Compound action potential (AP) tuning curves*

Peter Dallos and Mary Ann Cheatham

AuditoryPhysiology
Laboratory(Audiology)and Departmentof ElectricalEngineering,Northwestern
University,Evanston,Illinois 60201
(Received 18 September1975; revised 17 November 1975)
With a tone-on-tonemaskingprocedurethe compoundaction potential (AP), elicited by brief tone bursts
of set frequencyand intensity,was decreasedby a constantfraction. The frequency-intensitypairs formed
by the maskerthat yield this decreasegeneratethe AP tuningcurve.It is demonstrated
that suchtuning
curvesare very similar to both psychophysical
tuning curvesand singleVIIIth-nerve-fiber tuning curves.
Changesin the propertiesof thesecurvesare describedas functionsof stimulusfrequencyand level, mode
of masking(simultaneousand forward), and parametersof the masker.

SubjectClassification:[43]65.40,[43]65.42, [43]65.58.

INTRODUCTION

The compoundaction potential (AP) of the auditory

nerve

is known

to be diminished

when

noise

is added

to the stimulus which elicits the AP (Derbyshire and

Davis, 1935). This decrease, in analogy with the psychophysical process, is commonly referred to as maskingo The most frequently used masker in AP studies
has been wide-band noise, although narrow-band noise
maskers

have been

utilized

to determine

the cochlear

segmentscontributingto the total AP (Teas et al.,

1962; Bone et al., 1972). The usual masking method
is to present a continuous background noise, even
though forward masking of the AP has been demonstrated

over ten years ago (Coats, 1964). Explorations of

simultaneous and forward masking effects have been

provided by Spoor (1965), Spoor and Eggermont (1971),

Eggermontand Spoor (1973), and Eggermont and Odenthai (1974). Extremely limited work has been done
on tone-on-tone AP masking (Kupperman, 1971; Spoor
and Eggermont, 1971).
The AP was classically considered to be the response
to clicks

or to the wide-band

onset

transient

in tone

bursts, and thus to be made up from the contributions

of fibers originating in the basal turn of the cochlea

(Tasaki, 1954; Kiang et al., 1965; Dallos, 1973). It

is now fairly clear that carefully shaped tone bursts
of low or moderate intensity can activate limited cochlear segments, appropriate to their frequency, and thus

generatefrequen..cy-specific
AP responses
(Teas, 1966;
Mitchell, 1973; Ozdamar, 1973;Eggermont and Oden-

thai, 1974;de Boer, 1975;zdamar andDallos, 1976).

In psychophysics one of the best means of assessing
frequency selectivity can be obtained with tone-on-tone
masking procedures. Particularly interesting among
such procedures is the generation of so-called psycho-

physical tuning curves (Chistovitch, 1957; Small,

1959: Zwicker, 1974; Vogten, 1974). These curves
are considered analogous to single-unit tuning characteristics, and thus they provide a provocative link between psychoacoustic and physiological measures of

cochlear frequency selectivity.

Inasmuch as tuning curves are finding increased use
in human psychophysics, as noted above, and have been

introducedin animal psychophysics(McGee et al.,

1975; Dallos and Ryan, 1975), we considered it fruitful to explore this measure in the compound response

591

J. Acoust.Soc.Am.,Vol. 59, No. 3, March1976

of the auditory nerve. An additional motive for this

work was to provide new information on tone-on-tone
masking of whole-nerve action potentials.
I. METHODS

All reported data are based on measurements on

younganesthetisedguineapigs (anesthetic: 1600 mg/kg

urethane, or 800mg/kgurethaneplus64mg/kg ketamine
initially with additional ketamine doses when needed).
In all animals, tungsten electrodes were placed in
scalae vestibuli and tympani of the first cochlear turn
and the AP was derived from the summed output of
the two. In all preparation, the tensor tympani tendon
was severed, and the bulla was open during data collection. Sound stimuli were generated in a closed sys1

tern by a -in.

condenser microphone

nonlinearity-compensating

eta[.

driven from the

circuit described by Molnar

(1968). Soundpressure was measured with a

calibrated probe-tube microphone near the eardrum.

