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PubMed1996

Sensitivity to interaural intensity differences of neurons in primary auditory cortex of the cat. I. types of sensitivity and effects of variations in sound pressure level.

1. Interaural intensity differences (IIDs) provide the major cue to the azimuthal location of high-frequency narrowband sounds. In recent studies of the azimuthal sensitivity of high-frequency neurons in the primary auditory cortex (field AI) of the cat, a number of different types of azimuthal sensitivity have been described and the azimuthal sensitivity of many neurons was found to vary as a function of changes in stimulus intensity. The extent to which the shape and the intensity dependence of the azimuthal sensitivity of AI neurons reflects features of their IID sensitivity was investigated by obtaining data on IID sensitivity from a large sample of neurons with a characteristic frequency (CF) > 5.5 kHz in AI of anesthetized cats. IID sensitivity functions were classified in a manner that facilitated comparison with previously obtained data on azimuthal sensitivity, and the effects of changes in the base intensity at which IIDs were introduced were examined. 2. IID sensitivity functions for CF tonal stimuli were obtained at one or more intensities for a total of 294 neurons, in most cases by a method of generating IIDs that kept the average binaural intensity (ABI) of the stimuli at the two ears constant. In the standard ABI range at which a function was obtained for each unit, five types of IID sensitivity were distinguished. Contra-max neurons (50% of the sample) had maximum response (a peak or a plateau) at IIDs corresponding to contralateral azimuths, whereas ipsi-max neurons (17%) had the mirror-image form of sensitivity. Near-zero-max neurons (18%) had a clearly defined maximum response (peak) in the range of +/- 10 dB IID, whereas a small group of tough neurons (2%) had a restricted range of minimal responsiveness with near-maximal responses at IIDs on either side. A final 18% of AI neurons were classified as insensitive to IIDs. The proportions of neurons exhibiting the various types of sensitivity corresponded closely to the proportions found to exhibit corresponding types of azimuthal sensitivity in a previous study. 3. There was a strong correlation between a neuron's binaural interaction characteristics and the form of its IID sensitivity function. Thus, neurons excited by monaural stimulation of only one ear but with either inhibitory, facilitatory, or mixed facilitatory-inhibitory effects of stimulation of the other ear had predominantly contra-max IID sensitivity (if contralateral monaural stimulation was excitatory) or ipsi-max sensitivity (if ipsilateral monaural stimulation was excitatory). Neurons driven weakly or not at all by monaural stimulation but facilitated binaurally almost all exhibited near-zero-max IID sensitivity. The exception to this tight association between binaural input and IID sensitivity was provided by neurons excited by monaural stimulation of either ear (EE neurons). Although EE neurons have frequently been considered to be insensitive to IIDs, our data were in agreement with two recent reports indicating that they can exhibit various forms of IID sensitivity: only 23 of 75 EE neurons were classified as insensitive and the remainder exhibited diverse types of sensitivity. 4. IID sensitivity was examined at two or more intensities (3-5 in most cases) for 84 neurons. The form of the IID sensitivity function (defined in terms of both shape and position along the IID axis) was invariant with changes in ABI for only a small proportion of IID-sensitive neurons (approximately 15% if a strict criterion of invariance was employed), and for many of these neurons the spike counts associated with a given IID varied with ABI, particularly at near-threshold levels. When the patterns of variation in the form of IID sensitivity produced by changes in ABI were classified in a manner equivalent to that used previously to classify the effects of intensity on azimuthal sensitivity, there was a close correspondence between the effects of intensity on corresponding types of azimuthal and IID sensitivity

Animals↗
PubMed1997

Two-tone distortion on the basilar membrane of the chinchilla cochlea.

