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Some results on lateral suppression obtained in a

partialmasking lateralization paradigm

Citation for published version (APA):

Bezemer, A. W. (1981). Some results on lateral suppression obtained in a partialmasking lateralization

paradigm. Journal of the Acoustical Society of America, 70(4), 996-1002. https://doi.org/10.1121/1.387034

DOI:

10.1121/1.387034

Document status and date:

Published: 01/01/1981

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Some results on lateral suppression obtained in a partial-

masking lateralization paradigm

A. W. Bezemer

Institute for Perception Research, P.O. Box 513, 5600 MB Eindhoven, The Netherlands

{Received 8 Novembe• 1979; accepted for publication 7 July 1981)

Results were reported of psychophysical forward-masking experiments using a lateralization method. A general interpretation of masking was given, considering masking to be the combined result of three different mechanisms: the overlap mechanism, the adaptation mechanism, and the suppression mechanism. The aim of this study was to demonstrate the use of the lateralization method in a masking experiment. Masking was measured in a band-widening experiment using a test tone frequency of 3 kHz, which is the center frequency of the masking noise. It was found that the effect of the suppression mechanism depends in a complex way on the difference between masker level and test tone level, as does the bandwidth at which maximum masking occurs. These level effects could be described qualitatively by means of nonlinear excitation patterns. PACS numbers: 43.66.Dc, 43.66.Mk, 43.66.Pn, 43.66.Ba [FLW]

INTRODUCTION

This paper presents some tentative results from a study of the masking process. Its primary goal is to

describe a new measuring technique which uses a

lateralization paradigm in forward masking. The new technique is applied to a band-widening experiment.

The following paragraphs describe the theoretical and

experimental framework within which the results of the

experiment have been obtained. , For the interpretation of masking data we propose the following explicit hypothesis regarding underlying mechanisms. A test tone will be masked by a masker

whenever the "central detector" is unable to detect the

presence of the test tone in the responses

in the "audi-

tory channels." The channels can be identified with

primary auditory-nerve fibers. The detector has to detect the test tone response in neutral spike trains on the basis of either rate or synchrony information

(e.g., Siebert, 1970, 1972). In case of simultaneous masking the channels are excited by both test tone and masker. Detection performance will then depend on the

ratio of the responses to test tone and masker. This ,

consideration leads to the well-known hypothesis that

masking is due to the overlap of excitation patterns

across the channels. A test tone is masked when the .

masker's excitation pattern covers the test tone pat-

tern (e.g., Zwicker, 1958). Detection of the test tone

requires the signal-to-noise ratio to exceed a certain criterion value. We term this mechanism the overlap mechanism. In absence of the masker; its role in the

signal-to-noise ratio is played by internal noise. The

internal noise determines the absolute threshold.

Theoretically, the overlap mechanism can be treated

as a linear mechanism. There are, however, two

additional

factors that influence

the excitation

pa/terns,

viz., lateral suppression and adaptation, of which at

least suppression is essentially a nonlinear one. Lateral

suppression (e.g., Sachs and Kiang, 1968; Houtgast,

1974) is assumed to work instantaneously, and to pre-

cede the adaptation

mechanism. [It is reasonable

to as-

sume that suppression occurs at the input of the hair

cell (ef., Selliek and Russell, 1979), whereas adaptation

appears to occur at the hair cell output (el., Furukawa

and Matsuura, 1978)]. The change

in sensitivity of the

auditory system during stimulation, i.e., adaptation, follows a gradual time course, as does the recovery

after termination of the masker. The relatively slow

post-stimulatory recovery of sensitivity is displayed in

forward masking (de Mar•, 1940). The amount of

adaptation

is apparently related to the excitation level.

Thus, the results of a forward-masking experiment

reflect the masker's excitation pattern. We assume that partial masking reflects the same processes as threshold masking, but to a lesser extent.

