friend of Insects and Friend of People

Preface

The investigation of the relationships between a behavior pattern and its underlying sensory and neurophysiological mechanisms in both man and animals dates back well into the last century. However, the concepts and findings of ethology and experimental psychology, together with an improved understanding of how the nervous system is organized and how neurons interact with each other, have only in the last 30 years laid the groundwork for an in-depth analysis. The many technological advances achieved in neurophysiology and neuroanatomy have also played an important role in this. The study of the neuronal bases of behavior - for which the term "neuroethology" has been coined - has thus become one of the central themes of neuroscience.

Kenneth David Roeder, who died in 1979, was one of the pioneers of this field of research. It is to him that the contributions in this book are dedicated. K.D. Roeder was among the first to attempt to define the correlation between the natural behavior of an experimental animal and the activity of single sensory and nerve cells. The ques­tions he asked, his experimental approach, and his fundamental discoveries are pre­sented in an introductory chapter.

The contributions in this book are based on a symposium that was held in Mainz in September 1982 and that was made pOSSible by the kind support of the Stiftung Volkswagenwerk, the Max-Planck-Gesellschaft, and above all, the Mainz Academy of Science and Literature. Two of the goals of this gathering of experts were to sum­marize the present state of neuroethological knowledge and to point out new concep­tual and methodological directions based on the example of especially well-investi­gated model organisms and their behavioral strategies.

This book covers almost all levels of behavior. Its topics range from motor compo­nent systems to movement sequences subserving behavior patterns such as prey cap­ture, mate finding, and social interactions (including the orientational capabilities required by these behaviors) to the more complex levels of neural integration such as plasticity phenomena and learning.

Each chapter represents the personal bias ofits author, and the reader is challenged to follow the course of each analysiS to its conclusion. The reader is, thus, provided a picture of the new methods now being used in the field of quantitative behavioral phYSiology and neurobiology and is obliged to make a critical judgement about the direction of future experimental developments. As is readily apparent from many of the chapters, neuroethology is an interdisciplinary field. However, a gap still exists between ethology, psychology, behavioral ecology, and sociobiology on the one side and the palette of strictly neurobiological research directions on the other. This gap can be successfully bridged only if the versatile new techniques are employed on the

VIII Preface

basis of carefully designed laboratory experiments aimed at elucidating behavioral strategies used by a freely moving animal in its natural habitat.

The authors and editors hope that this book, in memory of K.D. Roeder, will point the way for further research into the many yet unanswered questions. It is addressed to undergraduate and graduate students of zoology, ethology, and psychology, as well as to students of all aspects of neurobiology and to experts in these disciplines.

We would like to express our thanks to Springer-Verlag for the excellent production of this volume.

Summer 1983 F. HUBER and H. MARKL

Contents

Kenneth David Roeder (1908-1979) Curriculum Vitae and List of Publications . . . . . . . . . . . . . . . . . . . . . . . . .. XV

K.D. Roeder's Impact on Insect Neuroethology Franz Huber. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

I. Neural Bases of Motor Systems

1.1 On the Way to Neuroethology: The Identified Neuron Approach (With 11 Figures) Graham Hoyle . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

1.2 Local Interneurones and the Control of Movement in Insects (With 8 Figures) Malcolm Burrows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 26

1.3 The Interaction of Peripheral and Central Components in Insect Locomotion (With 12 Figures) Gernot Wendler. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 42

1.4 A Neuroethological Approach to the Phylogeny of Leg Stridulation in Gomphocerine Grasshoppers (With 5 Figures) Norbert Elsner . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 54

II. Sensory Strategies of Adaptation

11.1 K.D. Roeder and the History of Chemoreception Vincent G. Dethier . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 71

11.2 Olfactory Food and Mate Recognition (With 9 Figures) Jtirgen Boeckh and Klaus-Dieter Ernst. . . . . . . . . . . . . . . . . . . . . . . .. 78

11.3 Species Recognition and Acoustic Localization in Acridid Grasshoppers: A Behavioral Approach (With 11 Figures) Dagmar v. Helversen and Otto v. Helversen. . . . . . . . . . . . . . . . . . . . .. 95

x

11.4 Neural Correlates of Orthopteran and Cicada Phonotaxis (Wi th 18 Figures)

Contents

Franz Huber. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 108

11.5 Frequency and Temporal Processing in the Auditory System of Anurans (With 11 Figures) Robert R. Capranica and Gary Rose . . . . . . . . . . . . . . . . . . . . . . . . .. 136