Basilar membrane responses to pairs of tones were measured, with the use of a laser velocimeter, in the basal turn of the cochlea in anesthetized chinchillas. Frequency spectra of basilar membrane responses to primary tones with frequencies (f1, f2) close to the characteristic frequency (CF) contain prominent odd-order two-tone distortion products (DPs) at frequencies both higher and lower than CF (such as 2f1-f2, 3f1-2f3, 2f2-f1 and 3f2-2f1). For equal-level primaries with frequencies such that 2f1-f2 equals CF, the magnitude of the 2f1-f2 DP grows with primary level at linear or faster rates at low stimulus levels, but it saturates or decreases slightly at higher levels. For a fixed level of one of the primary tones, the magnitude of the 2f1-f2 DP is a nonmonotonic function of the level of the other primary tone. For low intensities of the variable tone, the 2f1-f2 DP grows at a rate of approximately 2 dB/dB with f1 level and 1 dB/dB with f2 level. DP magnitudes decrease rapidly with increasing primary frequency ratio (f2/f1) at low stimulus levels. For more intense stimuli, DP magnitudes remain constant or decrease slowly over a wide range of frequency ratios until a critical value is reached, at which DP magnitudes fall with slopes as steep as -300 dB/octave. As stimulus level grows, DP phases increasingly lag for large f2/f1 ratios, but exhibit leads for small f2/f1 ratios. Cochlear exposure to an intense tone that produces large sensitivity losses for the primary frequencies (but only small losses for tones with frequency equal to 2f1-f2) causes a substantial decrease in magnitude of the 2f1-f2 DP. This result demonstrates that the 2f1-f2 DP originates at the basilar membrane region with CFs corresponding to the primary frequencies and propagates to the location with CF equal to the DP frequency. 2f1-f2 DPs on the basilar membrane resemble those measured in human psychophysics in most respects. However, the magnitude of basilar membrane DPs does not show the nonmonotonic dependence on f2/f1 ratio evident in DP otoacoustic emissions.

Animals↗
PubMed1997

Processing of interaural intensity differences in the LSO: role of interaural threshold differences.

Cells in the lateral superior olive (LSO) are known to be sensitive to interaural intensity differences (IIDs) in that they are excited by IIDs that favor the ipsilateral ear and inhibited by IIDs that favor the contralateral ear. For each LSO neuron there is a particular IID that causes a complete inhibition of discharges, and the IID of complete inhibition varies from neuron to neuron. This variability in IID sensitivity among LSO neurons is a key factor that allows for the coding of a variety of IIDs among the population of cells. A fundamental question concerning the coding of IIDs is: how does each cell in the LSO derive its particular IID sensitivity? Although there have been a large number of neurophysiological studies on the LSO, this question has received little attention. Indeed, the only reports that have directly addressed this question are those of Reed and Blum, who modeled the binaural properties of LSO neurons and proposed that the IID at which discharges are completely suppressed should correspond to the difference in threshold between the excitatory, ipsilateral and inhibitory, contralateral inputs that innervate each LSO cell. The main purpose of this study was to test the threshold difference hypothesis proposed by Reed and Blum by recording responses to monaural stimulation and to IIDs from single cells in the LSO of the mustache bat. Our results show that although the IID sensitivities of some LSO cells correspond to the difference in threshold between the excitatory and inhibitory ears, in the majority of cells the difference in thresholds did not correspond to the cell's IID sensitivity. The results lead us to propose two models to account for IID sensitivities. One model is similar to that proposed by Reed and Blum and emphasizes differences in the thresholds of the excitatory and inhibitory inputs. This model accounts for the minority of cells in which the IID of complete inhibition corresponded to the difference in threshold of the inputs from the two ears. The other model, which accounts for the cells in which the IID of complete inhibition did not correspond to the difference in the thresholds of the inputs from the two ears (the majority of cells), places emphasis on differences in latencies of the excitatory and inhibitory inputs. The models incorporate features that are concordant with the known properties of the neurons that project to the LSO and together can account for the diversity of IID sensitivities among the population of LSO neurons.

Animals↗
PubMed1997

Responses of neurons in the inferior colliculus to binaural masking level difference stimuli measured by rate-versus-level functions.

