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PubMed1993

High-frequency neurons in the inferior colliculus that are sensitive to interaural delays of amplitude-modulated tones: evidence for dual binaural influences.

1. Localization of sounds has traditionally been considered to be performed by a duplex mechanism utilizing interaural temporal differences (ITDs) at low frequencies and interaural intensity differences at higher frequencies. More recently, it has been found that listeners can detect ITDs at high frequencies if the amplitude of the sound varies and an ITD is present in the envelope. Here we report the responses of neurons in the inferior colliculi of unanesthetized rabbits to ITDs of the envelopes of sinusoidally amplitude-modulated (SAM) tones. 2. Neurons were studied extracellularly with glass-coated Pt-Ir or Pt-W microelectrodes. Their sensitivity to ITDs in the envelopes of high-frequency sounds (> or = 2 kHz) was assessed using SAM tones that were presented binaurally. The tones at the two ears had the same carrier frequency but modulation frequencies that differed by 1 Hz. This caused a cyclic variation in the ITD produced by the envelope. In this "binaural SAM" stimulus, the carriers caused no ITD because they were in phase. In addition to the binaural SAM stimulus, pure tones were used to investigate responses to ipsilateral and contralateral stimulation and the nature of the interaction during binaural stimulation. 3. Neurons tended to display one of two kinds of sensitivity to ITDs. Some neurons discharged maximally at the same ITD at all modulation frequencies > 250 Hz (peak-type neurons), whereas others were maximally suppressed at the same ITD (trough-type neurons). 4. At these higher modulation frequencies (> 250 Hz), the characteristic delays that neurons exhibited tended to lie within the range that a rabbit might normally encounter (+/- 300 microseconds). The peak-type neurons favored ipsilateral delays, which correspond to sounds in the contralateral sound field. The trough-type neurons showed no such preference. 5. The preference of peak-type neurons for a particular delay was sharper than that of trough-type neurons and was comparable to that observed in neurons of the inferior colliculus that are sensitive to delays of low-frequency pure tones. 6. At lower modulation frequencies (< 150 Hz) characteristic delays often lay beyond +/- 300 microseconds. 7. Increasing the ipsilateral intensity tended to shift the preferred delay ipsilaterally at lower (< 250 Hz), but not at higher, modulation frequencies. 8. When tested with pure tones, a substantial number of peak-type neurons were found to be excited by contralateral stimulation but inhibited by ipsilateral stimulation.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1993

Sensitivity of auditory nerve fibers to spectral notches.

1. Listeners use direction-dependent spectral cues introduced by the torso, head, and pinnae to localize the source of a sound in space. Among the prominent direction-dependent spectral features in the free field-to-eardrum transfer function are narrow regions of low acoustic energy referred to as spectral notches. In this paper, we studied the sensitivity of single auditory nerve fibers in the barbiturate-anesthetized cat to broadband noise that had been filtered by a function whose shape approximated natural notches in the free field-to-eardrum transfer function. 2. Two experimental paradigms were employed. The first was the repeated presentation of a burst of broadband noise filtered by the simulated-notch function. Center frequency of the notch was held constant at or around the fiber characteristic frequency (CF). We refer to this as a "stationary" notch stimulus. The second paradigm was the repeated presentation of a broadband noise that was constructed from noise segments, each filtered by the simulated notch, whose CF was incremented and then decremented in a systematic way. We refer to this as a "moving" notch stimulus. Results from these two paradigms were compared with respect to notch detection. 3. Data were obtained from 161 single auditory nerve fibers having CFs ranging from 0.4 to 40 kHz. Most fibers studied had CFs > 5 kHz, and they detected the presence of the spectral notch in an intensity- and frequency-dependent manner. Each fiber responded vigorously to the presence of broadband noise. When the CF of the notch encroached on the response area of the fiber, there was a demonstrable reduction in discharge rate. The greatest reduction in discharge rate occurred when the notch was centered at the fiber's CF and when the level of the notch signal was some 15-55 dB above the fiber's noise threshold. There was close association in the frequency-intensity plane between the position of the most effective notch and the fiber's threshold tuning curve. 4. For high-spontaneous rate fibers, a moving-notch stimulus, but not a stationary one, reduced the discharge below the spontaneous rate at and in the immediate vicinity of the most effective notch frequency. This increases sensitivity to a spectral notch and suggests a mechanism by which localization ability is enhanced when there is relative motion between a sound source and the head.(ABSTRACT TRUNCATED AT 400 WORDS)