11.6 Optics and Accommodation in Owls and Flying Foxes (With 5 Figures) Howard C. Howland. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 153

III. Neural Development and Behavioral Plasticity

111.1 Insect Ecdysis: A System for the Study ofInternal Chemicals That Control Behavior (With 3 Figures) James W. Truman. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 167

I1I.2 Maps in the Insect Nervous System, Their Implications for Synaptic Connectivity and Target Location in the Real World (With 6 Figures) Rodney K. Murphey . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 176

I1I.3 Neuroethological Studies of Associative Learning in Feeding Control Systems (With 9 Figures) Alan Gelperin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 189

IlIA Learning by Honeybees in an Unnatural Situation (With 5 Figures) Randolf Menzel and M.E. Bitterman. . . . . . . . . . . . . . . . . . . . . . . . .. 206

111.5 The Search for Neural Correlates of Learning in the Honeybee (With 4 Figures) Joachim Erber . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 216

IV. Predator - Prey - Relationships

IV.1 Moth Hearing and Bat Sounds: The History of a Collaboration (With 1 Figure) Asher E. Treat . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 231

IV.2 The Acoustical Image of Fluttering Insects in Echolocating Bats (With 7 Figures) Hans-Ulrich Schnitzler, Dieter Menne, Rudi Kober, and Klaus Heblich . .. 235

IV.3 How Insects Detect and Avoid Bats (With 9 Figures) Lee A. Miller . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 251

Contents XI

N.4 futerception of Flying Insects by Bats James A. Simmons and Shelley A. Kick. . . . . . . . . . . . . . . . . . . . . . .. 267

N.5 Echolocation and Adaptivity to Ecological Constraints (With 10 Figures) Gerhard Neuweiler. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 280

N.6 Neuroethology of Acoustic Prey Localization in the Barn Owl (With 10 Figures) Masakazu Konishi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 303

V. Ecological Aspects of Social Relations and Orientation

V.1 Strategies for Acoustic Communication in Complex Environments (With 5 Figures) Axel Michelsen and Ole N. Larsen ........ . . . . . . . . . . . . . . . . . .. 321

V.2 Vibrational Communication Hubert Markl . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 332

V.3 Chemical Manipulation, Enemy Specification and futercolony Communication in Ant Communities (With 3 Figures) Bert Holldobler ....................................... 354

V.4 Celestial and Terrestrial Navigation: Human Strategies -fusect Strategies (With 7 Figures) Rudiger Wehner. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 366

V.5 Towards Understanding the Flow of Information Between Objective and Subjective Space (With 13 Figures) Horst Mittelstaedt. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 382

Epilogue: Neurobiological Roots and Neuroethological Sprouts Theodore H. Bullock. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 403

Contributors

You will find the addresses at the beginning of the respective contributions

Bitterman, M.E. 206 Boeckh, Jiirgen 78 Bullock, Theodore H. 403 Burrows, Malcolm 26 Capranica, Robert R. 136 Dethier, Vincent G. 71 Elsner, Norbert 54 Erber, Joachim 216 Ernst, Klaus-Dieter 78 Gelperin, Alan 189 Heblich, Klaus 235 Helversen, Dagmar v. 95 Helversen, Otto v. 95 Holldobler, Bert 354 Howland, Howard C. 153 Hoyle, Graham 9 Huber, Franz 1, 108 Kick, Shelley A. 267

Kober, Rudi 235 Konishi, Masakazu 303 Larsen, Ole N. 321 Markl, Hubert 332 Menne, Dieter 235 Menzel, Randolf 206 Michelsen, Axel 321 Miller, Lee A. 251 Mittelstaedt, Horst 382 Murphey, Rodney K. 176 Neuweiler, Gerhard 280 Rose, Gary 136 Schnitzler, Hans-Ulrich 235 Simmons, James A. 267 Treat, Asher E. 231 Truman, James W. 167 Wehner, Rudiger 366 Wendler, Gernot 42

Kenneth David Roeder (1908-1979)

K.D. Roeder was born in Richmond, Middlesex, England on March 9, 1908. He went to Bembridge School, Isle of Wight, before entering Cambridge University, Natural Science Tripos (1926-1929) and studied Zoology (1930-1931). There he received the degrees of B.A. Hons.(1929) and of M.A. (1933). In 1931 he married Sonja von Cancrin, and in the same year left England to take over a teaching assistentship at Toronto University. After a few months he went to the United States and became in­structor in biology at Tufts University, where he stayed until his death in 1979. In 1950 Roeder was promoted to Professor of PhYSiology at Tufts, in 1951 he received the D.Sc. (Honorary) from Tufts University, was Chairman of the Department of Biol­ogy from 1959 to 1964, and from 1964 to 1975 Research Professor on a NIH Career Award.

