Forward masking of the compound action potential: thresholds for the detection of the N1 peak

A two-interval forced-choice method was developed that provides a rapid and objective computerized measurement of the threshold for detection of the N1 peak of the compound action potential (CAP) recorded in response to a probe tone. The CAP was recorded at the round window of anesthetized chinchillas and several gerbils. An adaptive threshold-tracking procedure was verified by comparing measured thresholds to those obtained from neurometric functions, which plot the proportion of correct detections of the probe as a function of probe intensity. The adaptive procedure was applied in a forward masking paradigm to study the growth of masking of the CAP as a function of masker intensity. Results indicate that growth of masking of the CAP more closely corresponds to that observed psychophysically, than does forward masking observed in the response of a single neuron. Implications for neural encoding mechanisms are discussed.     

Recovery from adaptation in human compound action potential following forward masking

Recovery from adaptation was studied as changes in compound action potential (CAP) in response to clicks after broadband noise masking in volunteers with normal hearing. CAP was transtympanically elicited from the promontory using a needle electrode. Preceding masking noise and subsequent click stimuli were delivered separately from 2 loud speakers 80 cm from the tested ear. We evaluated the effect of masking noise duration and intensity on CAP recovery from adaptation as a function of delta t, defined as the interval between masking noise offset and click stimulus onset. Results were as follows: At delta t shorter than 200 ms or less, recovery from latency prolongation and amplitude decrease slowed with increasing masking noise duration and intensity. At delta t longer than 200 ms, no significant difference was seen in CAP recovery based on changes in masking noise duration or intensity. Recovery from adaptation, which depended on click intensity, took about 700 ms at a 40 dBnHL click and about 300 ms at a 60 dBnHL click. The influence of the middle ear muscle reflex on CAP recovery patterns is neglected when considering the intensity both masking noise and click stimulus. In CAP recovery, accumulated effects of the relative auditory-nerve refractory period may adversely affect recovery at very short delta t, while recovery from short-term adaptation, assumed involved in cochlear-hair cell synapse function, may be an important factor at delta t longer than 30-40 ms. Based on animal experiments showing different recovery times dependent on different auditory nerve spontaneous rates(SR), we concluded that at delta t of 200 ms or less, CAP recovery reflects high SR neuron activity, while at delta t longer than 200 ms, it mainly reflects that of low SR neurons.     

Absence of tonic activity of the crossed olivocochlear bundle in determining compound action potential thresholds, amplitudes and masking phenomena in anaesthetised guinea pigs with normal hearing sensitivities

In Nembutal- or Urethane-anaesthetised guinea pigs N1 audiograms and N1 input-output functions were measured as were compound action potential (CAP) tuning curves under forward masking and simultaneous masking conditions. Then the crossed olivocochlear bundle was lesioned at the floor of the fourth ventricle and the cochlear responses were re-measured. There were never any changes in the N1 audiograms, input-output functions, or the CAP tuning curves. Thus, the crossed efferent pathways do not appear to play any tonic role in determining cochlear threshold sensitivities, selectivities or masking phenomena in anaesthetised guinea pigs with normal hearing sensitivities.     

Suprathreshold comparisons of derived and enhanced compound action potentials.

The Derived cochlear nerve compound action potential (CAP) and the Enhanced CAP are both measures which demonstrate the ability of a forward masker to increase the amplitude of the CAP produced by a probe stimulus. Enhancement occurs whenever the amplitude of a masked CAP is larger than that of a nonmasked CAP, whereas the derived CAP is produced by the subtraction of the entire masked CAP waveform from that of the nonmasked waveform. Therefore, a derived CAP is created whenever the masker produces a difference of amplitude, latency, and/or waveform shape. The present experiments compare these two measures by observing the effects of 13 kHz maskers varying from levels of -10 to +70 db SPL on CAPs produced by 50 or 60 dB SPL, 13 kHz probe stimuli. Enhancement is characterized by a nonmonotonic increase of CAP amplitude (and sometimes a decrease of latency) as a function of increasing levels of the forward masker, whereas this pattern seldom occurs with the derived CAP. Enhancement is typically seen with forward making, but seldom seen with simultaneous masking, whereas the derived CAP is very similar under these two types of masking.     