Acoustic Stimulation↗
PubMed1993

Virtual-space receptive fields of single auditory nerve fibers.

1. Sounds reaching the tympanic membranes are first modified by the acoustic properties of the torso, head, and external ear. For certain frequencies in the incident sound there results a complex, direction-dependent spatial distribution of sound pressure at the eardrum such that, within a sound field, localized areas of pressure maxima are flanked by areas of pressure minima. Listeners may use these spatial maxima and minima in localizing the source of a sound in space. The results presented describe how information about this spatial pressure pattern is transmitted from the cochlea to the central auditory system via single fibers of the auditory nerve. 2. Discharges of single fibers of the auditory nerve were studied in Nembutal-anesthetized cats [characteristic frequencies (CFs) ranged from 0.4 to 40 kHz]. Click stimuli were derived from sound-pressure waveforms that were generated by a loudspeaker placed at 1,800 locations around the cat's head and recorded at the tympanic membrane with miniature microphones. Recorded signals were converted to acoustic stimuli and delivered to the ear via a calibrated and sealed earphone. The full complement of signals is referred to as "virtual acoustic space," and the spatial distribution of discharges to this array of signals is referred to as a "virtual-space receptive field" (VSRF). 3. Fibers detect both pressure maxima and pressure minima in virtual acoustic space. Thus VSRFs take on complex shapes. 4. VSRFs of fibers of the same or similar CF having low spontaneous rates had the same overall pattern as those from high-spontaneous rate (HSR) fibers. For HSR fibers, the VSRF is obscured by the high background spike activity. 5. Comparison of the VSRF and isolevel contour maps of the stimulus derived at various frequencies revealed that auditory nerve fibers most accurately extract spectral information contained in the stimulus at a frequency close to or slightly higher than CF.

Animals↗
PubMed1994

Encoding of amplitude modulation in the cochlear nucleus of the cat.

1. Amplitude modulation (AM) is a pervasive property of acoustic communication systems. In the present study we investigate neural temporal mechanisms in the auditory nerve and cochlear nuclei of the pentobarbital sodium-anesthesized cat associated with the neural coding of 100% AM tones, both in quiet and in the presence of wideband, quasi-flat-spectrum noise. The AM carrier frequency was set to the neuron's characteristic frequency (CF) and the sound pressure level (SPL) of acoustic stimuli was varied over a wide dynamic range of intensities (< or = 40 dB). The temporal AM-encoding capability of auditory neurons was measured by computing the synchronization coefficient (SC) of the neural response to the signal's modulation and carrier frequency. The temporal modulation transfer function (tMTF) of a neuron was then computed by measuring the SC of the response to signals of variable fmod (50-2550 Hz). 2. Neurons in the cochlear nuclei synchronize on average more highly to the modulation frequency than fibers of comparable CF, threshold, and spontaneous rate in the auditory nerve. The disparity in performance is greatest at high SPLs and low signal-to-noise ratios. However, there is a significant degree of diversity in AM-encoding capability among neurons in both the cochlear nuclei and auditory nerve. Among auditory nerve fibers (ANFs), low- and medium-spontaneous-rate (SR) units (SR < 18 spike/s) phase-lock with greater precision than comparable high-SR units at any given frequency, particularly at moderate to high SPLs, consistent with previous studies. 3. The phase-locking capabilities of neurons in the cochlear nucleus are considerably more variable than in the auditory nerve. Moreover, the variability itself depends on two distinct measures of phase-locking performance. Most ANFs are capable of phase-locking to frequencies as high as 3-4 kHz. In the cochlear nucleus many unit types do not phase-lock to modulation frequencies > 1 kHz. As a result, phase-locking performance is measured on the basis of two parameters, maximum synchronization, irrespective of stimulus frequency, and the upper frequency limit for significant phase-locking. 4. Cochlear nucleus neurons may be divided into three distinct groups on the basis of maximum synchronization capability. In group 1 are the primary-like (PL) units of the anteroventral division, whose phase-locking capabilities are comparable with those of high-SR ANFs.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Effects of ear plugging on single-unit azimuth sensitivity in cat primary auditory cortex. II. Azimuth tuning dependent upon binaural stimulation.