The psychophysical detection threshold of a low-frequency tone masked by broadband noise is reduced by < or = 15 dB by inversion of the tone in one ear (called the binaural masking level difference: BMLD). The contribution of 120 low-frequency neurons (best frequencies 168-2,090 Hz) in the inferior colliculus (ICC) of the guinea pig to binaural unmasking of 500-Hz tones masked by broadband noise was examined. We measured rate-level functions of the responses to identical signals (So) and noise (No) at the two ears (NoSo) and to identical noise but with the signal inverted at one ear (NoS pi): the noise was 7-15 dB suprathreshold. The masked threshold was estimated by the standard separation, "D". The neural BMLD was estimated as the difference between the masked thresholds for NoSo and NoS pi. The presence of So and S pi tones was indicated by discharge rate increases in 55.3% of neurons. In 36.4% of neurons, the presence of So tones was indicated by an increase in discharge rate and S pi tones by a decrease. In 6.8% of neurons, both So and S pi tones caused a decrease in discharge rate. In only 1.5% of neurons was So indicated by a decrease and S pi by an increase in discharge rate. Responses to the binaural configurations were consistent with the neuron's interaural delay sensitivities; 34.4% of neurons showing increases in discharge rate to both So and S pi tones gave positive BMLDs > or = 3 dB (S pi tones were detected at lower levels than So), whereas 37.3% gave negative BMLDs > or = 3 dB. For neurons in which So signals caused an increase in the discharge rate and S pi a decrease, 72.7% gave positive BMLDs > or = 3 dB and only 4.5% gave negative BMLDs > or = 3 dB. The results suggest that the responses of single ICC neurons are consistent with the psychophysical BMLDs for NoSo versus NoS pi at 500 Hz, and with current binaural interaction models based on coincidence detection. The neurons likely to contribute to the psychophysical BMLD are those with BFs near 500 Hz, but detection of So and S pi tones may depend on different populations of neurons.

Animals↗
PubMed2000

Temporal discharge patterns evoked by rapid sequences of wide- and narrowband clicks in the primary auditory cortex of cat.

The present study investigated neural responses to rapid, repetitive stimuli in the primary auditory cortex (A1) of cats. We focused on two important issues regarding cortical coding of sequences of stimuli: temporal discharge patterns of A1 neurons as a function of inter-stimulus interval and cortical mechanisms for representing successive stimulus events separated by very short intervals. These issues were studied using wide- and narrowband click trains with inter-click intervals (ICIs) ranging from 3 to 100 ms as a class of representative sequential stimuli. The main findings of this study are 1) A1 units displayed, in response to click train stimuli, three distinct temporal discharge patterns that we classify as regions I, II, and III. At long ICIs nearly all A1 units exhibited typical stimulus-synchronized response patterns (region I) consistent with previously reported observations. At intermediate ICIs, no clear temporal structures were visible in the responses of most A1 units (region II). At short ICIs, temporal discharge patterns are characterized by the presence of either intrinsic oscillations (at approximately 10 Hz) or a change in discharge rate that was a monotonically decreasing function of ICI (region III). In some A1 units, temporal discharge patterns corresponding to region III were absent. 2) The boundary between regions I and II (synchronization boundary) had a median value of 39.8 ms ICI ([25%, 75%] = [20.4, 58. 8] ms ICI; n = 131). The median boundary between regions II and III was estimated at 6.3 ms ([25%, 75%] = [5.2, 9.7] ms ICI; n = 47) for units showing rate changes (rate-change boundary). 3) The boundary values between different regions appeared to be relatively independent of stimulus intensity (at modest sound levels) or the bandwidth of the clicks used. 4) There is a weak correlation between a unit's synchronization boundary and its response latency. Units with shorter latencies appeared to also have smaller boundary values. And 5) based on these findings, we proposed a two-stage model for A1 neurons to represent a wide range of ICIs. In this model, A1 uses a temporal code for explicitly representing long ICIs and a rate code for implicitly representing short ICIs.

Acoustic Stimulation↗
PubMedDate not supplied

A 'neural' response with 3-ms latency evoked by loud sound in profoundly deaf patients.

A large negative deflection with a latency of 3 ms was observed in the auditory brainstem response (ABR) waveforms of some patients with peripheral profound deafness. This deflection was termed the N3 potential. In this paper, we review patients with the N3 potential and discuss the characteristics of abnormal ABR waveforms. The origin of the N3 potential was also discussed, especially with respect to vestibular evoked potentials. In most of the patients, audiograms showed no response to the maximum output of an audiometer in the high-frequency range and a residual response in the low-frequency range. The N3 potentials were noted at intensities of 80 dB nHL or greater. As the stimulus intensity increased, the amplitude of the potential increased and the latency decreased. A high repetition rate (83.3/s) of the click stimulus influenced the latency and amplitude of the N3 potential. The potential was replicated on retest within less than a month, and had a consistent latency and amplitude over the scalp. The results indicate that the N3 potential is not an electrical artifact but a physiological neural response evoked by a loud sound. The N3 potential is most likely not an auditory evoked response from cochlear or a response from a semicircular canal, because it has a 3-ms latency, a sharp waveform, and is unassociated with vertigo. The results suggest that the N3 potential may be a saccular acoustic response.