Enhancement of the cochlear nerve compound action potential: sharply defined frequency-intensity domains bordering the tuning curve.

Forward masking can either decrease or increase the response to a subsequent stimulus. In the gerbil, frequency-intensity domains of the maskers that decrease the amplitude of the compound action potential (CAP) can be plotted as the sharply defined CAP tuning curve (TC). Regions were also found over which masking increases (enhances) the amplitude of the CAP. Center-frequency (CF) enhancement domains were found in approximately 2/3 of the animals tested, in response to maskers having frequencies very near that of the probe stimulus, at levels ranging from below the CAP detection threshold to just below the tip threshold of the TC. Approximately 2/3 of the animals showing CF enhancement also displayed low-frequency (LF) enhancement, in response to a domain which borders the low-frequency tail of the TC.     

Efferent tracts and cochlear frequency selectivity

The cochlear innervation of guinea pigs was sectioned medially in a rostrocaudal direction at the level of the floor of the fourth ventricle, to study the effects of efferent pathways on cochlear microphonic (CM) suppression, the compound action potential (CAP) masking phenomenon, the input-output CAP function, and cochlear frequency selectivity estimated with tuning curves of single auditory nerve fibers. Sectioning reduced CM suppression without having any effect on absolute CM amplitude; it also reduced CAP masking. The input-output CAP function was not changed at intensities below 75 dB, and the single-unit tuning curves recorded before and after nerve sectioning were unaffected. Thus, the crossed efferent tracts (i.e., mainly the medial system) seems to be involved in the masking function itself, rather than one of the mechanisms responsible for high frequency cochlear selectivity.     

Frequency-specific enhancement of the cochlear compound action potential: influence of the forward masker.

Under restricted frequency and intensity conditions, forward masking can result in the amplitude of the CAP being increased above its unmasked value (Henry, 1991). The present study provides a quantitative analysis of this enhancement effect. In response to forward maskers having the same frequency as the probe stimulus, central frequency (CF) enhancement varies as a function of the level of the forward masker: the lowest masker level at which it can reliably be detected is often well below the visual detection threshold of the CAP generated by the unmasked probe stimulus; the highest masker level at which it can reliably be detected corresponds to approximately 10 dB above the probe stimulus CAP threshold. A second low frequency (LF) enhancement region also exists, encompassing a narrow range of more intense maskers. CF enhancement can double the amplitude of the CAP, whereas LF enhancement is less pronounced. The magnitude of CF enhancement varies as a function of the duration of the forward masker, with longer durations generally increasing the magnitude of the effect. This duration effect, however, interacts with the level of the stimulus. Decreasing the interval between the end of the forward masker and the beginning of the probe increases the magnitude of CF enhancement.     

Latency and Amplitude Tuning Curves of the N1 and N2 Components of the Cochlear Nerve Compound Action Potential

Compound action potential tuning curves (CAP TCs) generated by masking the N1 component of the CAP provide a means of assessing the ability of the cochlea to selectively tune to certain stimuli. This paper examines some of the factors which can influence this TC when a moderately intense (i.e. 40–80 dB SPL) probe stimulus is used. At these levels, each of the four corners of the trapezoidal stimulus envelope is capable of generating a CAP. Also, short stimulus rise times can merge the CAPs produced by the first two corners, but this does not appear to have a major effect on the CAP TC. It was shown that the N2 component of the CAP for the first corner of the stimulus is equally capable of producing a well-tuned TC. Another study has shown that, in addition to amplitude decrements, one can use latency increases as a criterion for CAP TCs. We have demonstrated that latency TCs are more finely tuned than amplitude TCs at high levels, especially when the stimulus rise time is short.     

