Table Of ContentHandbook
of
Sensory Physiology
VolumeV/l
Editorial Board
H.Autrum R.Jung W.R. Loewenstein
D.M.MacKay H.L.Teuber
A uditory System
Anatomy
Physiology (Ear)
By
H.W.Ades A.Axelsson I.L.Baird G.v.Bekesy R.L.Boord
C. B. G. Campbell O. Densert D. H. Eldredge H. Engstrom
J.Fex J.M.Harrison O.W.Henson M.E.Howe S.Iurato
A. Michelsen A. R. M011er R. R. Pfeiffer S. and I. Rauch
E.A.G.Shaw J.Wersiill E.G. Wever
Edited by
Wolf D. Keidel and William D. Neff
With 305 Figures
Springer-Verlag Berlin Heidelberg New York 1974
Wolf D. Keidel
I. Physiologisches Institut der Universitat 852 Erlangen, UniversitatsstraJ3e 17 (Germany)
William D. Neff
Indiana University, Center of Neural Sciences Psychology Building 320, Bloomington, Indiana 47401 (USA)
ISBN-13: 978-3-642-65831-0 e-ISBN -13: 978-3-642-65829-7
DOl: 10.1007/978-3-642-65829-7
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 1974. Library of
Congress Catalog Card Number 74-415_ The use of general descriptive names, trade names, trade marks, etc. in this
publication, even if the former are not especially identified, is not to be taken as a sign that such names, as under
stood by the Trade Marks and Merchandise Marks Act, may accordingly be used freely by anyone. Typesetting,
Softcover reprint of the hardcover 1s t edition 1974
Preface
In planning The Handbook volumes on Audition, we, the editors, made the
decision that there should be many authors, each writing about the work in the
field that he knew best through his own research, rather than a few authors who
would review areas of research with which they lacked first hand familiarity.
For the purposes of the chapters on Audition, sensory physiology has been
defined very broadly to include studies from the many disciplines that contribute
to our understanding of the structures concerned with hearing and the processes
that take place in these structures in man and in lower animals. A number of
chapters on special topics have been included in order to present information
that might not be covered by the usual chapters dealing with anatomical, physi
ological and behavioral aspects of hearing.
We wish to thank all authors of the volumes on Audition for the contributions
that they have made. We feel confident that their efforts will also be appreciated
by the many scientists and clinicians who will make use of the Handbook for
many years to come.
WOLF D. KEIDEL WILLIAM D. NEFF
Erlangen Bloomington
August 1974
Contents
Chapter 1 Introduction. By G. v. BEKESY t. With 3 Figures. . . . . . . . 1
Chapter 2 Consideration of the Acoustic Stimulus. By R. R. PFEIFFER. With
19 Figures. . . . . . . . . . . . . . . . . . . . . . . . . 9
Chapter 3 Comparative Anatomy of the Middle Ear. By O. W. HENSON Jr. With
23 Figures. . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Chapter 4 The Morphology of the Middle Ear Muscles in Mammals. By O. DENSERT
and J. WERSALL. With 12 Figures ................. 111
Chapter 5 Anatomy of the Inner Ear. By H. W. ADES and H. ENGSTROM. With
26 Figures. . . . . . . . . . . . . . . . . . . . . . . . . . . . 125
Chapter 6 Anatomical Features of the Inner Ear in Submammalian Vertebrates.
By 1. L. BAffiD. With 18 Figures . . . . . . . . . . . . . . . . . . 159
Chapter 7 The Blood Supply of the Inner Ear of Mammals. By A. AXELSSON. With
36 Figures. . . . . . . . . . . . . . . . . . . . . . . . . 213
Chapter 8 Efferent Innervation ofthe Cochlea. By S. IURATO. With 11 Figures 261
Chapter 9 Anatomy of the Afferent Auditory Nervous System of Mammals. By J. M.
HARRISON and M. E. HOWE. With 25 Figures . . . . . . . . . . . . . 283
Chapter 10 Central Auditory Pathways of Nonmammalian Vertebrates. By C. B. G.
