Hearing by Bats

526 Pages·1995·27.644 MB·English

by  Alan D. Grinnell (auth.), Arthur N. Popper, Richard R. Fay (eds.)

SPRINGER HANDBOOK OF
AUDITORY RESEARCH
Series Editors: Richard R. Fay and Arthur N. Popper
SPRINGER HANDBOOK OF AUDITORY RESEARCH
Volume 1: The Mammalian Auditory Pathway: Neuroanatomy
Edited by Douglas B. Webster, Arthur N. Popper, and Richard R. Fay
Volume 2: The Mammalian Auditory Pathway: Neurophysiology
Edited by Arthur N. Popper and Richard R. Fay
Volume 3: Human Psychophysics
Edited by William Yost, Arthur N. Popper, and Richard R. Fay
Volume 4: Comparative Hearing: Mammals
Edited by Richard R. Fay and Arthur N. Popper
Volume 5: Hearing by Bats
Edited by Arthur N. Popper and Richard R. Fay
Forthcoming volumes (partial list)
Auditory Computation
Edited by Harold L. Hawkins, Theresa A. McMullen, Arthur N. Popper,
and Richard R. Pay
Clinical Aspects of Hearing
Edited by Thomas R. Van de Water, Arthur N. Popper, and Richard R. Fay
Arthur N. Popper
Richard R. Fay
Editors
Hearing by Bats
With 138 Illustrations
Springer-Verlag
New York Berlin Heidelberg London Paris
Tokyo Hong Kong Barcelona Budapest
Arthur N. Popper Richard R. Fay
Department of Zoology Parmly Hearing Institute and
University of Maryland Department of Psychology
College Park, MD 20742, USA Loyola University of Chicago
Chicago, IL 60626, USA
Series Editors: Richard R. Fay and Arthur N. Popper
Cover illustration: A big brown bat, Eptesicusjuscus, is about to capture a mealworm hanging
on a fine filament. Photograph by S. P. Dear and P. A. Saillant. This figure appears on p. 148
of the text.
Library of Congress Cataloging in Publication Data
Hearing by bats / Arthur N. Popper, Richard R. Fay, editors.
p. cm. - (Springer handbook of auditory research: v. 5)
Includes bibliographical references and index.
ISBN-13: 978-1-4612-7577-0 e-ISBN-13: 978-1-4612-2556-0
DOl: 10.1007/978-1-4612-2556-0
1. Bats-Physiology. 2. Echolocation (physiology) 3. Hearing
Behavior. I. Popper, Arthur N. II. Fay, Richard R. III. Series.
QL737.C5H435 1995
599.4'041825 -dc20 94-41860
CIP
Printed on acid-free paper.
© 1995 Springer-Verlag New York, Inc.
Softcover reprint of the hardcover 1st edition 1995
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Series Preface
The Springer Handbook oj Auditory Research presents a series of com
prehensive and synthetic reviews of the fundamental topics in modern
auditory research. It is aimed at all individuals with interests in hearing
research including advanced graduate students, postdoctoral researchers,
and clinical investigators. The volumes will introduce new investigators to
important aspects of hearing science and will help established investigators
to better understand the fundamental theories and data in fields of hearing
that they may not normally follow closely.
Each volume is intended to present a particular topic comprehensively,
and each chapter will serve as a synthetic overview and guide to the
literature. As such, the chapters present neither exhaustive data reviews nor
original research that has not yet appeared in peer-reviewed journals. The
series focuses on topics that have developed a solid data and conceptual
foundation rather than on those for which a literature is only beginning to
develop. New research areas will be covered on a timely basis in the series
as they begin to mature.
Each volume in the series consists of five to eight substantial chapters on
a particular topic. In some cases, the topics will be ones of traditional
interest for which there is a solid body of data and theory, such as auditory
neuroanatomy (Vol. 1) and neurophysiology (Vol. 2). Other volumes in the
series will deal with topics which have begun to mature more recently, such
as development, plasticity, and computational models of neural processing.
In many cases, the series editors will be joined by a co-editor having special
expertise in the topic of the volume.
Richard R. Fay
Arthur N. Popper
v
Preface
Of all vertebrate groups, the echo locating mammals have probably pro
vided the most intrigue to auditory researchers. The biological significance
of echolocation sounds is obvious, yet the sounds are generally inaudible to
humans and are used in ways that seem alien to our auditory experience.
Of the echolocators, bats are the most widely and intensively studied, and
are the best understood. In fact, we probably know and understand more
about the acoustic neuroethology of bats than of any other vertebrate
group.
Our decision to include a volume on bats in the Springer Handbook oj
Auditory Research was motivated by several considerations. First, to our
knowledge, there has been no modern volume that addresses hearing and
the neuroethology of echolocation in bats in a systematic and comprehen
sive way. Thus, this volume provides detailed insights into the current state
of our knowledge on these fascinating species, as well as a guide to areas for
future research. Second, and as pointed out by George Pollak, Jeffery