Latency and amplitude tuning curves of the N1 and N2 components of the cochlear nerve compound action potential

Compound action potential tuning curves (CAP TCs) generated by masking the N1 component of the CAP provide a means of assessing the ability of the cochlea to selectively tune to certain stimuli. This paper examines some of the factors which can influence this TC when a moderately intense (i.e. 40-80 dB SPL) probe stimulus is used. At these levels, each of the four corners of the trapezoidal stimulus envelope is capable of generating a CAP. Also, short stimulus rise times can merge the CAPs produced by the first two corners, but this does not appear to have a major effect on the CAP TC. It was shown that the N2 component of the CAP for the first corner of the stimulus is equally capable of producing a well-tuned TC. Another study has shown that, in addition to amplitude decrements, one can use latency increases as a criterion for CAP TCs. We have demonstrated that latency TCs are more finely tuned than amplitude TCs at high levels, especially when the stimulus rise time is short.     

Acoustic hemifields in the spatial release from masking of speech by noise

The Hearing-in-Noise Test (HINT) is able to measure the benefit to speech intelligibility in noise conferred when the noise masker is displaced 90 degrees in eccentricity from a speech source located at zero degrees azimuth. Both psychoacoustic and neurophysiological data suggest that the perceptual benefit of the 90-degree azimuth separation would be greatest if the speech and noise were presented in different acoustic hemifields, and would be smallest if the two sources were in the same acoustic hemifield. The present study tested this hypothesis directly in ten normal-hearing adult listeners. Using the HINT stimuli, we confirmed the hypothesis. Release from masking scores averaged 8.61 dB for "between-hemifield" conditions, 6.05 dB for HINT conditions, and 1.27 dB for "within-hemifield" conditions, even though all stimulus configurations retained a 90-degree angular separation of speech and noise. These data indicate that absolute separation of speech and noise alone is insufficient to guarantee a significant release from masking, and they suggest that what matters is the location of the stimulus elements relative to the left and right spatial perceptual channels.     

Role of the efferent medial olivocochlear system in contralateral masking and binaural interactions: an electrophysiological study in guinea pigs.

Contralateral broadband noise (BBN) elevates ipsilateral auditory thresholds (central masking) and reduces the amplitude of ipsilateral brainstem auditory evoked potentials (BAEPs). Binaural interactions are complex psychophysical phenomena, but binaural interaction components are easily extracted from BAEPs to monaural versus binaural click stimulation. However, contralateral, or binaural, acoustical stimulation is known to activate simultaneously the crossed and uncrossed medial olivocochlear (MOC) efferent systems and decrease activity in both cochleas. Particularly, contralateral BBN stimulation suppresses in part ipsilateral peripheral activity. What is the role of such contralaterally induced peripheral suppression in the overall changes in central BAEPs observed during contralateral masking or binaural stimulation? Compound action potentials (CAPs) of the auditory nerve and BAEPs were recorded simultaneously in awake guinea pigs from electrodes chronically implanted on the round window of the cochlea and the surface of the brain. Peripheral and central measures of contralateral masking and binaural interactions were obtained from responses to monaural or binaural clicks, with or without contralateral BBN, recorded before, during, and after the reversible blockade of the MOC function following a single intramuscular injection of gentamicin. Contralateral BBN effectively reduced the amplitudes of CAP and of all BAEP peaks. CAP to ipsilateral click did not, however, change significantly from monaural to binaural click stimulation; still, normal binaural interaction components developed in the BAEPs. When the medial efferent function was blocked by gentamicin, the normal contralateral BBN suppression of CAP and of the earliest BAEP peak was lost; however, the later BAEP peaks were suppressed by contralateral BBN as before gentamicin, and the central binaural interaction components were unchanged. In these experimental conditions, the MOC efferent system seems to play little role in centrally recorded contralateral masking and binaural interactions.     