It is a psychophysical challenge to try to separate the

role of the three mechanisms experimentally. The

obvious difference between simultaneous and forward

masking is that in forward masking there is no direct

interaction between the excitation pattern of the test

tone and the masker. Thus, in forward masking, de-

tection is determined by the recovery state of the sen-

sitivity and is limited by internal noise. The overlap

mechanism is inoperative except for this internal noise

floor. Hence, the comparative study of the two gives in-

formation about the overlap mechanism. An additional

difference, and thereby a factor complicating the above analysis, is that the test tone in a simultaneous-masking

situation is subjected to the suppression mechanism, whereas this is not the case in forward masking. It

appears difficult to interpret data obtained in a simul-

taneous-masking paradigm in terms of perstimulatory adaptation and suppression because both test tone and masker are affected. The net effect of adaptation and

suppression in the signal-to-masker ratio as seen by the detector is then greatly, if not completely, reduced.

We decided to explore an additional technique where the test tone would be presented to the contralateral

ear (henceforth this tone is called the calibration tone).,

The calibration tone should scan the excitation level of

the stimulus or masker in the ipsilateral ear. This is achieved by adding a suprathreshold test tone to the masker. The ipsilateral test tone will be affected by

the masker. This effect is measurable in terms of the

lateral image produced by test tone and calibration tone. The experimental method is described in detail in Sec. I B. The l•/teralization method is applicable as

a simultaneous as well as a nonsimultaneous technique.

It yields a method to determine effects perstimulatory.

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If one assumes that the lateral position of test and cali- bration tone is determined by their relative loudnesses, ß it is obvious that the results of this experiment are re- lated to partial-masking data.

As an example of the applicability of the lateralization paradigm, we present some data from a band-widening experiment. In this paper we do not explore the time parameter, but we investigate the effect of both masker

level and test tone level in a forward-masking situation. In 1970 Greenwood and Goldberg found, as a result of

their physiological experiments, that the firing rate

measured in neutrons of the cochlear nucleus first in-

creases with increasing bandwidth of a noise stimulus

and then decreases when the noise bandwidth increases

beyond a certain value. The noise had a constant spec-

tral density. Ruggero (1973) obtained a corresponding result from primary auditory-nerve fibers.

Comparable psychophysical results have been obtained with nonsimultaneous masking methods. Houtgast (1974) performed the band-widening experiment with the pulsa-

tion-threshold method and obtained the same nonmono-

tonic result that was obtained physiologically. Schreiner

(1977) corroborated Houtgast's results. He observed

that the bandwidth at which maximum masking occurs

corresponds

to the critical band. Houtgast (1974) as-

sumed that the decrease of masking beyond a certain

bandwidth is caused by lateral suppresssion. Leshowitz

and Lindstrom

(•977), Terry and Moore (1977), and

Weber (1978) carried out band-widening experiments with a conventional forward-masking paradigm. Their

experiments produced results that are qualitatively

similar to results obtained with the pulsation-threshold

method. Following Houtgast, s interpretation, the dif-

ference found between maximum masking and masking

measured at the widest noiseband used can be con-

sidered a measure of the lateral suppression effect.

We expect a similar trend in lateralization data. The

amount of nonmonotonicity may be different, however.

In this experiment we explore level parameters. Ob- viously, a follow-up study investigating temporal

parameters is called for.

I. METHOD

A. Experimental setup and procedure

In all our experiments the masker consisted of band- pass-filtered noise with a variable bandwidth centered around 3 kHz. This noiseband was obtained by multiply- ing lowpass-filtered noise having a variable cutoff fre- quency by a pure tone having a frequency of 3 kHz. The

multiplication was carried out with a 12-bit digital

modulator (Willeros et al., 1977). After filtering, the

slope steepness of the noiseband was about 5 dB/100

Hz. Test tone and calibration tone were pure tones at the center frequency of the noiseband, i.e., 3 kHz.

