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potentials
Peter Dallos, Mary Ann Cheatham, and John Ferraro
Auditory ResearchLaboratory,Departmentof ElectricalEngineering,* NorthwesternUniversity,Evanston,
Illinois
60201
INTRODUCTION
of CM data diffi-
cult, and the development of one-to-one relationships between CM and the motion of one point of the basilar membrane, or CM and the response of a single auditory
nerve
fiber
hazardous.
These
two factors
are
the com-
behavior
neural events.
mechanical
or
siderable attention during the past several years (Whitfield and Ross, 1965; Weiss et at., 1971; Dallos et at.,
1972; Haas,
membraneous cochlea.
It has also
597
J.Acoust.
Soc.Am.,Vol.55,No.3, March1974
METHOD
which
the CM was
eliminated.
The
details
of ani-
constant
then
middle
ear
influences
are
eliminated
Copyright
1974bytheAcoustical
Society
of America
597
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598
598
Middle
fective input to the cochlea is pressure at the oval window, and this pressure is proportional to the product of
stapes velocity and cochlear input impedance. In the
guinea pig the acoustic input impedance of the cochlea
is resistive and constant at all significant audio frequen-
transfer
SPL
ear
function
correction
SPL
correction
displacement
SPL
SPL
400
400
400
frequency (Hz)
FIG. 1.
are
do.
two sets of available
direct
measurements
their
use.
The
main
characteristic
of both of
these functions is that when sound pressure is held constant at the eardrum the stapes displacement remains
constant up to about 400 Hz, beyond which it decreases
at an approximate
rate of 8-9 riB/octave.
Thereare
some resonant peaks superimposed on this general pattern, but presumably the magnitude and frequency of
such peaks is highly variable from animal to animal
of dB re 1 .
II.
RESULTS
AND
DISCUSSION
Von Bksy's (1960, pp. 680-682) classic experiments utilizing the vibrating electrode provided the first
direct indication of the relationship between basilar
membrane motion and cochlear potentials.
He demonstrated that a trapezoidal displacement of the cochlear
partition produced a trapezoid-shaped microphonic response. The conclusion that can be drawn from these
experiments, now widely accepted as one of the .funda-
1972b; Dallos,
1973). This modified description of the functional relation between microphonic time patterns and basilar
membrane motion merely extends yon Bksy's observations and is in no way in conflict with them. To
explain, consider that the electrical output of the outer
hair cells is considerably in excess of what is generated
by the inner hair cells. This has been demonstrated by
velocity plots are sought then the input sound level is de-
creased at a rate of 6 riB/octave up to 400 Hz and increased at a rate of 2 riB/octave above this corner fre-
quency.
One can glean from the data of both groups of investigators that at low frequencies the rms amplitude of
rejected. The insert in the figure shows the cochleogram of the abnormal animal whose CM function is presented. This plot demonstrates that in the neighbor-
8000-Hz
elec-
599
Dallas,Cheatham,
andFerraro:
Cochlear
mechanics
andpotentials
i
100i--::.-.-.-.-.-.-.-.-.-
,,-;,-,-':-':.
-
! i
.
i i
i i
1973).
i
- 5
OHC
zo
I0
so
15
DISTANCE
FROMAPEX
1
,20
f =8000Hz
2
SPL
E[G. 2.
Input-output [unctons [o C
ioo
12o
(dB)
at 8000 Hz :eco:ded
[:om the fi:st turn of the cochlea. Heavy filled c:cles :ep:esent data [:om one andrea[whose coch[ea ws poisoned by
Eanamycn njecHons. The coch[eog:am of ths andrea[ s
shown n the nse:t. ote that n the vcty of the :eco:dng
e[ect:ode pa: (a::ow) on[y nne: ha[: ceUs ([HC) a:e p:esent.