CAMPBELL and R. L. BOORD. With 4 Figures. . . . . . . . . . . . . 337
Chapter 11 Anatomy of the Descending Auditory System (Mammalian). By J. M.
HARRISON and M. E. HOWE. With 10 Figures. . . . . . . . 363
Chapter 12 Hearing in Invertebrates. By A. MICHELSEN. With 18 Figures 389
Chapter 13 The Evolution of Vertebrate Hearing. By E. G. WEVER. With 14 Figures 423
Chapter 14 The External Ear. By E. A. G. SHAW. With 22 Figures . . . . 455
Chapter 15 Function of the Middle Ear. By A. R. MOLLER. With 19 Figures 491
Chapter 16 The Acoustic Middle Ear Muscle Reflex. By A. R. MOLLER. With 13 Figures 519
Chapter 17 Inner Ear - Cochlear Mechanics and Cochlear Potentials. By D. H.
ELDREDGE. With 9 Figures . . . . . . . . . . . . . . . . . . . . 549
Chapter 18 Neural Excitatory Processes of the Inner Ear. By J. FEX. With 10 Figures 585
Chapter 19 Physico-Chemical Properties of the Inner Ear Especially Ionic Transport.
By S. and 1. RAUCH. With 13 Figures. . . . . . . . . . . . . . . . 647
Author Index 683
Subject Index 708
List of Contributors
ADES, Harlow W., Department of Electrical Engineering, Bioacoustics Research Laboratory,
University of Illinois, Urbana, Illinois 61801, USA
AXELSSON, Alf, University of Washington, Department of Otolaryngology, BB 1165 RL·30,
Seattle, Washington 98195, USA
BAIRD, Irwin, Department of Anatomy, The Milton S. Hershey Medical Center, Pennsylvania
State University, Hershey, Pennsylvania 17033, USA
t BEKEs¥, Georg von, Laboratory of Sensory Sciences, University of Hawaii, Honolulu,
Hawaii 96822, USA
BOORD, Robert, Department of Biological Sciences, University of Delaware, Newark, Delaware
19711, USA
CAMPBELL, C. B. G., Department of Neurological Surgery, University of Virginia, Charlottes·
ville, Virginia 22901, USA
DENSERT, Ove, Department of Otolarynology, Karolinska Sjukhuset, S·10401 Stockholm 60,
Sweden
ELDREDGE, Donald H., Central Institute for the Deaf, 818 South Euclid, St. Louis, Missouri
63110, USA
ENGSTROM, Hans, Department of Oto.Rhino-Laryngology, Uppsala University, University
Hospital, S-75014 Uppsala, Sweden
FEX, Jargen, Laboratory of Neuro-otolaryngology, National Institute of Neurological Diseases
and Stroke, National Institutes of Health, Bethesda, Maryland 20014, USA
HARRISON, J. M., Department of Psychology, Boston University, Boston, Massachusetts
02215, USA
HENSON, Jr., Odell, W., The University of North Carolina at Chapel Hill, Division of Health
Affairs, Department of Anatomy, 306 Mac Nider, Chapel Hill, North Carolina 27514, USA
HOWE, M. E., New England Regional Primate Research Center, Harvard Medical School,
Southborough, Massachusetts, USA
IURATO, Salvatore, Cattedra di Bioacustica dell' Universita di Bari, Policlinico, 70124 Bari,
Italia
MICHELSEN, Axel, Institute of Biology, Odense Universitet, 5000 Odense, Denmark
MOLLER, Aage R., Fysiologiska Institutionen II, Karolinska Institutet, 10401 Stockholm 60,
Sweden
PFEIFFER, Russel R., Department of Electrical Engineering and Physiology and Biophysics,
The Washington University, St. Louis, Missouri 63130, USA
RAUCH, S. and 1., 3011 Bern, ZeughausstraBe 20, Switzerland
SHAW, Edgar A. G., Acoustic Section, Division of Physics, National Research Council
Canada, Ottawa, Ontario KIA OR6, Canada
WERSALL, Jan, Department of E. N. T., Huddinge sjukhus, 14186 Huddinge, Sweden
WEVER, E. G., Auditory Research Laboratories, Princeton University, Princeton, New Jersey
08540, USA
Chapter 1
Introduction
GEORG VON BEKESY t, Honolulu, Hawaii (USA)
With 3 Figures
Contents
Historical Background of Research in Hearing . . . 1
Impact of Physical and Telephone Engineering Concepts on Research in Hearing . 3
The Impact of Instrumentation on Hearing Research 5
The Data Explosion . 6
References . . . . . . . . . . . . . . . . . . . 7
Since scientific progress is not a continuous flow, the best time to write a
handbook is when the number of findings reaches a maximum and the newer
experiments show a certain redundancy. VON HELMHOLTZ'S (1863) book on hearing
is a classical example of a book published at the right time. But so also is the
handbook published by BETHE and BERGMANN (1928). Perhaps the publisher of
these three volumes is correct in his judgment that we have reached such a maximum
and tha,t now is the correct time to summarize the data. This is a brave enterprise
of the publisher, the editors, and the authors; their efforts will be highly apprecia
ted by the coming generation. The material in the two volumes on audition will
form the platform from which new advances can be made, as was the case with
the books of VON HELMHOLTZ and BETHE and BERGMANN.