The duration of the masker was 290 ms, including

rise and fall times of 20 ms (see Fig. 1). Test tone

and calibration tone had a duration of 20 ms with rise

and fall times of 10 ms. All signals had Gaussian- shaped onsets and offsets. Test tone and calibration

o

. M

290 time(ms)

time (ms)

FIG. 1. Time course of the stimulus with the lateralization

method. R: stimulus to the right ear, L- stimulus to the left ear, M. masker, T. test tone, C: calibration tone.

tone were presented simultaneously 25 ms after the starting point of the masker's offset ramp.

The bandwidth of the noise was the independent variable and could vary from 60 to 2000 Hz. The

spectral level of the noise masker and the level of the

test tone were independent parameters and could be

equal to, respectively, 50, 30, 10dB/Hz and 75, 65,

55, 35 dB (all dB's re 20 btPa).

The subject was seated in a sound-insulated booth. The stimulus was presented through Pioneer SE-700 headphones. The subject was asked to set, by means of adjustment, the level of the calibration tone that was

required to perceive the hearing sensation in the median

plane of his head. Between two stimulus presentations

there was always a fixed silent interval of 1190 ms. All

measuring points were presented three times in a

pseudorandom

sequence. Three subjects, having

normal audiograms and considerable experience in

psychophysical experiments, participated in the ex-

periments.

B. Description of the lateralization method

The lateralization method was used as long as 25 years

ago for measuring auditory fatigue or adaptation in non-

simultaneous masking conditions (Hood, 1950; Harris

and Rawnsley, 1953). More recently, Houtgast

(1977),

Jesteadt and Jayel (1978), and Kearney (1979) have

reported on the application of a corresponding lateraliza-

tion paradigm for measuring suppression in a simul-

taneous-masking experiment.

The lateralization method is based on the fusion

phenomenon that occurs when a pure tone is presented diotically. To one ear of a subject, a masker and a test tone are presented; and simultaneously with the test tone, a calibration tone is presented to the contralateral

ear. The calibration tone is equal to the test tone except

for its amplitude. The test tone is masked partially by the masker (in this study, in a forward-masking para- digm). In general the test tone remains well perceptible. Thus, the lateralization method is a suprathreshold method. The subject's task is to adjust the level of the

calibration tone is such a way that he perceives the

hearing sensation which belongs to the combination of test tone and calilJration tone, at an arbitrary point of the 'median plane of his head. We assume that in this condition test tone and calibration tone have equal loud- ness. Then the adjusted physical level difference be-

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tween test tone and calibration tone is a measure of the

amount of (partial) masking caused by the masker.

Besides a level difference between test tone and cali-

bration tone, other factors can also influence the position of the hearing sensation. Examples are a phase differ- ence between the tones, or a difference in time between

the presentation of the tones to the ears. When such a difference occurs in our experiments, this may in- fluence the results. These two possible artifacts are

discussed below.

In spite of sufficient control of the physical signals used, an internal phase difference might occur between

test tone and calibration tone due to interaction between

test tone and masker. Therefore, it is important to

know what the influence of such a difference between the

tones is. The effect of a phase difference on the position of the hearing sensation depends on frequency. This dependence has been tested experimentally. In accord-

ance with results of other investigators (e.g., Licklider

and Webster, 1949; Zwislocki and Feldman, 1956), we

found that such a phase difference is significant only at frequencies below 1500 Hz. We performed our experi- ments in a frequency region above 1500 Hz, so the pos- sible occurrence of a phase difference cannot have a significant influence on our results.

When a time difference is present between the onsets of both tones, the hearing sensation will be found at the

side of the ear first stimulated. This effect is related

closely to the precedence effect (Wallach et al., 1949).

From an explorative experiment, it followed that with

our signals a time difference smaller than 2 ms has

very little influence on the position of the hearing sen- sation. With time differences greater than 2 ms the two contralateral tones no longer fuse perfectly, and

it is clear to the listener that a time difference is

present. From this it follows that interaural time dif- ferences, which are easily controlled to .values smaller than 1 ms, will be negligible.