The nput-output [uncton :ep:esented by [gh[ symbols s a
median no:m[ p[ot (a[ongwth nte:qua:t[e :ae).
hair
cells
are
about
It is a reasonable
recorded by the difby the inner hair
conclude that the
30 dB more
sensitive
than the
one-tenth
of the
contribution
of the outer
hair
stapesdisplacement
of 10 ., andat the approximate
stapesvelocityof 25000,/sec. Whensoundpressure
is kept constant, the CM magnitude rises at an approxi-
cations.
300
that
lo-
coMtant
stapes
displacement
I00
"'30-
:o -
stapes
velocity
TI
t,
o.,
o.s
Frequency
such cancellation
constant
3-
the contrast
electrode
cells.
Thus the CM generated by the inner hair cells has negligible influence on the total, normal, response. We can
thus say, in harmony with yon B&k&sy, that in a normal
cochlea (dominated by outer hair cells) the CM is proportional with basilar membrane displacement.
Another
conclusion that can be drawn from the plots of Fig. 2 is
that schemes that attempt to explain the declining segment of the CM input-output function by postulating a
cancellation between the CMs produced by the two cell
20 I 0 I 6 I 8 I
599
FIG. 3.
constant
SPL
,o
(kHz)
10 andvelocityof 25000k/sec.
600
Dallos,Cheatham,and Ferraro:Cochlearmechanics
andpotentials
Basilor
membrane
displacement
20
io
ments
.--
"'
CM at constant
stapes
displacement
at 4.0
-2:0',
'
Ol
0.2
FIG.
4.
'
'
'
'
'
'
I0
20
50
Comparison
ear
structures
and basilar
mem-
mm
0.5
Frequency
of both middle
.-I0
600
(kHz)
stapesdisplacement
(1 .) from one of our guineapigswith
basilar membrane displacement divided by incus displacement
as measured in a guinea pig by Wilson and Johnstone (1972).
In both experiments the point of measurement is approximately
the same. The ordinate is arbitrary logarithmic scale.
cells
distributed
over
a considerable
extent
of the inner-
trodes. Since the mechanical resonance is highly localized and since the electrical recording is not, it is
reasonable to expect that the latter should reflect a
spatial average and thus to result in a flatter function.
A similar argument can explain the discrepancy between
the high-frequency slopes of the CM and the basilar
membrane
functions.
membrane
set
to incus displacement.
of mechanical
data
was
chosen
This parbecause
it is
tions
in both the CM
are
similar
below
motion
18000
Hz.
func-
At
The major
(Dallos and Durrant, 1972) that this expectation is largely borne out. It was shown that with a triangular time
pattern of stapes displacement the electrical response
clearly approximates a square-wave, indicating the
fundamental validity of the assumption that CM is proportional to stapes velocity.
An interesting observation
was the approximately 0.9-msec time delay between the
third turn and the first turn square-wave responses. The
delay corresponds to the travel time that a disturbance
in response magnitude around the characteristic frequency is not matched by the CM. While there is an
must
take
to move
between
locations
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601
T2
I T
TI
30
ioooI_40
40
Not
30
20
300 - 20
I
601
observations
" IOO
that
can be made
about
these
families
of
3o
I0-
O.2 0.
'r
O.5
I0
20
Frequency
FIG.
5.
{ kHz)
theparameterin dBre I
This square displacement wave travels along the cochlear partition producing the square-wave-like
CM,
stapes of the chinchilla is moved by a triangular displacement, the resulting CM is not a pure square wave
but it possesses pronounced impulse components at the
transitions. We have suggested some years ago that
these discrepancies simply reflect the varied influence
lower frequencies, the bunching of the graphs immediately above the frequency of the low-level peak, and actual
decreases
in response
withincreasedinput-againjust
above the original peak. The pronounced shape changes
in the plots with intensity indicate that only at the lowest
stimulus levels can one presume that nonlinear effects
negligibly influence the CM response. Some departures
fromlinea.rityoccuraboveapproximately
1 . of stapes
displacement.' This meansthat whentranslatedinto the
more familiar sound-pressure measure, one can not
expect truly linear CM response from the third cochlear
turn above 30 dB SPL, from the second above 55 dB,
and from the first above 75 dB. Early onset of nonlinearity, however, occurs in a rather limited frequency region: just above the best frequency of the electrode location. Far distant from this region the response remains markedly linear up to extremely high driving
that indicate
that except
being a linear.response.
plots obtained from the basal turn of one guinea pig are
is manifested by the decline of the low-frequency response and the departure from the rule that the CM is
proportional to stapes velocity.