Aside from accurately establishing the sound-pressure

levels available throughout the frequency range of interest, baseline measurements, consisting of microphonic magnitude and phase determinations and AP
thresholds, were also routinely performed on all animals. Only subjects that showed normal CM and AP
patterns were utilized in this study.
All AP measurements were carried out by averaging
from 8 to 256 responses. While the summed output
of the two electrodes contains very little CM, this
potential was further eliminated by presenting the
stimulus tone bursts in random phase and thus averaging out the residual ac receptor potential.
The stimulus
used to elicit the AP was always a short tone burst

(usually 15 msec in duration)with linear-ramp rise

and fall patterns of 1 msec each. This stimulus structure generates a well-developed AP at all frequencies.
While the 1-msec rise-fall times are sffficiently long
to eliminate spectral spread above 4 kHz, such spectral smearing does undoubtedly occur at the low frequencies. The consequence of such is that as energy

spreads to other than the signal frequency a broader

segment of the organ of Corti could be excited;
the tone-on-tone masking procedure Could yield
poorer index of frequency selectivity than what
be obtained in the absence of spectral spread.

thus
a
could
Another
factor that influences the results at low frequencies

Copyright
1976by the Acoustical
Societyof America

591

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592

P. Dallos and M. A. Cheatham: Compound AP tuning curves

STIMULUS

592

+2 dB/octabove400 HZo The rationalefor describing

MASKER

input quantities in terms of stapes velocity instead of

(a)

sound-pressure

level is elaborated

in ])allos el al.

(1974).
II.

RESULTS

AND

DISCUSSION

All data reported in this article are expressed in

terms of the masker level required to yield a predetermined reduction in the magnitude of the AP. In
practice, the averaged AP response was displayed on
an oscilloscope screen, adjusted to a convenient size;
then the masker level was changed until the displayed
averaged response was reduced to the desired fraction
of the response without masking. Since only relative
measures were thus utilized, the electrical signal was
prefiltered in various ways to provide stable baseline
readings. The AP was always expressed as the differ-

(b)

(c)

ence between baseline (level prior to stimulus delivery)

and the N peak.

A. Tuning curves:simultaneousmasking
FIG. 1.

Time patterns of probes and maskers used in this

study: (a) Continuousmasker; stimulus tone bursts are 15

mse lon and are separated by 300 mse. Stimulus rise-

f11 time: 1 mse. (b) Pulsed simultaneous masking. Tone

bursts are overlapped by the masker, whose duration is
mseo Silent interval:

300 mse; 11 rise-all times:

mseo () Forward masking. Masker duration (TM) and

masker-probe interval (T) are variable probe 15 mse
long silent interval

300 mse. All rise-fll

times are

msec.

Many of our data were collected with the stimulus,

or probe, tone 10 to 15 dB above AP threshold. In
such cases, the most effective masker frequency is
almost invariably the same as the probe frequency, and
for any degree of masking the absolute level of the
masker is lower than that of the probe. How much
lower depends on the amount of reduction one wishes
to specify. Thus, for example, for our most often used

probe frequency (f0) of 8 kHz, and a reduction of the

N[ by one-third,
stems from the effects of randomized

of the stimulus.

starting phase

Wang (1971) has shown that below

2 kHz the starting phase influences the magnitude of

the Nz, and that phase-locked signals starting either

with 0 (rarefaction) or r (condensation) phases yield

larger action potentials than the ones with random
phase. The reduced amplitude and inherent time smear
of action potentials generated by random phase signals
does potentially reduce the expected sharpness of tuning at low frequencies. Thus, it is likely that because
of both spectral and time effects, our low-frequency
results will underestimate the frequency selectivity of
low-frequency AP. Three types of masking stimulus
were utilized: continuous, pulsed simultaneous, and

forward maskingstimuli. The three paradigmsare

schematized in Fig. 1.

The only variables were the

masker duration (TM) and the time interval (AT) between masker and signal; both ranged from 5 to 1000
msec. The OFF time was kept at 300 msec.
Unless specifically noted, all stimulus and masker
level designations are expressed in decibels relative

to a specified stapes velocily. The reference is 1 nm/

sec (10 i/sec).

sound-pressure

Stapesvelocitiesare established
from
measurements with the aid of an ideal-

ized guineapig middle-ear transfer function[described

in Dallos el al. (1974) and based on the data of John-

stoneand Taylor (1971)]. The correction factors for

obtainingthe equivalentSPL (decibelsre 20 /N/m2)

level

and effective

the median difference between probe

masker

level

is 18 dB.

On either

side of f0 the required masker level rapidly increases.