1. Single-unit recordings were carried out in primary auditory cortex (AI) of barbiturate-anesthetized cats. Observations were based on a sample of 131 high-best-frequency (> 5 kHz), azimuth-sensitive neurons. These were identified by their responses to a set of noise bursts, presented in the free field, that varied in azimuth and sound-pressure level (SPL). Each azimuth-sensitive neuron responded well to some levels at certain azimuths, but did not respond well to any level at other azimuths. 2. Unilateral ear plugging was used to infer each neuron's response to monaural stimulation. Ear plugs, produced by injecting a plastic ear mold compound into the external ear, attenuated sound reaching the tympanic membrane by 25-70 dB. The azimuth tuning of a large proportion of the sample (62/131), referred to as binaural directional (BD), was completely dependent upon binaural stimulation because with one ear plugged, these cells were insensitive to azimuth (either responded well at all azimuths or failed to respond at any azimuth) or in a few cases exhibited striking changes in location of azimuth function peaks. This report describes patterns of monaural responses and binaural interactions exhibited by BD neurons and relates them to each cell's azimuth and level tuning. The response of BD cells to ear plugging is consistent with the hypothesis that they derive azimuth tuning from interaural level differences present in noise bursts. Another component of the sample consisted of monaural directional (27/131) cells that derived azimuth tuning in part or entirely from monaural spectral cues. Cells in the remaining portion of the sample (42/131) responded too unreliably to permit specific conclusions. 3. Binaural interactions were inferred by statistical comparison of a cell's responses to monaural (unilateral plug) and binaural (no plug) stimulation. A larger binaural response than either monaural response was taken as evidence for binaural facilitation. A smaller binaural than monaural response was taken as evidence for binaural inhibition. Binaural facilitation was exhibited by 65% (40/62) of the BD sample (facilitatory cells). Many of these exhibited mixed interactions, i.e., binaural facilitation occurred in response to some azimuth-level combinations, and binaural inhibition to others. Binaural inhibition in the absence of binaural facilitation occurred in 35% (22/62) of the BD sample, a majority of which were EI cells, so called because they received excitatory (E) input from one ear (excitatory ear) and inhibitory (I) input from the other (inhibitory ear). One cell that exhibited binaural inhibition received excitatory input from each ear.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1994

Two separate inhibitory mechanisms shape the responses of dorsal cochlear nucleus type IV units to narrowband and wideband stimuli.