Since the amount of forward masking of the test tone depends on masker level, the decreases in calibration tone level required to center the hearing sensation at different masker levels could conceivably be perceived as different time delays. This is a problem for all experiments measuring time-dependent phenomena, but it should have only a minor effect in the present experiment. It can be minimized by choosing a test tone that is as short as possible, without being too broad

spectrally (e.g., a Gaussian-shaped 5-10-ms signal).

To compare the variability of the lateralization method as described above with the variability of the conventional threshold method, an exploratory forward- masking experiment was performed with both methods.

A noiseband with a bandwidth from 2 to 4 kHz and a

spectral level equal to 30 dB/Hz was used as a masker.

The test tone frequency was at the center frequency of

the noise mask•r, i.e., 3 kHz. With the lateralization

method, ten different calibration tone levels were

pseudorandomly presented ten times each (with a fixed test tone level of 75 dB). The subject was asked to

indicate if he perceived the fused hearing sensation at

the left- or at the right-hand side of the median plane of his head. With the threshold method, ten different test tone levels were pseudorandomly presented ten times each. The subject was now asked to indicate if he could perceive the test tone. From these results a psycho-

metric function was determined for both methods. For

the given conditions both functions coincide almost com- pletely, and a standard deviation of 1.5 dB was found graphically for both methods. It might be expected that with the lateralization method, the level of the test tone has some influence on the slope of the psychometric function. However, we suppose that this will be the case

for very low levels (near threshold) only. For all other levels, the decision "left or right" seems to be rather

independent of level.

C. Some perceptual impressions

The position of the hearing sensation, produced by a pure tone which is presented diotically through head- phones, is somewhere at the median plane of the listen-

er's head. We found that the precise position was de-

pendent on the frequency of the tone. When a low fre- quency is applied, the hearing sensation is perceived

at the back of the head (near the nape of the neck). With increasing frequency the hearing sensation shifts up- wards over the back of the crane to the front. The up-

ward shift has also been found with experiments on

sound localization. With these experiments, stimuli are presented through loudspeakers. In 1930 Pratt ob-

served that "high tones are phenomenologically

higher

in space than low tones." His observations were con- firmed by Roffler and Butler (1968).

Further, it was noticed that at a relatively high fre-

quency (3 kHz), the hearing sensation seems to exist

in a sharp focused point. At lower frequencies the di- mensions of this point seem to grow, so that a less pre- cise spot is perceived.

After some training, subjects preferred experiments

in which the lateralization method was applied to simi- lar experiments that used the classical threshold paradigm. Threshold methods require in general more

concentration and time. II. RESULTS

In a first experiment the influence of masker level

on the masking effect was studied, starting with a fixed

test tone level (65 dB). Results are shown in Fig. 2.

Because there is no indication that the standard deviation

varies systematically with the bandwidth of the noise,

we computed an estimation of the mean standard devia-

tion for each curve. Twice this standard deviation is indicated at the left-hand side of each curve.

The curves for the lowest noise level (10 dB/Hz) show

that masking increases with increasing bandwidth. Max- imum masking occurs at the largest bandwidth (2000 Hz) and amounts to 9-14 dB, depending on the subject. As

the noise level increases to 30 dB/Hz, more masking

is found for all bandwidths. In addition, we observe that

the curves tend to flatten ou• and for subject AB there

is even a decrease. The curves measured at the highest noise level (50 dB/Hz) follow a somewhat different

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25 2O 15 10 o 20 = 15 o .c_ 2O N O (dB/Hz) L T (dB $PL) x 50 65 O 30 65 [] 10 65 HZ : •. ! : ... lO s

%0

... i'oo

bafidwidth (Hz)

FIG. 2. Masking curves measured as a function of the band-

width of the masking noise for three subjects. Three noise

levels No, fixed test tone level

course. The rising character with increasing band- width is only initially present in the results of subject AB, at bandwidths larger than 400 Hz the curve bends

downwards. Results of H Z show an almost continuous

falling character, and JV's results seem to be fairly

independent of bandwidth. Results obtained at this

noise level (except for JV) are in agreement with ex-

pectations based on lateral suppression: masking effec- tiveness of the central part of a noiseband is reduced by the outer parts, if the bandwidth is large enough. In the figures, this results in a decrease of masking at larger

bandwidths.