At f0 = 8 kHz, the medianhigh-frequencyslope is 90 dB/

oct, and the low-frequency slope tends to be somewhat

shallower. At approximately an octave below f0 the

function flattens out, and the difference between masker level at f0 and at the plateau is about 30 dB+ 10 dB;
the lower the probe level, the greater this difference
tends to be.

The plots shown in Fig. 2 provide a picture of the

most common features of the tuning curves; they also
illustrate the effect upon these curves of the criterion
suppression level. To explain, the two plots shown
were obtained by plotting the masker levels required
for depressing the AP by one-third and for complete
elimination of the response. The overall shape of the
two curves is similar, there is a vertical shift in excess of 10 dB between them, and it can be noted that
the upper curve, which represents the complete masking situation, is sharper than the one belonging to the

partial masking case. Inasmuch as it is hard to judge

the level required for complete masking, many of the
data reported here were collected with a partial masking
criterion; either one-third or one-half down. The latter criterion would produce a curve fitting between the
two plots of Fig. 2.

from the velocity figures reported here are -? dB at

It is apparent that these AP tuning curves possess the

same features as the well-known single-unit tuning
curves, or the psychophysical tuning curves. There

400 Hz, -7 dB+6 dB/oct below 400 Hz, and -? dB

is a relatively sharp tip segment, an elongated low-fre-

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P. Dallos and M. A. Cheatham: Compound AP tuning curves

593

593

8O

8 -

7O

6 -

60
o

50

ao-

8
0.2

0.5

10

20

Signol
,-

(I) O0

cD

Frequency

(kHz)

FIG. 4. Individual Qt0 values obtained from several preparations with simultaneous masking. For comparison, we show
the median single unit Qt0 for guinea pigs deduced from the

30

work of Evans and Wilson

(1973).

20-

0.5

Mosker

I III

.5

Frequency

tween the plateau and the lowest point on the tip) also

I0

tends to be greater in single unit plots.

20

(kHz)

FIG. 2. Masker stapesvelocity (decibels .e 1 nm/sec) versus

masker frequency necessary to decrease the magnitude of an

Art response elicited at 10 kHz, 15 dB above AP threshold, by

(o-o), andto eliminatethe AP altogether(-e).

In this

and succeeding figures the hexagonal symbol signifies the frequency and level of the probe tone.

quency tail, and no masking effect at high frequencies

The major difference between AP and single-unit tuning
curves is the sharpness of tuning. Thus, while the
general features are similar, the single-unit curves
are more sharply tuned. The vertical extent of the

high-frequency tip segment (that is, the difference be-

The well-known graduation in sharpness of tuning

that is reflected in basilar membrane vibration patterns

(Tonndorf and Khanna, 1968) and in single-unit tuning

curves (Kiang et al., 1965; Evans and Wilson, 1973)
is apparently represented in the AP plots as well.
This frequency dependence is further highlighted in
Figs 3 and 4. In the first of these figures, six tuning
curves obtained from a single animal at different frequencies-all with the probe 15 dB above AP threshold-are shown.

There is a clear-cut

gradation in the

sharpness of tuning, and a conversion from the simple

V-shaped pattern to an asymmetrical one, as one progresses up the frequency scale. The two highest-frequency plots demonstrate the sometimes observed
slight detuning of the curves, whereby the lowest point
is found either somewhat below or above the probe

go-

frequency. One popular measure of the sharpness of

tuning is the Q0 value, numerically obtained as the

ratio of the centerfrequencyand the bandwidth

mea-

7oo_
....-o.
"-

/'
"--o /

,'/'l,

"-/
/

!.1

'-.

sured 10 dB above the lowest point. In Fig. 4, the Q0

computed from numerous tuning curves is compared
with the median plot that can be gleaned from the data

,;/

presented by Evans and Wilson (1973) for single-unit

' 1

tuning curves from guinea pigs. In both types of data,

there is a hint that sharpest tuning is obtained around
8 kHz, with significant flattening toward the low frequencies. The numerical values of AP Q0's are about
one-third to one-half of those of the corresponding
single-unit curves.