Winer, and William O'Neill in Chapter 10, bats are exceptionally interesting
and useful model systems for the study of mammalian hearing in general,
and as such, this volume provides investigators interested in other species
with detailed insights into the auditory mechanisms and capabilities of this
model system. Finally, as pointed out elegantly by Alan Grinnell in Chapter
1, our understanding of hearing and echolocation by bats is " . . . one of
the triumphs of neuroethology . . . " and, as such, bats provide one of the
few comprehensive "stories" about the uses of sound in the auditory
literature.
Grinnell (Chapter 1) provides an overview of what is known about
hearing and echolocation by bats, and provides a valuable perspective on
the questions and controversies of greatest interest. In Chapter 2, Fenton
discusses the natural history and evolution of echolocation and provides an
introduction to the myriad species that are discussed in the rest of this
volume. The behavioral capabilities of bats are discussed in Chapter 3 by
Moss and Schnitzler, and discrimination is treated in detail by Simmons and
his colleagues in Chapter 4. The structure and function of the auditory
vii
viii Preface
system is dealt with in detail in Chapters 5 through 9. In Chapter 5, Kossl
and Vater describe the inner ear. The brainstem and lower auditory
pathways are discussed by Covey and Casseday in Chapter 6, while Pollak
and Park describe the inferior colliculus in Chapter 7. The thalamus is
treated in Chapter 8 by Wenstrup and the auditory cortex in Chapter 9 by
O'Neill. Finally, Pollak, Winer, and O'Neill give an overview of the
mammalian auditory system and provide compelling arguments for the use
of bats as a model system for mammalian hearing.
In inviting the authors to write chapters for this volume we asked them to
not only consider the species upon which they work, but also to provide a
comparative overview of the auditory system of the wide variety of bat
species. Thus, the volume not only deals with bat hearing per se, but also
provides a perspective that shows the wide range of variation in detection
and processing mechanisms even within this one amazing group of mam
mals.
The editors would like to express their gratitude to George Pollak for his
guidance, insight, and help in developing this volume. Dr. Pollak has a
unique and invaluable perspective on bats, and he shared his thoughts with
us freely and generously when we invited him to contribute chapters to this
volume.
Arthur N. Popper
Richard R. Fay
Contents
Series Preface ..................................................................... v
Preface.............................................................................. vii
Contributors. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi
Chapter 1 Hearing in Bats: An Overview .............................. 1
ALAN D. GRINNELL
Chapter 2 Natural History and Biosonar Signals..................... 37
M. BROCK FENTON
Chapter 3 Behavioral Studies of Auditory Information
Processing ........................................................ 87
CYNTHIA F. Moss AND HANS-ULRICH SCHNITZLER
Chapter 4 Auditory Dimensions of Acoustic Images in
Echolocation ..................................................... 146
JAMES A. SIMMONS, MICHAEL J. FERRAGAMO,
PRESTOR A. SAILLANT, TIM HARESIGN,
JANINE M. WOTTON, STEVEN P. DEAR,
AND DAVID N. LEE
Chapter 5 Cochlear Structure and Function in Bats ................. 191
MANFRED KbsSL AND MARIANNE VATER
Chapter 6 The Lower Brainstem Auditory Pathways ............... 235
ELLEN COVEY AND JOHN H. CASSEDAY'
Chapter 7 The Inferior Colliculus ........................................ 296
GEORGE D. POLLAK AND THOMAS J. PARK
Chapter 8 The Auditory Thalamus in Bats ............................ 368
JEFFREY J. WENSTRUP
Chapter 9 The Bat Auditory Cortex ..................................... 416
WILLIAM E. O'NEILL
ix
x Contents
Chapter 10 Perspectives on the Functional Organization
of the Mammalian Auditory System: Why Bats
Are Good Models............................................... 481
GEORGE D. POLLAK, JEFFERY A. WINER,
AND WILLIAM E. O'NEILL
Index ................................................................................ 499
Contributors
John H. Casseday
Department of Neurobiology, Duke University Medical Center, Durham,
NC 27710, USA
Ellen Covey
Department of Neurobiology, Duke University Medical Center, Durham,
NC 27710, USA
Steven P. Dear
Department of Neuroscience, Brown University, Providence, RI 02912,
USA
M. Brock Fenton
Department of Biology, York University, North York, Ontario, Canada
M3J IP3
Michael J. Ferragamo
Department of Neuroscience, Brown University, Providence, RI 02912,
USA
Alan D. Grinnell
Department of Physiology, UCLA School of Medicine, Jerry Lewis
Neuromuscular Research Center, Los Angeles, CA 90024, USA
Tim Haresign
Department of Neuroscience, Brown University, Providence, RI 02912,
USA
Manfred Kossl
Zoologisches Institut, Universitat Munchen, D8000 Munchen 2, Germany
xi