K.D. Roeder was member of the National Academy of Sciences, honorary member of the Royal Entomological Society (London) and member of the Akademie der Natur­forscher, Leopoldina, Halle. He was also member of A.A.A.S., the American Society of Zoologists, the Entomological Society of America, Sigma Xi, the Society for General PhYSiologists, Phi Beta Kappa, and Fellow of the American Academy of Arts and Sciences. In 1946 he acted as member of the DDT Panel, in 1950 of the Codification Committee of the National Research Council, became member of the Study Section of ParaSitology and Tropical Medicine and that of PhYSiology of the NIH, and member of the Cotton Insect Control Panel of the Office of Science and Technology (1961-1963).

K.D. Roeder's research interests focused on the neural mechanisms of insect behav­ior, in which field he became a pioneer and leader. He published 88 research papers and reviews, was editor and author of Insect Physiology (Wiley 1953), and author of Nerve Cells and Insect Behavior (Harvard 1st edn 1963, 2nd edn 1967). From 1960 to 1964 he was on the Editorial Board of the Annual Review of Entomology; he also served on the Advisory Boards of the Journal of Comparative PhYSiology and the Journal of Insect Physiology. In 1953 he was a Fulbright Scholar, in 1946 Lowell lec­turer (Boston), in 1960 AIBS Lecturer (San Francisco), in 1966 Founders' Memorial Lecturer, Entomological Society of America (Oregon) and in 1974 Griswold Lecturer (Cornell University, Ithaca).

List of Publications of K. D. Roeder

(Without short notes and abstracts of papers presented at meetings)

Roeder KD (1934) A simple method for measuring small time intervals. Science 79:82-83 Roeder KD (1935) An experimental analysis of the sexual behavior of the praying mantis. BioI

Bull 59:203-220 Roeder KD (1936) Raising the praying mantis for experimental purposes. Science 83:582-583 Roeder KD (1937) The control of tonus and locomotor activity in the praying mantis. J Exp Zool

76:353-374 Roeder KD (1939a) The action of certain drugs on the insect central nervous system. BioI Bull 76:

183-189 Roeder KD (1939b) Synchronized activity in the optic and protocerebral ganglia of the grasshop­

per, Melanoplus femur·rubrum. J Cell Comp PhysioI14:299-307 Roeder KD, Roeder S (1939) Electrical activity in the isolated ventral nerve cord of the cockroach.

1. The action of pilocarpine, nicotine, eserine and acetylcholine. J Cell Comp Physiol14: 1-12 Roeder KD (1940a) A rubber cast of the dogfish spiral valve. Science 91:321 Roeder KD (1940b) The origin of visual rhythms in the grasshopper, Melanoplus femur·rubrum.

J Cell Comp PhysioI16:1-2 Roeder KD (1941a) A case for the cockroach. Tuftonian 1 :18-20 Roeder KD (1941b) The effect of potassium and calcium on the spontaneous activity of the iso-

lated crayfish nerve cord. J Cell Comp PhysioI18:1-9 Roeder KD (1946) Fine-tapered silver electrodes for physiologist's work. Science 104:425-426 Roeder KD, Weiant EA (1946) The site of action of DDT in the cockroach. Science 103:304-306 Roeder KD, Kennedy NK, Samson EA (1947) Synaptic conduction to giant fibers an4 the action

of anticholinesterases. J Neurophystoll0: 1-10 Roeder KD, Wellman B (1947) An electronic counter for rapid impulses. Electronics October 1947 Roeder KD (1948a) The effect of potassium and calcium on the nervous system of the cockroach,

Periplaneta americana. J Cell Comp Physio131:327 -338 Roeder KD (1948b) Organization of the ascending giant fiber system of the cockroach, Periplaneta

americana. J Exp ZooI108:243-261 Roeder KD (1948c) The effect of anticholinesterases and related substances on nervous activity in

the cockroach. Bull Johns Hopkins Hosp 83:587-600 Roeder KD, Edwards GA, Weiant EA, Slocombe AG (1948) The action of ryanodine on the con­

tractile process in striated muscles. Science 108:330-332 Roeder KD, Weiant EA (1948) The effect of DDT on sensory and motor structures in the cock­

roach leg. J Cell Comp Physiol 32: 175-186 Roeder KD, Weiant EA (1950) The electrical and mechanical events of neuromuscular transmission

in the cockroach, Periplaneta americana. J Exp BioI 27: 1-13 Roeder KD (1951) Movements of the thorax and potential changes in the thoracic muscles of in­

sects during flight. BioI Bull 100:95-106 Roeder KD, Weiant EA (1951) The effect of concentration, temperature, and washing on the time

of appearance of DDT-induced trains in sensory fibers of the cockroach. Ann Entom Soc Am 44:372-380

Roeder KD (1952) Insects as experimental material. Science 115:275-280 Roeder KD (1953a) Electric activity in nerves and ganglia. In: Roeder KD (ed) Insect Physiology.