60

4o

/ --

11 --1

I
All

3o

.2

0.5

Masker

/ q ;

I0

20

Frequency (kHz)

FIG. 3. Family of AP tung cues at different probe frequencies, all at 15 dB ave AP threshold. Ordinate is stape
velociW of masker; abscissa is masker frequency. Hexon
symls ave tung cue tips show level and frequency of
probe; numbers ne tips give 0 vues.

the data

shown

thus far

were

obtained

with

the

probe slightly above the AP threshold. The question

arises as to how, if at all, the tuning curves change
as the probe level is raised. Figures 5 and 6 address
this issue. In Fig. 5, a detailed view is provided of
the tip segment of five tuning curves obtained at 10-dB
increments of the probe level. It is notable that the
first four plots largely parallel one another without
either a significant broadening or sharpening. The top
graph appears shallower; in fact, its Q0 is 2.8 as opposed to the average of 3.3 for the other four. More
impressive than this broadening at the highest levels
is the difference in spacing between the four lowest

J. Acoust. Soc. Am., Vol. 59, No. 3, March 1976

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594

P. Dallos and M. A. Cheatham: Compound AP tuning curves

594

6 of Eggermont and Spoor (1973). They also noted the

relationship between the different rates of masking and
the two segments of the AP magnitude functions. These
relationships are relatively easy to explain in light of

90

/e (2.8)

our recent suggestion()zdamar andDallos, 1976) that

the low-level segment of the AP magnitude function is

70-

to be associatedwith the dischargesof a relatively lo-

',,

calized group of neurons, operating within the tip segment of their tuning curves, while the high-level segment reflects the recruitment of ever-increasing
numbers of neurons having best frequencies above the stimulus frequency and operating on the tail segment of their
tuning curves. Accordingly, while the AP to be masked
is made up from the contributions of a spatially limited
set of neurons (on the low-level segment of the magnitude function), effective masking can be achieved by
only moderately increasing the masker that acts on

,o'; (3.2)

60-

,,, 50-

that spatial region (the rate of increase, in fact, is

40

I0

very similar to the rate of growth of the AP magnitude

function). When, however, the AP is dominated by responses from neurons located over wide regions of the

(probe stapesvelocity in decibels -e 1 nm/sec is given as parameter). The numbers in paretheses showthe applicable Q10

basilar membrane (as on the high-level segment of

the magnitude function), correspondingly large numbers
of neurons need to be deactivated (or alesynchronized)
by the maskero Hence, the necessity for its rapid

values.

growth.

Masker
FIG. 5.

Frequency

(kHz)

AP tuning curves obtained at various probed levels

We see on the basis of the various data presented

plots and that between the top one and its. neighbor.

At

low probe levels, 10-dB increase in probe intensity

is accompanied by a much lesser increase in masker
level required to produce the criterion decrease in
N. Apparently, at sufficiently high probe levels this
relationship is reversed and, especially in the vicinity
of f0, more than 10 dB of rise in masker level is necessitatedby a 10-dB increase in the strength of the
probe. This contention is seen even better in Fig. 6,
where a plot of masker level versus probe level is given for a criterion decrease of one-half in the N. For
this plot both probe and masker are at 8 kHz. At up

thus far that simultaneous, tone-on-tone masking of

the compound action potentials of the auditory nerve
is a highly predictable and orderly process. This
masking paradigm yields patterns that are at least
qualitatively similar to both psychophysical and auditory-nerve-fiber tuning curves. These findings reinforce the notion that tone bursts are capable of synchronizing unit activity in relatively limited segments

- 3OO

to 75 dB re 1 nm/sec, the slope of the maskingfunction

is 3 dB/10 dB, while above this level the function rapidly accelerates. This behavior is obtained with maskers at other frequencies as well; notably, those maskers on the tip of the tuning curve produce parallel
masking curves, while slopes of the curves obtained
with lower-frequency maskers are somewhat shallower,

- 200

about 2 dB/10 dB. We thus see that tone-on-tone masking of the AP is a highly nonlinear process in that the
growth of the effective masker is much slower than

--

I00

<

that of the probe.

The shallow and accelerating segments of the masking functions are extremely well correlated with the
low-level and high-level segments of the AP input-

output function of the probe. To explain, when AP magnitude is plotted as a function of signal level, first a
gradual growth is seen, followed by a much steeper
segment above a critical signal input level. This two-

segmentednature of the AP magnitude (and latency)

function is very well known, and information on the

phenomenonis summarized by Dallos (1973, pp. 334347). A similar change in the rate of growth in masking as shown in Fig. 6 can also be deduced from Fig.

20

: 0 _1

I
4O

Signel
FIG. 6.

I
6O

Level

I
80

I_0
I00

(dB re I nm/sec)

Growth of masker (8 kHz) necessary to yield the

same decreasein N1 at various probe (8 kHz) levels (- ).