In engineering and the exact sciences, a new handbook would mean essentially
the addition of some new facts to the already acquired ones. Unfortunately, this
is not so in the field of hearing. Hearing research at the moment is a complicated
interaction between physics, anatomy, physiology, and psychology. We cannot
separate certain variables to the degree that is possible in physics. Furthermore,
our measurements are not so precise, and the range of validity is not so well
defined. Therefore, we often have to modify our earlier findings in light of the new,
at least in the range of validity. In this way, a new handbook in the field of hearing
is almost always a reevaluation of all the known facts. It is this balancing of one
finding against another that makes modern handbook writing difficult.
Historical Background of Research in Hearing
The main basic problem in hearing today is almost exactly the same as it was
a century ago. Perhaps it can be illustrated best by an experiment. If we have a
2 G. VON BEKESY: Introduction
speaker in a normal living room and we listen to him monaurally or binaurally
first from a distance of one meter and then from three meters, we notice hardly
any difference except for a small drop in loudness at the greater distance. But if
we have two identical microphones, one placed one meter away from the speaker
and the second three meters away, then the recordings show two different sound
pressure patterns over time as illustrated in Fig. 1. There is a small time delay
for the lower trace which was recorded from the more distant microphone. It is
difficult to understand how such different stimuli as the sound patterns in the
upper and lower traces can produce the same sensations. Much research was done
to find the reasons why this is possible.
We inherited from the last century three basic experiments. One was done
by SAVART (1840), one by OHM (1843; 1844), and one by SEEBECK (1841, 1843,
1844a, b). All three experiments were repeated over and over again and proved
to be correct. In recent years, the validity range was also precisely determined.
SAVART found that the pitch of a tone, with or without overtones, is determined
with the precision of 10% in two cycles. Since resonators or small band pass
filters have to accumulate energy to produce a frequency analysis, the ear cannot
possibly consist of such frequency analyzers, since two cycles do not permit the
necessary accumulation of energy. Therefore, the short onset and offset time of
pitch discrimination of the ear contradicts the resonance theories of hearing. The
above experiment is so simple that it did not become controversial and, therefore,
it is seldom quoted.
OHM introduced the FOURIER analysis into the field of hearing, showing that
li>uch a complicated pressure pattern as speech can be represented as the sum of
many sinusoidal tones. Since sinusoidal electric currents could be easily measured,
OHM'S law became the basis of modern telecommunication planning. When com
bined with the fact that phase differences between two tones cannot be detected
Fig. 1. Section of,a recording of speech by two condenser microphones, one, I m away from the
speaker (upper recording) and, another, 3 m away (lower recording); both of them in the median
plane of the head and spoken toward the direction of the microphones. There is a small time
delay between the two recordings as a result of the traveling time of the s01md waves; but,
otherwise, they would be similar if there would be no sound reflections from the wall of the
ordinary living room. The problem of hearing research is that listening monaurally or binau·
rally to both sounds, which show such a strong difference in the oscillographic recording, we
are unable to detect by hearing a difference except the change in loudness in agreement with
the distance increase
Impact of Physical and Telephone Engineering Concepts on Research in Hearing 3
with a single ear and, therefore, phase measurements can be omitted, OHM'S law
made possible a quantitative description of communication systems. It was known,
however, that OHM'S law is not valid in all situations. A FOURIER analysis of a
complex tone overemphasizes the steady state and is of little use for transients.