John Wiley and Sons, New York, pp 423-462

List of Publications of K.D. Roeder XVII

Roeder KD (1953b) Reflex activity and ganglion function. In: Roeder KD (ed) Insect physiology. John Wiley and Sons, New York, pp 463-487

Roeder KD (1955) Spontaneous activity and behavior. Sci Mon 80:362-370 Roeder KD, Kennedy NK (1955) The effect of certain trisubstituted phosphine oxides on synaptic

conduction in the roach. J Pharmacol Exp Therap 114:211-220 Hodgson ES, Lettvin JY, Roeder KD (1955) Physiology of primary chemoreceptor unit. Science

122:417 -418 Hodgson ES, Roeder KD (1956) Electrophysiological studies of arthropod chemoreception. I. Gen­

eral properties of the labellar chemoreceptors of Diptera. J Cell Comp PhysioI48:51-76 Roeder KD, Treat AE (1956) Electrical response of the noctuid tympanum to ultrasonic stimula­

tion. Proc Xth Int Congr ZooI2:117 -120 Twarog BM, Roeder KD (1956) Properties of the connective tissue sheath of the cockroach abdom­

inal nerve cord. BioI Bull 111:278-286 Rilling S, Mittelstaedt H, Roeder KD (1957) Prey recognition in the praying mantis. Behaviour 14:

164-184 Roeder KD, Treat AE (1957) Ultrasonic reception by the tympanic organ of noctuid moths. J Exp

Zool134:127 -158 Twarog BM, Roeder KD (1957) Pharmacological observations on the desheathed last abdominal

ganglion of the cockroach. Ann Entomol Soc Am 50:231-237 Roeder KD (1958) The nervous system. Ann Rev EntomoI3:1-18 Roeder KD (1959) A physiological approach to the relation between prey and predator. Smithson

MiscCollect 137:287-306 Roeder KD, Treat AE (1959) A nervous element of unknown function in the tympanic organ of

moths. J Insect PhysioI3:262-270 Milburn N, Weiant EA, Roeder KD (1960) The release of efferent nerve activity in the roach, Peri­

planeta americana, by extracts of the corpus cardiacum. BioI Bull 118: 111-119 Roeder KD (1960) The predator and display strikes of the praying mantis. Med BioI Illus 10: 172-

178 Roeder KD, Tozian L, Weiant EA (1960) Endogenous nerve activity and behaviour in the mantis

and the cockroach. J Insect PhysioI4:45-62 Roeder KD, Treat AE (1961a) The detection and evasion of bats by moths. Am Sci 49:135-148 Roeder KD, Treat AE (1961b) The reception of bat cries by the tympanic organ ofnoctuid moths.

In: Rosenblith W (ed) Sensory communication. MIT Press, Cambridge, MA and Wiley, New York, pp 545-560

Roeder KD (1962a) Ultrasonic interaction of bats and moths. In: Bernard EE, Kare MR (eds) Biological prototypes and synthetic systems, vol I. Plenum Press, New York London, pp 54-57

Roeder KD (1962b) Neural mechanisms of animal behavior. Am ZooI2:105-115 Roeder KD (1962c) The behaviour of free flying moths in the presence of artificial ultrasonic

pulses. Anim Behav 10:300-304 Milburn N, Roeder KD (1962) Control of efferent activity in the cockroach terminal abdominal

ganglion by extracts of corpora cardiaca. Gen Comp Endocrinol 2: 7 0-7 6 Roeder KD (1963) Nerve cells and insect behavior. Harv Books BioI Nr 4, Harvard University Press,

Cambridge Mass Roeder KD (1963a) Ethology and Neurophysiology. Z TierpsychoI20:434-440 Roeder KD (1963b) Echoes of ultrasonic pulses from flying moths. BioI Bull 124:200-210 Roeder KD (1964a) Night fighters in a sonic duel. Nat Hist January 33-39 Roeder KD (1964b) Aspects of the noctuid tympanic nerve response having significance in the

avoidance of bats. J Insect Physioll0:529-546 Roeder KD (1965a) Moths and ultrasound. Sci Am 212:94-102 Roeder KD (1965b) Epilogue. In: Treherne JE, Beament JWL (eds) The physiology of the insect

central nervous system. Academic Press, London New York, pp 247-252 Dunning DC, Roeder KD (1965) Moth sounds and the insect catching behavior of bats. Science

147:173-174 Roeder KD, Payne RS (1965) Acoustic orientation of a moth in flight by means of two sense cells.