At the lowest noise level, suppression has no observ-

able influence. To determine whether there is no sup-

pression at all at such a low level (10 dB/Hz) or whether

its influence is not measurable with a relatively loud

test tone (65 dB), we repeated the foregoing experiment,

but now with a fixed difference between masker level and test tone level. This difference was chosen in such

a way that at every masker level the test tone remained clearly perceptible. The results of this experiment are shown in Fig. 3. We observe at once that with the sub- jects AB and HZ, in all three conditions, masking first increases with increasing bandwidth and then de- creases. At all three levels, lateral suppression has

an observable effect. Further we note that least

masking is found at the highest noise level. JV's data differ markedly from the data of the other. To a cer- tain extent this was already the case in the previous experiment, and we will observe similar discrepancies in the next experiments.

The set of data collected up to now can easily be ex-

999

J. Acoust.

Soc.

Am., Vol. 70, No. 4, October

1981

3O 25 20 15 10 0 m 30

• 25

e• 20 15 _.e 10 ß ½- 0 25 2o 15 10 No (dB/Hz) L T (dB SPL) x 50 75 ¸ 30 55 [] 10 35 '. : : :::111 I I I HZ I i i ::: ::: bandwidth (Hz)

FIG. 3. As Fig. 2, but three conditions with a fixed difference

between spectral noise levelN 0 and test tone level Lr.

tended by performing two series of measurements, one at a masker level of 30 dB/Hz in combination with a test tone level of 75 dB, and another at a masker level of 50 dB/Hz in combination with a test tone level of 55

dB. Results obtained from these measurements have

been combined with results from Figs. 2 and 3, yielding

two sets of curves measured at fixed noise levels (30

and 50 dB/Hz) with test tone level as parameter. These

two sets of curves are shown in Figs. 4 and 5. For all

three subjects, it holds that at a noise level of 30 dB/Hz

(Fig. 4), maximum masking is measured at the lowest

test tone level (55 dB). However, at larger bandwidths

the curves approach each other; this is mainly due to the fact that masking measured at the lowest test tone level

decreases relatively strongly (except for JV). It is of

particular interest to note that the three curves in each panel do not parallel each other. This can also be ob- served in Fig. 5, which shows results obtained at a noise level of 50 dB/Hz. The curves obtained at this level also tend to coincide at larger bandwidths. Results of JV are somewhat different again; the differing order of the curves is especially noteworthy.

III. DISCUSSION

The lateralization paradigm produces data with a

reproducibility similar to that of a threshold paradigm. The stimulus provides an additional parameter, viz.,

the level of the test tone. This is inherent to partial-

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3O 25 2O 5 25

• 2o

= 15 • lO ß 0 ._ 25 20 15 10 5 N O (dB/Hz) L T (dB SPL) 30 75 30 65 30 55 AB : i '. '.T, III•' I I I I {I!11 I I I HZ JV

...

...

i'oo ' ' '

bandwidth (Hz)

FIG. 4. As Fig. 2, but with a fixed spectral noise level N O

and three test tone levels L T .

masking paradigms. After discussion of the data of this particular experiment, we will return to the potential possibilities of the lateralilation method.