Masker stapes velocity is shown on left ordinate, signal
stapes velocity on abscissa. For comparison, the actual N 1
magnitude (o--o) (in/V) generated by the probe signal is
shown in the right ordinate. Both functions show two distinct

segments, with the break being at 75 dB, re 1 rim/sec.

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595

P. Dallos and M. A. Cheatham: Compound AP tuning curves

90

595

psychophysics, we were interested in examining AP

tuning curves generated by this masking process. It
is known that A P mas king does occur when the mas ke r

80

precedes the probe (Coats, 1964), and some parametric

data are also available (Spoorand Eggermont, 1971;

70

60

50

Eggermont and Spoor, 1973).

Assuming that the silent interval between successive
presentations of the masker-probe pair is long enough
to ensure virtual independence of these pairs, the
relevant parameters in a forward masking paradigm

are masker duration (T), time betweenmasker and

probe (AT), and probe duration. Inasmuchas we are

dealing with the compound AP, which is an onset response and largely independent of stimulus duration,

40

this parameter is of second-order importance and is

not studied here. Tuning curves are presented below

20-I

I
i

2
Mosker

I.

! ilil

Frequency

I0

i
2o

(khz)

FIG. 7. Comparison of AP tuning curves obtained with pulsedsimultaneous (e--e) and forward masking (o-o) paradigms.
In this case the masker duration was 75 msec in both cases;
stimulus duration: 15 msec. The probe signal appeared 50
msec after the onset of the masker in the simultaneous case,
while AT was 7.5 msec for the forward masking case. Signal
level is the same in both cases (shown by hexagonal symbol).

of the auditory nerve and that they thus generate frequency-specific APo The frequency dependence of the
tone-on-tone masking effect mimics that of isorate re-

with both T and AT as parameters, in order to assess

their influence upon the shape and sharpness of these
characteristic
plots. The first question that needs to
be answered is whether the overall shapes of the AP
tuning curves remain invariant when the experimental
paradigm changes from simultaneous to forward masking. Figure 7 demonstrates that in the forward masking case as well as in the simultaneous one, the tuning
curve is composed of a sharply tuned tip region and a
shallow, low-frequency tail. The most apparent difference

between

the two

curves

is the

masker

level

at

any given frequency that is required to produce the

criterion decrease in the N 1. Forward masking invariably requires more intense maskers. One might
conjecture, on the basis of the similarity between the
configuration of the tuning curves, that the peripheral

sponsecurves of VIIIth-nerve units. Thus, presumably,

mechanisms that result in the decrease of the N1 associ-

the masker interferes with the prob in a manner that

ated with f0 in the presence of f (that is, masking) are

parallels the basic excitatory process of single units.

In other words, as the intensity of a variable-frequency
tone needs to be altered to maintain uniform output

similar whether the two signals are present simultane-

ously or are separated in time. Of course, as Eggermont and Spoor (1973) stated, the masker must pre-

(dischargerate) from a neuronof best frequencyf0,

cede the probe.

the level of a variable-frequency

masker must be
changed in a similar manner in order to produce a

ing of the N[.

constant reduction in the response (AP) to a probe tone

of frequency f0. Of course, since the AP is a population response, there is some degree of smoothing
or averaging in its generation, and thus in the toneon-tone masking process as well.
We assume that
the generally broader tuning curves produced in AP
masking reflect the averaging of numerous sharper
response processes belonging to the population of affected neurons. It is possible that, especially at low

probe frequencies, somewhat sharper AP masking

could have been obtained with a phase-locked probe
signal. It is worth remarking that the shape of the AP
tuning curve appears to be a better reflection of the ex-

citatory area of single VIIIth-nerve neurons than twotone suppressioncontours (e.g., Sachsand Kiang, 1968).
Thus masking curves do not show the typical two-lobed
suppression patterns, which are also not seen in other
peripheral responses: CM and SP.