1. The principal cells of the dorsal cochlear nucleus (DCN) are mostly inhibited by best frequency (BF) tones but are mostly excited by broadband noise (BBN), producing the so-called type IV response characteristic. The narrowband inhibitory responses can be explained by the inhibitory influence of interneurons with type II response characteristics. However, it is not clear that all the details of the type IV responses can be accounted for by this neural circuit. In particular, many type IV units are inhibited by band-reject noise (notch noise); type II units tend to be only weakly excited by these stimuli, if at all. In this paper we study the relationships between the narrowband, inhibitory and the wideband, excitatory regimens of the type IV responses and present the case for the existence of a second inhibitory source in DCN, called the wideband inhibitor (WBI) below. 2. Type IV units were studied using pure tones, noise bands arithmetically centered on BF, notch noise centered on BF, and BBN. We measured the rate-level function (response rate as function of stimulus level) for each stimulus. This paper is based on the responses of 28 type IV units. 3. Evidence for low-threshold inhibitory input to type IV units is derived from analysis of rate-level functions at sound levels just above threshold. Notch noise stimuli of the appropriate notch width produce inhibition at threshold in this regime. When BBN is presented, this inhibition appears to summate with excitation produced by energy in the band of noise centered on BF, resulting in BBN rate-level functions with decreased slope and maximum firing rate. A range of slopes and maximal firing rates is observed, but these variables are strongly correlated and they are negatively correlated with the strength of the inhibition produced by notch noise; this result supports the conclusion that a single inhibitory source is responsible for these effects. 4. By contrast, there is a weak (nonsignificant) positive correlation between the strength of the inhibitory effect of notch noise and the slope/maximal firing rate in response to narrowband stimuli, including BF tones. The contrast between this positive nonsignificant correlation and the significant negative correlation mentioned above suggests that more than one inhibitory effect operates: specifically, the type II input is responsible for inhibition by narrowband stimuli and a different inhibitory source, the WBI, produces inhibition by notch stimuli. 5. Several lines of evidence are given to show that type II units cannot produce the inhibition seen with notch noise stimuli.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗
PubMed1997

Intracellular response properties of units in the dorsal cochlear nucleus of unanesthetized decerebrate gerbil.

Intracellular recording experiments on the dorsal cochlear nuclei of unanesthetized decerebrate gerbils were conducted. Acceptable recordings were those in which resting potentials were -50 mV or less and action potentials (APs) were > or = 40 mV. Responses to short-duration tones and noise, and to current pulses delivered via recording electrodes, were acquired. Units were classified according to the response map scheme (types I-IV). Ninety-two acceptable recordings were made. Most units had simple APs (simple-spiking units); nine units had both simple and complex APs, which are bursts of spikes embedded on slow, transient depolarizations (complex-spiking units). Of 83 simple-spiking units, 46 were classified as follows: type I/III (9 units), type II (9 units), type III (25 units), type IV (2 units), and type IV-T (1 unit). One complex-spiking unit was classifiable (a type III unit); six were unclassifiable because of weak acoustic responses. Classifying 39 other simple-spiking units and 2 complex-spiking units was impossible, because they were either injured or lost before sufficient data were acquired. Many simple-spiking units showed depolarization or hyperpolarization (approximately 5-10 mV) during acoustic stimulation; some were hyperpolarized during the stimulus-off period. Type I/III units were not hyperpolarized during off-best-frequency (off-BF) stimulation. In contrast, many type II units were hyperpolarized by off-BF frequencies, suggesting that they received strong inhibitory sideband inputs. When inhibited, some type III units were hyperpolarized. Type IV units were hyperpolarized during inhibition even at low levels (<60 dB SPL); sustained depolarizations occurred only at higher levels, suggesting that they receive strong inhibitory and weak excitatory inputs. Several intracellular response properties were statistically different from those of extracellularly recorded units. Intracellularly recorded type II units had higher thresholds and lower maximum BF-driven and noise-driven rates than their extracellularly recorded counterparts. Type I/III units recorded intracellularly had lower maximum BF-driven rates. Type III units recorded intracellularly had higher maximum noise rates compared with those recorded extracellularly. Weaker acoustic responses most likely result from membrane disruption, but heightened responses may be related to weakened chloride-channel-dependent inhibition due to altered driving forces resulting from KCl leakage. Firing rates of simple-spiking units increased monotonically with increasing levels of depolarizing current pulses. In contrast, many complex-spiking units responded nonmonotonically to depolarizing current injection. The monotonic rate-versus-current curves and the nonmonotonic rate-versus-sound level curves of type IV and III units suggest that the acoustic behavior is the result of extrinsic inhibitory inputs and not due solely to intrinsic membrane properties.