Our results shown in Fig. 2 indicate that the effect of the suppression mechanism, as apparent from the non- monotonic behavior, increases with increasing masker

level. This agrees with results of Houtgast (1974), Terry and Moore (1977), and Weber (1978). Results

from experiment• on two-tone suppression point to a

corresponcling level effect (Duifhuis, 1980). Further-

more, our results tend to show that if a maximum oc-

curs in the masking curves, it shifts towards greater

bandwidths with decreasing noise level. This would

become more prominent if we speculate that at the

lowest noise level (10 dB/Hi), a maximum Will be

present at some bandwidth greater than 2000 Hi. (Un-

fortunately, for practical reasons it was impossible to carry out measurements at bandwidths larger than

2000 Hi.) Although less clear, Houtgast (1974), using

the pulsation-threshold method, also observed such a

shift of the maximum. In the results of forward-mask-

ing experiments by Weber (1978), no maximum shift

is observable. In both studies, masker levels ranging from about 20-50 dB/I-Iz were used.

Comparing Figs. 2 and 3, we note that the difference between masker level and test tone level is of impor- tance for the observation of lateral suppression. At a

masker level of 10 dB/Hz no suppression effect is ob-

served in Fig. 2, but in Fig. 3 it is clearly present. However, because suppression is assumed to be an

instantaneous

effect (cf., Introduction), it is not pos-

25 2O 15 lO .e 20 ;• 15 ß 10 25 2O 10 No(dB/Hz) L T (dB SPL) x 50 ¸ 50 [] 50 75 65 ; I I : ; iiii I i I bandwidth (Hz)

FIG. 5. As Fig. 4, but with a different spectral noise level

N O ß

sible that the presentation of the subsequent test tone influences the preceding suppression. Consequently, we are led to the conclusion that the sensitivity for

measuring suppression depends on test tone level (and

test tone versus masker level difference).

The influence of test tone level on the observation of

the suppression effect is demonstrated more clearly by the results shown in Figs. 4 and 5. It is not a very sur- prising effect if one recalls that in general partial

masking

decreases

as the signal-to-noise ratio •n-

creases (Zwicker, 1958; Scharf, 1964). Thus it would

be a consistent result if the measured interaural level

difference would be found to decrease with increasing test tone level. This does appear to be the case for small bandwidths only, however. The behavior at the largest bandwidths shows not much effect of level. This nonuniform behavior may be interpreted on the basis of excitation patterns which are level dependent.

It can be concluded from our previous results (Beze- mer, 1978) that the suppression mechanism has its

maximum effect at the central frequency region of the masking noise. From this it follows that a test tone with an excitation pattern that occurs just within this frequency region will show the effect of the suppression mechanism most clearly. In general, with increasing signal level, the excitation pattern of a signal becomes

progressively broader (e.g., Zwicker and Feldtkeller,

1967). In our case, with increasing test tone level, the

excitation pattern of the test tone will extend over a

much wider frequency range than just the central part of the masking noise. Therefore, the total average

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sensitivity to suppression (within the preceding masker)

will then become smaller. This trend describes the

results shown in the upper anti middle panels of Figs. 4 and 5: the largest difference between maximum mask- . ing and masking measured at the widest noise band

(2000 Hz) is found at the lowest test tone level (55 dB).

In line with this result, the maximum in the masking curve shifts towards a higher bandwidth when the level

of the test tone increases.

However, there is a complication in the above inter- pretation because there are two opposing effects. With increasing bandwidth, and to a certain extent also with

increasing level, there is not only an increase in sup-

pression but also a broadening of the excitation pattern

of the noise band. The broadening can cause more

masking; this depends on the difference between test

tone level and masker level. It is not easy to predict whether the suppression or broadening effect will dom- inate. A quantitative description of these effects would be required. At this point such a description is not -

available, so for the moment our reasoning is supported

by the results showed in this study only. From this it follows that implications about suppression based on threshold measurements cannot directly be extended to

above-threshold levels.