Above
frequency
plotsare shownfor three electrodelocations.
As a means
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602
Dallos,Cheatham,
andFerraro:Cochlearmechanics
andpotentials
1.0
602
their primaries then all orders of harmonics and combination tones peak at the same frequency, which depends primarily on the location of the recording elec-
-I0,0
I0
20
50
ment of 10 from three electrode locations. The fundamental plots show the now familiar steady growth toward
a peak and the subsequent rapid decline. The animals
were chosen to be representative, thus the differences
in magnitudes from turn-to-turn that are seen here are
a reasonably consistent finding. Accordingly the peak
response tends to be the greatest in the second turn and
0.01
0.1
02 0.
0.5
I0
Frequency
20
(kHz)
stapesdisplacement
in dBre I ). Plotsarefromoneindividual guinea pig, recording is from the basal turn. In the lower
plot a "degree of nonlinearity" function is presented. This
function is defined by the ratios of the normalized CM mag-
smallest
in the first
turn.
The second
harmonic
func-
of quantifyingthese contentionswe now introducea "degree of nonlinearity function." This function is obtained
by forming the ratio between the normalized magnitudes
of any two CM plots. For convenience we choose the
ratio betweennormalizedCM at 1 and 10 stapesdisplacements and present the resulting function on the bottom of Fig. 6. A value of one indicates linear response,
and the more nonlinear the CM becomes the larger the
numerical value of the function. It is apparent that we
have a rather sharply peaked plot whose maximum occurs at 14000 Hz, somewhat above the peak of the lowlevel CM .function. Highly similar functions can be obtained
from
the other
cochlear
The fact
frequency region than the primaries; to state it differently, they show sharper tuning. This suggests that the
distortion component amplitude is not a simple function
of the magnitude of the primary CM, or of the input
stapes displacement.
It is worth reemphasizing that
all orders of distortion
turns.
T$
T2
It is tempting to associate the pronounced frequencydependent nonlinearity of the CM with the somewhat simi-
cesses, that is, from distortion tied in with the transducer mechanism as opposed to hydraulic or mechanical
in the same
manner.
This
behavior
,,
,,
i
i
can be summarized
O.
0.1
I
0.
0.5
,5
Frequency
13
I0
20
(kHz)
animals)as thefunction
of frequency
at a constant
10 stapes
displacement. Both fundamental (solid symbols) and second
harmonic components (open symbols) are shown. The second
harmonic functions are plotted at the frequency of the fundamental.
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603
Dallos,Cheatham,
andFerraro:Cochlear
mechanics
andpotentials
i
3o
AVE(10.)
603
ti0n. Thus while the absolute magnitude of the distortion component is greatest at the frequency where the
fundamental CM is maximum, the relative distortion is
most pronounced where the degree of nonlinearity is the
largest.
"'
SECOND
TURN
.I
.-.
These peaks are clearly better correlated with the frequency region of greatest nonlinearity than with the
fundamental
CM
maximum.
oJ- I \ !!
%2fo
(10,)-j3o
.
{ ?_fj,.)'
'1I i
The final nonlinear phenomenon in need of a brief discussion is the elusive "interference
effect."
First noted
i0 .
(")
3 '
tone
-30
is diminished
when
a second
tone
is introduced.