Animals↗
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

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↗
PubMed2005

Key processing precedes emotional categorization of Western music.

To investigate whether key processing precedes the appraisal of valence in music, participants listened to pairs of clips of same or different valence, played either in the same key or one semitone apart. They judged whether the second clip expressed the same emotion as the first one. Our predictions were confirmed: the response times obtained were shorter when both clips were played in the same key than when they were played one semitone apart.

Acoustic Stimulation↗
PubMed1989

Influence of stimulus intensity on AEP components in the 80- to 200-millisecond latency range.

The influence of the intensity of passively perceived 1-kHz tones on the latencies and amplitudes of AEP components in the 80- to 200-ms latency range recorded from frontal, central and temporal electrode locations was investigated in healthy adult subjects. At stimulus intensities from 30 to 70 dB SL the latencies of the frontally and centrally recorded N100 and P175 waves decreased, their amplitudes increased. At stimulus intensities from 70 to 90 dB the latencies of N100 and P175 increased, N100 amplitude declined and P175 amplitude increased at a slower rate than between 30 and 70 dB. The latency of the N140 wave recorded at temporal electrode locations decreased markedly with increasing stimulus intensity.

Adult↗
PubMed1991

Application of phase-induced lateralization to the Stenger test.

The Stenger test was employed to estimate the genuine hearing thresholds in normally hearing volunteer subjects simulating a total unilateral loss. The test was carried out in its standard form and in a modified form in which a phase shift was introduced between the signal delivered to the two ears, set to produce phase-induced lateralization towards the 'poor' ear. The standard test estimated the thresholds at a mean of 13.5 dB above the true thresholds at five frequencies from 250 Hz to 4 kHz. Thresholds at the different frequencies were compared, and although thresholds were lower for the higher frequencies, the apparent effect of frequency was not statistically significant. The modified test, using a 90 degrees phase shift, was found to enhance the test at 250 and 500 Hz (thresholds estimated at about 7 dB above true values), but not significantly at 1 kHz.

Adult↗
PubMedDate not supplied

Electrocochleographic analysis of the suppression of tinnitus by electrical promontory stimulation.

To investigate the origin, and evaluate the mechanism by which tinnitus is suppressed we performed electrical promontory stimulation (EPS) in 56 patients with tinnitus, and measured the compound action potential (CAP) using electrocochleography before and after EPS. In the group of patients in whom tinnitus was suppressed, the CAP amplitudes increased significantly, whereas the latencies showed no remarkable change. In the group of patients in whom tinnitus was not suppressed, both the CAP amplitudes and latencies exhibited no significant change. These data indicate that the effect on the cochlear nerve plays an important role in the suppression of tinnitus by EPS. The CAP reflects the number of the auditory nerve fibers which discharge synchronously. It is speculated that an increase of the CAP amplitudes is caused by synchronizing discharges of the auditory nerve fibers, and that the mechanism by which EPS suppresses tinnitus may be related to synchronizing these discharges.

Adult↗
PubMed2004

Cross-modal integration of simple auditory and visual events.

Responses are typically faster and more accurate when both auditory and visual modalities are stimulated than when only one is. This bimodal advantage is generally attributed to a speeding of responding on bimodal trials, relative to unimodal trials. It remains possible that this effect might be due to a performance decrement on unimodal ones. To investigate this, two levels of auditory and visual signal intensities were combined in a double-factorial paradigm. Responses to the onset of the imperative signal were measured under go/no-go conditions. Mean reaction times to the four types of bimodal stimuli exhibited a superadditive interaction. This is evidence for the parallel self-terminating processing of the two signal components. Violations of the race model inequality also occurred, and measures of processing capacity showed that efficiency was greater on the bimodal than on the unimodal trials. These data are discussed in terms of a possible underlying neural substrate.

Attention↗