Functional properties of the crossed part of the medial olivo-cochlear bundle

The efferent innervation of guinea pig cochleas was sectioned medially, at the level of the floor of the fourth ventricle, to study the effects of the crossed part of the medial efferent pathway on the compound action potential (CAP) masking phenomenon. Sectioning reduced CAP masking for a masker level varying with the frequency of the masker and the time elapsed between the masker onset and the probe onset. Functional properties of the crossed part of the medial efferent tracts: latency, thresholds and frequency selectivity, could be deduced from these data.     

Auditory Nerve Frequency Tuning Measured with Forward-Masked Compound Action Potentials

Frequency selectivity is a fundamental cochlear property. Recent studies using otoacoustic emissions and psychophysical forward masking suggest that frequency selectivity is sharper in human than in common laboratory species. This has been disputed based on reports using compound action potentials (CAPs), which reflect activity in the auditory nerve and can be measured in humans. Comparative data of CAPs, obtained with a variety of simultaneous masking protocols, have been interpreted to indicate similarity of frequency tuning across mammals and even birds. Unfortunately, there are several issues with the available CAP measurements which hamper a straightforward comparison across species. We investigate sharpness of CAP tuning in cat and chinchilla using a forward masking notched-noise paradigm—which is less confounded by cochlear nonlinearities than simultaneous masking paradigms and similar to what was used in the psychophysical study reporting sharper tuning in humans. Our parametric study, using different probe frequencies and notch widths, shows relationships consistent with those of auditory nerve fibers (ANFs). The sharpness of tuning, quantified by Q ₁₀ factors, is negatively correlated with probe level and increases with probe frequency, but the Q ₁₀ values are generally lower than the average trend for ANFs. Like the single fiber data, tuning for CAPs is sharper in cat than in chinchilla, but the two species are similar in the dependence of tuning on probe frequency and in the relationship between tuning in ANFs and CAP. Growth-of-maskability functions show slopes <1 indicating that with increasing probe level the probe is more susceptible to cochlear compression than the masker. The results support the use of forward-masked CAPs as an alternative measure to estimate ANF tuning and to compare frequency tuning across species.     

The effects of moderate cooling on gross cochlear potentials in the gerbil: basal and apical differences.

Changes in the threshold of the compound action potential (CAP) response in the gerbil to low- and high-frequency tonebursts were monitored during uniform cooling of the cochlea by 7-8 degrees C below normal body temperature. Recordings of the endocochlear potential (EP), cochlear microphonic (CM), and summating potentials (SP) were also obtained from the base and apex of the cochlea under the same conditions. Cooling-related changes in the CAP, as well as the CM and SP response obtained near the best frequency of the recording location, were greater in the base than in the apex. In contrast, reductions in the EP appeared uniform throughout the cochlea. Thus the greater vulnerability of CAP thresholds in the base does not result from a greater vulnerability of the stria vascularis in this region. Our results suggest that the enhanced susceptibility to cooling of the CAP in the cochlear base reflects changes in hair cell mechanisms.     

Offset tuning curves generated by simultaneous masking are more finely tuned than those generated by forward masking