Symp Soc Exp Bioi 20:251-272

XVIII List of Publications of K.D. Roeder

Payne RS, Roeder KD, Wallman J (1966) Directional sensitivity of the ears of noctuid moths. J Exp Bioi 44:17 -31

Roeder KD (1966a) Acoustic sensitivity of the noctuid tympanic organ and its range for the cries of bats. J Insect PhysioI12:843-859

Roeder KD (1966b) Interneurons of the thoracic nerve cord activated by tympanic nerve fibers in noctuid moths. J Insect PhysioI12:1227-1244

Roeder KD (1966c) A differential anomometer for measuring the turning tendency of insects in stationary flight. Science 153: 1634-1636

Roeder KD (1966d) The auditory system ofnoctuid moths. Science 154:1515-1521 Roeder KD (1967a) Nerve cells and insect behavior. Harvard Books in Biology, 2nd edn. Harvard

Univ Press, Cambridge Mass Roeder KD (1967b) Turning tendency of moths exposed to ultrasound while in stationary flight.

J Insect Physiol13:873-888 Roeder KD (1967c) Prey and predator. Bull Entomol Soc Am 13:6-9 Roeder KD (1968a) Three views of the nervous system. James Arthur Lecture, American Museum

of Natural History NY 28 pp Roeder KD (1968b) Interactions of bats and moths. Kybernetik 75-87 Roeder KD, Treat AE, Vande Berg JS (1968) Auditory sense in certain sphingid moths. Science

159:331-333 Roeder KD (1969a) Acoustic interneurons in the brain of noctuid moths. J Insect Physiol15:

825-838 Roeder KD (1969b) Brain interneurons in noctuid moths: Differential suppression by high sound

intensities. J Insect Physiol15: 1713 -1 718 Roeder KD (1970) Episodes of the insect brain. Am Sci 58:378-389 Roeder KD, Treat AE (1970) An acoustic sense in some hawkmoths. J Insect Physiol 16:1069-

1086 Roeder KD, Treat AE, Vande Berg JS (1970) Distal lobe of the pilifer: An ultrasonic receptor in

choerocampine hawkmoths. Science 170:1098-1099 Roeder KD (1971a) Insect flight behavior: Some neurophysiological indications of its control. Prog

Physiol PsychoI4:1-36 Roeder KD (1971b) Acoustic alerting mechanisms in insects. Ann NY Acad Sci 188:63-79 Roeder KD (1972) Acoustic and mechanical sensitivity of the distal lobe of the pilifer in choero­

campine hawkmoths. J Insect PhysioI18:1249-1264 Roeder KD (1973a) Brain interneurons in noctuid moths: binaural excitation and slow potentials.

J Insect Physiology 19:1591-1601 Roeder KD (1973b) A dog's world view. Nat Hist , August 1973 Roeder KD, Fenton MB (1973) Acoustic responsiveness of Scoliopteryx libatrix L. (Lepidoptera:

Noctuidae), a moth that shares hibernacula with some insectivorous bats. Can J ZooI51:681-685

Fenton MB, Roeder KD (1974) The microtymbals of some Arctiidae. J Lepidopt Soc 28:205-211 Roeder KD (1974a) Some neural mechanisms of simple behavior. Adv Study Behav 5:1--A6 Roeder KD (1974b) Responses of the less sensitive acoustic sense cells in the tympanic organs of

some noctuid and geometrid moths. J Insect PhysioI20:55-66 Roeder KD (1975a) Acoustic interneurons responses compared in certain hawkmoths. J Insect

PhysioI21:1625-1631 Roeder KD (1975b) Neural transactions during acoustic stimulation ofnoctuid moths. Adv Behav

Bioi 15:99-115 Roeder KD (1975c) Feedback, spontaneous activity, and behavior. In: Baerends G, Beer C, Man­

ning A (eds) Function and evolution in behaviour. Essays in honour of Professor Niko Tinber­gen, FRS. Oxford Univ Press, pp 55-70

Roeder KD (1976) Joys and frustrations of doing research. Perspect BioI Med pp 231-245