Acoustic Stimulation↗
PubMedDate not supplied

Preattentive memory-based comparison of sound intensity.

Changes in the intensity of repeated, ignored sounds elicit the mismatch negativity (MMN) brain response which reflects preattentive detection of the change. It is generally assumed that the MMN in response to intensity changes reflects a memory-based comparison mechanism rather than being due to differential states of refractoriness of intensity-specific cortical neurons. In the present study, an experimental protocol consisting of 4 oddball blocks and 1 control block was used in order to separate memory-comparison-related effects from refractoriness-related ones. This design allowed an assessment of intensity MMN using physically identical stimuli with equal probability of occurrence in separate blocks, while avoiding contamination by refractoriness. Results were consistent with an MMN in response to intensity change that reflects genuine memory-based comparison.

Acoustic Stimulation↗
PubMed2006

Evaluation of the mixing point in tinnitus sound therapy by a psychoacoustic matching protocol with a digital tinnitus evaluation system.

Directive counseling and sound therapy have been reported to effectively alleviate tinnitus suffering. Without objective evaluation, researchers doubt the real effect of sound therapy. This study was designed to evaluate the frequency and intensity of the theoretical 'mixing point' (MP) in sound therapy and to investigate its relationship with the minimal masking level (MML) of tinnitus. The tinnitus tones of 133 patients were successfully matched in frequency (dominant in 4-10 kHz) and loudness (83.09 +/- 12.04 dB) using a psychoacoustic matching protocol and a newly designed tinnitus evaluation system. The matching rate was 83%. The relationship between the MP and MML is described by a linear regression equation: MP = 0.90 x MML + 1.98 (r2 = 0.82, p < 0.0001). The results of matching and correlation analysis confirmed the consistency of the MP in sound therapy. The psychoacoustic characteristics of tinnitus can be objectively evaluated with our matching protocol and evaluation system.

Acoustic Stimulation↗
PubMed2006

Measuring the refractoriness of the electrically stimulated auditory nerve.

Intracochlear recordings in cochlear implant recipients provide access to the electrically evoked compound action potential (ECAP). ECAP thresholds are potential predictors of speech processor map's threshold and comfortable loudness levels. The auditory nerve's refractory properties can influence these levels due to high-rate stimulation with interpulse intervals within the relative refractory period. Recovery functions were investigated at 84 stimulation sites in 14 Nucleus CI24 recipients using neural response telemetry and a modified forward masking technique. This technique introduces a reference masker-probe interval (MPI). In our study, an appropriate value between 300 and 375 micro s was determined for this reference MPI, and the use of a reference MPI of 300 micro s is suggested for recovery and amplitude growth functions. A median absolute refractory period of about 390 micro s and a median time constant of about 425 micro s were obtained by fitting an exponential model to the data. Hence, the auditory nerve is usually in relative refractory state when standard neural response telemetry forward masking is selected because of its default MPI of 500 micro s. This can bias the measurement of ECAP thresholds. Additionally, the shape of standard forward masking recovery functions was explained by the influence of latency shift of the neural response.

Action Potentials↗
PubMedDate not supplied

Noise-induced cochlear hypoxia is intensity dependent, correlates with hearing loss and precedes reduction of cochlear blood flow.