The rapid ending of a tone produces an evoked potential which has different properties than that which is produced by the sudden onset of a tone. At the level of the round window, the offset N1 N2 follows the ending of the cochlear microphonic (CM) by approximately the same amount of time as does the onset N1 N2 to the onset of the CM. Both onset and offset responses are abolished with cochlear lesion. Continuous masking was used to generate tuning curves (TCs) from the NI-PI component of the evoked potential recorded from the round window of the gerbil. Those evoked potentials generated in response to the tone onset were complementary in appearance to those generated in response to the tone offset. TCs generated by continuous masking of the NI-PII component of the auditory brainstem response (ABR) of the gerbil show the same pattern. When it is generated by simultaneous masking, the midfrequency offset TC in the gerbil and mouse is W-shaped. It has two well tuned tips which occur at frequencies below and above that of the probe stimulus used to generate the TC. It also has an even better tuned peak occurring at or slightly above the probe stimulus frequency, which becomes sharper as the masker sound pressure level (SPL) is increased from 50 to over 80 dB. Because the midfrequency onset response is approximately 40 dB lower than the midfrequency offset response, probe stimuli for onset TCs are generally set at lower SPLs. When the onset probe stimulus is set to the same level as that of the offset probe, the Q10 dB of the offset TC may be up to 10 times the value of the Q10 dB of the onset TC. The offset TC generated in the CBA/J mouse by forward masking is quite different from that produced by simultaneous masking. Both forward and simultaneous conditions utilized a 40 ms duration tone to mask the PI-NI component of offset and onset ABRs of the mouse which were evoked by a 10 ms duration, 32 kHz tone, presented at an interstimulus interval of 160 ms. Forward masking (when compared with simultaneous masking) resulted in a more sharply tuned onset TC. But the offset TC was much less sharply tuned in the forward masking condition. This suggests that the offset response may reflect functions which are involved with fine tuning at moderate to high intensities in the presence of simultaneous sounds of similar spectral characteristics.(ABSTRACT TRUNCATED AT 400 WORDS)     

Continuum of impulsiveness caused by auditory masking

OBJECTIVE: Impulsivity is a hallmark of attention deficit/hyperactivity disorder (ADHD). Various auditory masking procedures can quantify the impulsivity caused by distracting background sounds. This study compares the impulsiveness and distraction caused by informational masking (unpredictable tones) with previously published data on central masking (contralateral noise) in children with and without ADHD. METHODS: Twenty-six normal and 14 children diagnosed as having ADHD (combined type), all between the ages of 7 and 13, indicated whether they heard a 512-ms, 500-Hz pure tone in a single-interval task under conditions of informational masking and in quiet. The masker consisted of 10 randomly selected frequencies between 1,000 and 2,500 Hz presented simultaneously at an overall level of 60 dB SPL. A maximum-likelihood method estimated thresholds and false alarm rates. RESULTS: There were no differences due to ADHD in thresholds or false alarm rates either with informational masking or in quiet. With informational masking, normal children had high false alarm rates, similar to those from children with ADHD under central masking. With informational masking, all children tended to say a stimulus was present when it was not. CONCLUSIONS: All children behave impulsively under some conditions. Pediatric patients with attention disorders can thus be reassured that impulsiveness with unpredictable background sounds is normal, to some extent. Response biases of children with ADHD may only diverge from normal in situations where distracting external stimuli have an intermediate level of predictability. A previous study showed that with central masking, children with ADHD are more impulsive than normal. There appears to be a limit to the uncertainty in auditory masking that can be tolerated by children. Children with ADHD become impulsive at lower levels of uncertainty than normal. Increasing the predictability of distracting background sounds may thus improve the performance of children with ADHD. Informational masking may, for normal listeners, mimic something of what it is like to have an attention deficit. ADHD can be profitably studied with auditory tasks.     

The effect of perceived spatial separation on informational masking of Chinese speech

The effect of perceived spatial separation, induced by the precedence effect, on release from noise or speech masking was investigated. Listeners were asked to orally repeat Chinese nonsense sentences, which were spoken by a female talker and presented by both the left (-45 degrees) and right (+45 degrees) loudspeakers, when maskers, which were either speech-spectrum noise sounds or Chinese nonsense sentences spoken by two other female talkers, were presented by the same two loudspeakers. Delays between identical sounds presented over the two loudspeakers were used to control the perceived locations of the target (right only) and masker (right, center, or left). The results show that perceived 45 degrees and 90 degrees separations of target speech from masking speech led to equivalently marked improvement in speech recognition, even though the degree of improvement was smaller than that reported in [J. Acoust. Soc. Am. 106 (1999) 3578 (using English nonsense speech)]. When the masker was noise, however, perceived separation only marginally improved speech recognition. These results indicate that release from informational masking, due to perceived target/masker spatial separation induced by the precedence effect, also occurs for tonal Chinese speech. Compared to the 45 degrees perceived within-hemifield separation, the 90 degrees perceived cross-hemifield separation does not produce further unmasking.     