It
10 stapesdisplacement
as thefunction
of frequency.Both
0.31
I
, ,, &,,,
2
, I,
, ,,11
5
Frequency (kHz)
-I00
I
2 3
I I
5 7
The vertical
interrupted
ratioof thenormalized
CMmagnitude
at i and10 stapesdisplacements,
andthepercent
second
harmonic
content
at 10
stapes displacement.
constant
10 stapesdisplacement.All measurements
are from
the same
animal.
that interference
stapesdisplacement
of 1 .
The interferingtone'sfre-
stantstapesdisplacement
of 10 ,. The ordinateshows
fundamentalfrequency. To aid in demonstratingthe difference and [o provide a summary of much of our previous discussion Fig. 8 is presented. In this figure various plots, all obtained from one animal, are given. In
the left panel, fundamental and second harmonic magni-
10 . fundamental
CM graphshowsthe signsof marked
nonlinear influences. The peak shifts down to 1250 Hz,
and a dip develops at 3500 Hz. In the center panel the
degree of nonlinearity function clearly indicates that the
most nonlinear frequency region is between 3000 and
3500 Hz. This region is above the low-intensity fundamental peak of 2500 Hz. Also included in the center
panel is a plot showing the percent second harmonic dis-
tortionat 10 . stapesdisplacement. This plot hasvirtually the same shape as the degree of nonlinearity func-
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604
Dallos,Cheatham,
and Ferraro:Cochlearmechanics
andpotentials
25
20
'E
0.3
604
0.03
I0
20
left panel:
I0
15
FIG. 9.
20
(kHz)
to a 7000-Hz(arrow)tonepresented
at I stapesdisplacement
level as the functionof thdfrequencyof an interferingtonewhich
is presented
at a constant
stapesdisplacement
of 10. Recording is from the first turn; right panel: Combined data
from three animals showing the relation between the frequency
of a tone, maximaltonalinterferencewith its CM occurs above the best frequency associated with the location of the recording electrode, in other words in the
aforementioned maximally nonlinear frequency band.
ACKNOWLEDGMENT
quencies
presented
at constant
I , andinterfering
frequencies
at constant
10 stapesdisplacement..
*It is likely that above approximately 20 kHz the cutoff rate
becomes considerably greater.
ly linear rule. The mechanism underlying this relationship is not at all evident, it is, however, quite clear
that effective interference
tone is within a region where our "degree of nonlinearity" function is large. It is interesting to note that we
never see significant interference below the best frequency, thus we can not confirm the existence of the
double-lobed
interference
functions
III.
location
is thus a well
demonstrated
fact.
SUMMARY
from
normal
cochleas
is overwhelm-
37-46.
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605
Dallos,
Cheatham,
andFerraro:
Cochlear
mechanics
andpotentials
Polarization
of the Cochlear
Partition
on the Sum-
mating Potentials,"
J. Acoust. Soc. Am. 52, 542-552.
Durrant, J., and Dallos, P. (1972b). "The Effects of dc
Current Polarization on Cochlear Harmonics,"
J. Acoust.
Soc. Am. 52, 1725-1728.
Durrant, J., and Dallos, P. (1974). "Modification of DIF
Summating-Potential Components by Stimulus Biasing, "J.
Acoust. Soc. Am. (to be published).
Engebretson, A.M.,
of Cochlear
Potentials,"
Guinan, J., and Peake, W. T. (1967). "Middle Ear Characteristics of Anesthetized Cats," J. Acoust. Soc. Am. 41,
1237-1261.
Hair Cells,"
Legouix, J.P.,
"Interference
Am.
53,
J. Acoust. Soc.
409-419.
605
Wang, C. Y.,
Wever, E.G.,
Whitfield, I. C., and Ross, H. F. (1965). "Cochlear Microphonic and Summating Potentials and the Outputs of Individual
Generators,"
J. Acoust. Soc. Am. 38, 126-131.
Wilson, J.P.,
and Johnstone, J. R. (1972). "Capacitive
Probe Measures of Basilar Membrane Vibration,"
in Hearing Theory (Institute for Perception Research, Eindhoven),
Hair-cell
172-181.
Northwestern
1818-1830.
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