Anesthetized and artificially ventilated guinea pigs were exposed to broad-band noise of 95, 101, 106 or 115 dB SPL for 30 min and studied for 180 min after cessation of noise. The partial pressure of oxygen (pO2) in the perilymph, the cochlear blood flow (CoBF) and auditory-evoked potentials were continuously recorded. Arterial blood pressure, electrocardiogram, inspiratory and expiratory gas levels, arterial blood gas levels and acid-base status were kept stable to exclude influences of these parameters on cochlear parameters. Exposure to 95 dB SPL did not affect perilymphatic pO2 or CoBF. Cochlear microphonics (CMs) were reduced, but compound action potentials of the auditory nerve (CAPs) and auditory brainstem potentials (ABRs) increased after exposure to this low-level noise. Perilymphatic pO2 decreased during exposure to 101 dB SPL and then further decreased during the subsequent 60 min after cessation of the noise. CoBF did not change significantly during and 30 min after noise but then paralleled the decline of perilymphatic pO2. However, both parameters showed a clear indication of recovery in the second and third hours after noise. At 101 dB SPL, CMs were again reduced immediately, CAPs were unaltered and ABRs again increased. Exposure to 106 and to 115 dB SPL resulted in a decrease in both perilymphatic pO2 and CoBF; this decrease began during the exposure but became progressively worse after the noise. Hearing loss was observed immediately with exposure and showed no signs of further deterioration after cessation. The observed time courses of changes are important. They reveal that hearing loss and cochlear hypoxia precede reduction in CoBF due to noise exposure. The potential mechanisms underlying these effects are discussed.

Animals↗
PubMedDate not supplied

Neurophysiological correlate of the auditory after-image ('Zwicker tone').

The 'Zwicker tone' (ZT) is an auditory after-image that can be evoked most effectively when a band-suppressed noise (relative width of gap 1/3 octave) presented for a certain period of time has been switched off. The sensation of this purely monaural phenomenon is that of a pure tone with a frequency corresponding to the center frequency of the gap and an equivalent level of 10-15 dB above auditory threshold. The sensation decays gradually; it may last as long as 10 s depending on how long the evoking noise was presented. The search for a physiological correlate has been futile so far, probably because the search was confined to more peripheral levels of the auditory system (inferior colliculus). A neuromagnetic study was performed in normal-hearing subjects in order to look for a neurophysiological correlate of the ZT in the auditory cortex. With a stimulation paradigm especially designed for this study, we have been able to isolate poststimulus activity which appears to be related to the ZT and which originates in the supratemporal auditory cortex. It is a sustained neuromagnetic activity that shows a clear-cut dipolar field distribution, and it appears that this activity has certain similarities with the tone-evoked auditory sustained response. The hypothesis is put forward that during the sensation of the ZT a process takes place in the auditory cortex which is similar to that underlying the sustained response, and which gives rise to the sensation of the ZT. In contrast to the sustained response, however, which is due to neural activity evoked by an external acoustic stimulus, the sustained activity associated with the ZT is due to a temporary absolute or relative reduction of neural activity originating from those regions in which the ZT exciting stimulus caused an adaptation. These differences in neural activity cannot be distinguished by the auditory system from a corresponding external acoustic signal. Preliminary studies in patients suffering from tonal tinnitus yielded results which exhibit a certain similarity with those obtained in the ZT experiment.

Acoustic Stimulation↗
PubMedDate not supplied

2f1-f2 distortion product otoacoustic emissions in White Leghorn chickens (Gallus domesticus): effects of frequency ratio and relative level.

The effects of primary tone frequency ratio (f2/f1 ratio) and relative level (L2/L1) on the amplitude of the cubic difference tone (CDT: 2f1-f2) distortion product otoacoustic emissions (DPOAEs) were investigated in adult White Leghorn chickens (Gallus domesticus). In experiment 1, 9 f2/f1 ratios ranging from 1.05 to 1.8 were investigated. Measurements were obtained from both ears of 4 chickens at 7 f1 frequencies ranging from 0.8 to 4.0 kHz. The primary tones were equal in level, and varied from 20 to 80 dB SPL. The mean CDT amplitude increased with increasing primary tone level once the measurement noise floor was exceeded. The input/ output functions assumed one of two shapes: one in which there was a systematic increase in DPOAE amplitude with increasing primary tone level, and the other in which there was a plateau in the input/output function near 65-70 dB SPL. At the highest primary tone level (80 dB SPL), there was a decrease in the CDT amplitude with increasing f2/f1 ratio. At high primary tone levels, the f2/f1 ratio which produced the largest CDT was 1.05 or 1.1, while at lower primary tone levels the largest CDT occurred at f2/f1 ratios of 1.2-1.3. In experiment 2, L2 was held constant at 70 dB SPL, and L1 varied from 50 to 80 dB SPL. For f1 frequencies of 0.8 and 3.2 kHz, there was an increase in the CDT amplitude with increasing L1, followed by an asymptote at higher levels. In contrast, for 1.6 and 2.0 kHz f1 frequencies, the amplitude increased, plateaued and then increased again at higher levels. Informal measurements suggest that spontaneous otoacoustic emissions (SOAEs) are rarely seen in chickens. However, a reliable SOAE was observed in 1 chicken, which could be suppressed by external sounds and anoxia.