Returning to the results shown in Fig. 3 (constant level difference between test tone and masking noise), it

may be remarked that least masking is found at the highest noise level. This suggests that compared to the excitation pattern of the masking noise, the test tone pattern becomes relatively broader when both the level

of the test tone and the masker are increased. As the

high-frequency slope of the excitation pattern is strong-

ly level-dependent (Zwicker and Feldtkeller, 1967), this

seems quite plausible, taking into account that the test tone level is higher than the masker level. At this point it is necessary to assume that the high-frequency slope of the noise is determined mainly by its spectral level

(and not by its overall level). Also, with these results,

only a small suppression effect is observable at the highest test tone level. This agrees with the results shown in Figs. 4 and 5.

The curves measured for subject JV differ in many aspects from the curves measured with the other sub- jects. Very often his results seem to be nearly inde- pendent of bandwidth. Compared with other studies on band widening, this is rather exceptional. The fact that

no or only little influence of suppression (decreasing masking with increasing bandwidth) is observable might

be less exceptional. Great differences between subjects

also have been found in psychophysical studies on two-

tone suppression (Shannon, 1976; Duifhuis, 1980). How-

ever, it is still possible that influence of the suppression mechanism is present in JV's results. The influence might be derived from the relative positions of the mea- sured curves, eog., Fig. 2 lower panel. The interin-

dividual differences found with suppression experiments

pose an interesting point for future research.

IV. CONCLUSIONS

The binaural lateralization method has proved itself a convenient and useful tool in partial-masking experi-

ments.

From measured psychometric functions it can be concluded that the sensitivity of the lateralization method is about equal to the sensitivity of the conven-

tional threshold method.

The occurrence of lateral suppression in a band- widening experiment can be measured by making use

of the lateralization method. The influence of masker

level on the suppresssion effect has been studied by means of this method at a fixed test tone frequency of 3 kHz.

At a fixed test tone level, the greatest suppression effect is measured at the highest masker level. At a fixed masker level, the greatest suppression effect is

measured at the lowest test tone level. With different combinations of masker level and test tone level with a

fixed difference between these levels, the size of the suppression effect is not constant. Implications about suppression based on threshold measurements cannot directly be extended to above-threshold levels.

The bandwidth of the noise masker at which maxi-

mum masking occurs depends on the difference between masker level and test tone level; as this dif- ference increases, the bandwidth at which maximum masking occurs decreases.

The results are interpreted qualitativity in terms of nonlinear excitation patterns. This interpretation is based on three mechanisms: the overlap mechanism, the suppression mechanism, and the adaptation mechan- ism. However, the interpretation within the given framework is not completed yet.

The effect of lateral suppression differs considerably between different subjects.

ACKNOWLEDGMENTS

The author would like to thank.H. Duifhuis for his

stimulating help in preparing this paper. He also grate- fully acknowledges J. v. d. Vorst and H. W. Zelle for

participating in the experiments. This research was

supported by the Netherlands Organization for the Advancement of Pure Research (Z.W.O.).

Bezemer, A. W. (1978). "Lateralisation applied to masking experiments," IPO Ann. Prog. Rep. 13, 13-19.

de Mar•, G. (1940). "Fresh observations as to the so-called

masking effect of the ear and its possible diagnostic signifi- cance," Acta Oto-Laryngol. 28, 314-316.

Duifhuis, H. (1980). "Level effects in psychophysical two- tone suppression," J. Acoust. Soco Am. 67, 914-927. Furukawa, T., and Matsuura, S. (1978). "Adaptive rundown

of excitatory post-synaptic potentials at synapses between hair cells and eight nerve fibres in the goldfish," J. Physiol.

276, 193-209.

Greenwood, D. D., and Goldberg, J. M. (1970). "Response

of neurons in the cochlear nuclei to variations in noise band-

width and to tone-noise co. mbinations," J. Acoust. Soc. Am.

47, 1022-1040.

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Harris, J. D., and Rawnsley, A. I. (1953). "'The locus of

short duration auditory fatigue or 'adaptation'," J. Exp.

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