SEE:aECK found that for speech and for certain complex sounds which show a
definite periodicity, the ear hears tones which do not show in the FOURIER analysis
of these complex sounds. In other words, the ear does not do a long-term frequency
analysis; its analysis is made in a few cycles.
All three of these basic experiments indicate that oscillographic recordings,
as shown in Fig. 1, are not good speech extractors.
Impact of Physical and Telephone Engineering Concepts
on Research in Hearing
Assuming that, during the evolution of the animal kingdom, the most efficient
system always survived, it seems probable that the laws of physics were followed
to build up maximal efficiency. Therefore, we expect that the physical laws served
as guidelines to the evolution of the structures and functions of the middle and
inner ear. The sensitivity of the ear is so great that we know nature has outdone
man in our attempts to build sound transmission systems based on these same
physical laws. For example, to obtain maximum sensitivity in hearing, none of
the sound energy hitting the eardrum should be reflected but should be transmitted
in its full amount to the inner part of the ear, the cochlea. THEVENIN (1883) has
shown that maximum sensitivity can be obtained by means of a mechanical trans
former which provides a proper matching between the relatively soft air of the
middle ear and the much harder fluid of the inner ear. THEVENIN'S theorem, which
is so widely used by man in telephone engineering, also helps to explain the struc
tures and functions that nature has provided for the middle ear.
It is relatively easy to measure the electrical activity inside the cochlea and the
energy of this activity for a given part of the cochlea. It turned out that the energy
activity inside the cochlea produced by a tone is very much higher than the energy
of the tone which is absorbed by the eardrum. Therefore, in the inner ear we do not
have a passive system but something like an amplifier. ]'rom telephone engineer
ing, it is known that it is of no use to amplify indefinitely because every amplifier
also produces noise. This holds for the human ear where the cochlea has a blood
supply with pulse waves resulting from the streaming of the fluid in the capillaries.
One of the most important technical concepts, therefore, became the so-called
signal-to-noise ratio. Nature applied this concept by reducing the sensitivity of
the ear for the lower frequency range where noise is introduced by the circulatory
system.
One of the principles which had a tremendous impact on hearing research was
the Fourier analysis. The Fourier analysis, as mentioned above, showed that any
complex sound can 1:>e considered as a sum of sinusoidal sounds. Since the ear does
perform to a certairi degree exactly that type of analysis, application of Fourier
analysis became one of the most successful tools in hearing and speech research.
Pitch discrimination seemed to be simply explained by assuming for every fre-
4 G. VON BEKESY: Introduction
quency a certain resonator which produced sensations only in that frequency
dominion.
However, there were two difficulties. One was that no proper resonator could
be found inside the cochlea. Most of the tissue in the cochlea is either too soft or
too hard to function as a resonator. The other difficulty was that there is a well
established law in telephone engineering that a filter with a given frequency band
width needs given onset and offset times. This difficulty was already known a
century ago, long before it was possible to formulate it mathematically. It is the
short onset time of the perception of speech which makes it unlikely that resona
tors inside the inner ear determine pitch perception. In this case, the application
of Fourier analysis was not successful.
But there are other principles, like the feedback principle in the auditory
system, which are only beginning to be developed and which will contribute to
the further understanding of cochlea activity. Pattern recognition similar to that
found in vision certainly plays a great role in the field of hearing; but, it will take
a long time before it can be evaluated. In vision, it is already clear that the visual
sensations are produced mainly by edges in the brightness distribution and not by
the brightness itself. The importance of similar phenomena in the auditory system
is only beginning to be investigated.
Compared with machines, one of the most surprising features of sense organs
is the tremendous range of physical events to which a given sense organ is sensitive.
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