Acoustic Stimulation↗
PubMedDate not supplied

Effect of trial-by-trial variation in center frequency on detection of interaural temporal and intensitive differences in bands of noise.

Sensitivity to interaural temporal disparities (ITDs) and interaural intensitive disparities (IIDs) was measured as a function of duration in conditions where the center frequency (CF) of the stimuli was constant and, separately, in conditions where the CF of the stimuli was varied on a trial-by-trial basis. Stimuli were bands of noise centered at either 300, 1200, 2400, or 4800 Hz with the bandwidth of the noise being 40% of their CF. The largest effects of shortening duration and varying center CF of the stimuli were degradations of sensitivity to ITDs for stimuli centered at 2400 or 4800 Hz. Overall, the data confirm and extend previous findings which indicate that ITDs and IIDs are processed separately within the central nervous system.

Acoustic Stimulation↗
PubMedDate not supplied

A new method of measurement of most comfortable loudness.

A new computer-assisted method for the measurement of most comfortable loudness (MCL) is presented, where not only MCL but also its reliability can be graphically displayed and measured. Two unique features concerning the reliability of MCL measurement are obtained in this method: sharpness and stability. Sharpness is defined by the rate of repeat selection of the same level as MCL, and stability is used to express the consistency of a subject's response during the experiment. A method of analyzing sharpness and stability was explained by showing two contrasting cases. Based on the graph configurations, the subjects were divided into two major categories, type A and type B, defined by sharpness of judgment. Type A subjects' judgments are sharper than those of type B. Decreasing the frequency increased the occurrence of type A subjects. Retest results revealed that this method has a high degree of reproducibility of MCL judgment, which supports the reliability of this method.

Adolescent↗
PubMedDate not supplied

Detection of the acoustic reflex below 80 dB HL.

A new method for detecting the acoustic reflex that utilizes standard otoacoustic emissions recording techniques is introduced and discussed. Two successive identical tone bursts of 100 ms duration and 10 ms interstimulus interval are presented in the occluded ear canal at a repetition rate of one per second. If the acoustic reflex is elicited, the contraction of the stapedius muscle is delayed with respect to the onset of the first stimulus. Hence, the acoustic compliance in the ear canal decreases primarily during the second stimulus. The difference of the microphone signals produced by the two stimuli is computed and averaged across a certain number of repetitions of the sequence. The presentation level is increased until this difference is larger than -40 dB (with respect to the stimulus level) and if its signal-to-noise ratio exceeds 20 dB. For normal-hearing subjects, the acoustic reflex threshold measured with this method is on average 8 dB lower than in a standard clinical setup. In 5 out of the 10 tested hearing-impaired subjects, the new method could detect an acoustic reflex at one or more frequencies where no reflex was detected in the clinical setup.

Adult↗
PubMed1980

Audiological considerations in Ménière's disease.

The audiological findings in Ménière's disease are reviewed particularly in respect of loudness recruitment and speech discrimination loss. Although recruitment is pathognomonic of all cochlear lesions, certain distinguishing features are apparent in Ménière's disease which differ from those in other non-metabolic disorders of the cochlea. With mild degrees of hearing loss, recruitment appears to aid speech discrimination supporting the suggestion that it represents the ears adjustment to failing hearing. With more profound degrees of hearing loss, there is a linear inverse relationship between maximum discrimination score and hearing level. Actual hearing aid gains preferred by patients correlate well with those predicted from a consideration of recruitment curves.

Hearing↗