Study of tinnitus masking by self-recording audiometer]

The tinnitus masking test, in which the minimum masking levels of tinnitus by various pure tones and band noises are measured and used to produce tinnitus masking curves, is one of the methods for evaluating the character of tinnitus. At present, the tinnitus masking test is usually performed using a pure tone audiometer. In this study, tinnitus masking curves were produced using a self-recording audiometer (Bekesy audiometer) in 22 cases of tinnitus, and the basic nature of the tinnitus masking curves from the self-recording audiometer was investigated and compared with those from a pure tone audiometer. The results showed no changes in the masking level, and the amplitudes of the tinnitus masking curves from the self-recording audiometer were observed to be at the tinnitus pitches. The amplitude of the tinnitus masking curve showed a negative correlation with tinnitus loudness as measured by the loudness balance test for tinnitus, that is, the amplitude tended to decrease as the tinnitus became louder. This suggests that the loudness of the masking tone influences the tinnitus masking phenomenon.     

Compound action potentials and single unit responses to a 1 kHz haversine in the cat

The single-cycle 1 kHz haversine (one cycle of a 1 kHz sine wave beginning at -90 degrees) is a low-frequency impulsive stimulus which has been little use, but which has significant potential applications both as a clinical and a research tool. The auditory nerve compound action potential (CAP) and single unit discharge patterns evoked by a single-cycle 1 kHz haversine stimulus were studied in anesthetized cats. The haversine CAP waveform consisted of two or three short latency peaks with peak to peak intervals of about 1.0 ms. Latencies of the CAP peaks decreased with increased stimulus intensity and were also strongly dependent on stimulus polarity. Typically, CAP peak latencies changed by about 0.5 ms with stimulus polarity reversal. Single unit responses were classified by the peak latency pattern of their haversine post-stimulus time histograms (PSTHs). Low CF units had low thresholds and PSTHs resembling their click responses. High CF units had high thresholds and PSTHs comprised of one or two short latency peaks whose latencies were polarity-sensitive. Some units in an intermediate CF range (approximately 1.5-3.0 kHz) had PSTHs which were a transitional form between the high and low CF types of response. The unit discharge patterns strongly suggested a low frequency origin for the haversine CAP at all intensities.     

Endocochlear potentials and compound action potential recovery: functions in the C57BL/6J mouse

The C57BL/6J mouse suffers from cochlear degeneration beginning at an early age and has been used as a model of age-related hearing loss (presbyacusis). Here, the endocochlear potential (EP) and compound action potential (CAP) responses were determined in one-, four-, 12- and 24-month-old C57BL/6J mice. CAP measures included thresholds to tone pips, input/output (I/O) functions, and recovery functions to conditioning tones. EP values among the four age groups did not differ significantly (P>0.05) in either the basal or apical turns. CAP thresholds were increased significantly by 10 to 30 dB in the four-month group compared to the one-month controls at 11.3, 16, 20, and 22.6 kHz. CAP I/O functions were shallower in the four-month group compared to controls at all frequencies. In the 12- and 24-month-old mice, CAP responses were absent, despite normal EP values in these animals. Recovery functions after conditioning tones were obtained at 8, 16, 20 and 22.6 kHz; the functions had fast and slow components at all frequencies tested in both the one- and four-month-old groups. The corresponding recovery curves were identical for both age groups, even with significant threshold shifts in the older group. The two component recovery curves provide the first physiological evidence that different spontaneous rate (SR) classes of auditory neurons exist in the C57BL/6J mouse. Moreover, the unchanged recovery functions in the older group suggest that there was no loss of activity of the low-SR fiber population with age under conditions where the EP remains stable, in contrast to the gerbil model of presbyacusis where there is a loss of low-SR fiber activity and EP does decline with age.