One does not observe backward

mask-

In Fig. 8, three families of tuning curves are shown

to demonstrate

various

subtle

effects

that

can be seen

when T and AT are varied. In each nesting family of

curves, T is constant and AT is the parameter.
The
three T values utilized are 5, 50, and 500 msec,
while AT was chosen to cover the range appropriate for
a given T. In order to see better what that range is,
plots of masker level versus AT are given in Fig. 9
for the above three T values when f =f0 = 8000 Hz.
The shorter the masker duration, the faster the masking
effects dissipate. In other words, with a T = 500 msec,
a AT of 5 or 10 msec is almost equally effective in producing masking. In contrast, with T = 5 msec, at AT
= 10 msec one requires over 20 dB more masking than
at 5 msec. The rate of cutoff is approximately the
same for each magnitude of T; thus, for the T = 1
msec condition, which we included for comparison, the
signal is no longer maskable when the AT is as short
as 5 msec.

B. Tuning curves: Forward masking

Inasmuch as various forward masking paradigms
are increasingly becoming important tools of auditory

In returning to Fig. 8 we note two interesting trends.

First, for short AT, the Q[0 (indicated in parentheses

next to each curve) is generally greater than in a si-

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596

P. Dallosand M. A. Cheatham' CompoundAP tuning curves

596

5o

5oo

I0O

.. 90

10(5.3)

80

FIG.

8.

Three

families

of for-

ward masking functions for TM

70

=5, 50, and 500 msec. Parameter is AT, shown next to the
plots. For comparison, tuning

:,

curves

30(5.2)

_ 60

Probe

50

I
4O

50-

5 (.)

'are

also

shown

for

contin-

uous masking (included in the

= 5 msec family), as well as for
pulsed simultaneous masking with
the appropriate T. Numbers in
parentheses show Q10values.
(8000 Hz) level:

50 dB

1 nm/sec.

1(.7)

com,uous f s,umEOUS
i

i I i i i

I0

Mosker

multaneous masking situation.

II

I0

Frequency

Second, as AT becomes

longer, Qz0tends to decrease. The very long (500

msec) musket condition seems to escape the above
trends; here simultaneous and forward masking situations all produce curves with approximately the same
sharpness. For relatively short masking durations,
however, the trends are very clear. Highest Qz0values are seen with very short muskets and brief silent
intervalso The Qz0decreases with the increase of both
Ts and AT. In general, for the longest AT values for
any given Ts, the Qz0 is quite comparable to its magnitude in the simultaneous masking case. Some of the
effects seen here apparently do have some phycho-

I0

(kHz)

taneous and forward masking, their resemblance to the

response areas of single fibers, and the apparent discrepancy between their shapes and single-unit two-tone
suppression areas all tend to suggest that compound

AP tuning curves rflect local excitatory processesin

the cochlea, Accordingly, one may conjecture that
AP tuning curves can provide a measure of cochlear
I00

physical analogs. Thus Houtgast (1973) notedthat psy-

./

chophysical tuning curves are sharper when obtained

with a forward masking paradigm, and Wightman (1975)

indicates that the Qz0decreases with increasing AT.
The mechanisms that produce these effects are not

clear.

The present demonstrations indicate, however,

that peripheral
III.

processes need to be involved.

SUMMARY

Action potential "tuning curves" are generated as

the plot of masker intensity versus masker frequency
required to produce a preselected decrease in the N,
elicited by a tone burst of constant frequency and level.
The resulting plots strongly resemble the shape of
single-unit tuning curves, having a sharply tuned tip
segment and an extended low-frequency tail. Tuning
curves can be generated by both simultaneous and forward masking paradigms
The similarity

of tuning curves obtained with simul-

40i

I0

20

50

Time

Deloy

(msec)

FIG. 9. Masker {f= 8 kHz) level as the function of AT with

masker duration (Ts) as the parameter. Probe is at 8 kI-Iz,
15 dB above N 1 threshold.

J. Acoust. Soc. Am., Vol. 59, No. 3, March 1976

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597

P. Dallos and M. A. Cheatham: Compound AP tuning curves

frequency selectivity.
Inasmuch as AP can easily be
measured and is the most informative response component studied in electrocochleography, it is hoped.
that AP tuning curves might become useful in this diagnostic process as indices of peripheral frequency
selectivity.

597

Kiang, N.Y.

--S.,

Watanabe, T.,

ca1 Tuning Curves of Chinchillas,"

So40(A).

ACKNOWLEDGMENT

Mitchell, C. (1973).

We thank). zdamar for his helpin this project.

Thomas, E. C.,

and

Clark, L. F. (1965) Discharge Patterns of Single Fibers in

Cat's Auditory Nerve (Res. Monograph 35, MIT Press, Cambridge, MA), pp. 29-31.
Kupperman, R. (1971). "Cochlear Masking and Adaptation,"
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