Temporal synchronization in the primary auditory response in the pigeon

Spike potentials were recorded from single fibres in the auditory nerve of the pigeon. In responses elicited by tonal stimuli, the timing of each spike relative to stimulus waveform was measured and period histograms were constructed. Phase locking of spikes was estimated in terms of a synchronicity index obtained by vector addition within the period histogram. A second measure of synchrony in the spike responses was obtained, that of temporal dispersion. For a population of fibres, vector strength of phase locking decreased for frequencies above 1 kHz, as reported for several other species. Temporal dispersion, however, also decreased with frequency, indicating enhanced temporal synchrony as frequency increased within the bandwidth of phase locking. The upper frequency limit of phase locking appears to depend on irreducible jitter of biological origin in the timing of spikes. For individual fibres, the bandwidth of synchronization of spikes consistently exceeds the response area, covering in addition the areas of suppression adjacent to the response area. Spike trains suppressed by a tonal stimulus become synchronized to that stimulus. Phase angles of synchronized responses systematically change as a function of tone level, when tone frequency is above or below CF, as reported for other avian species. Synchronicity and phase angle intensity functions are quite independent of spike rate intensity functions.     

Enhancement and suppression in the auditory midbrain nucleus (MLD) of the pigeon.

Auditory evoked potentials (AEPs) and spike responses were recorded from the same recording site in the nucleus mesencephalicus lateralis pars dorsalis (MLD) in pigeons with a tungsten microelectrode. Depending on the recording sites within the MLD, enhancement and suppression of the AEPs in response to clicks were observed at particular frequencies of a background continuous pure tone. Post stimulus time histograms (PSTs) of the spike responses, if available in such cases, were recorded from the same position by the same electrode. Suppression of the AEPs always occurred but enhancement occurred in only 21% of the trials. The frequencies of tone bursts that caused maximum AEP were vaguely related to the frequencies of continuous pure tones that elicited maximum suppression of the AEPs in response to clicks. However, enhancement was produced by a continuous pure tone of approximately 1.5 kHz, independent of the frequencies of tone bursts that produced maximum AEPs. Most of the PSTs in such instances showed parallel relations between the spike responses and the amplitudes of the AEPs. The nature of the enhancement and suppression of the click evoked AEPs during continuous pure tones was clearly different from those in recordings from the nucleus magnocellularis, nucleus angularis and Field L in respect to the probability of occurrence of enhancement and suppression.     

Induced suppression in spike responses to tone-on-noise stimuli in the auditory nerve of the pigeon

Spike potentials were recorded from single, afferent fibres in the pigeon auditory nerve. Pure-tone stimuli were presented in quiet and in combination with wide band noise. Presented alone, tones produced tuned response areas; noise generally drove spike rate to well above the spontaneous rate measured in quiet. When presented in combination with noise, tones up to 75 dB SPL at frequencies far from the fibre's response area had no effect on the noise-driven spike rate. As the tone frequency was shifted towards the response area, from above or below CF, suppression of the noise-driven spike rate became stronger until the tone reached the edge of the response area. Suppression of the noise-driven rate was directly proportional to the level of the tone. Within the area of response to the tone, tone-driven spike rates generally were unchanged or variably decreased (occasionally slightly increased) by tone-on-noise stimulation, depending on the relation of the tone frequency to CF and the level of the tone relative to that of the noise. Tuning properties were unaffected. It is suggested that in the pigeon, the suppression of driven spike rate during presentation of combination stimuli, which is common to all fibres, depends on the same mechanism as the suppression of spontaneous firing by tones that is observed in a proportion of fibres (Temchin, A.N. (1988), J. Comp. Physiol. A 163, 99-115; Hill et al., (1989) Hear. Res. 39, 37-48).     

Two-tone suppression, excitation and the after effect in rate responses in auditory nerve fibres in the cat.

Responses were recorded from single, auditory nerve fibres in the anaesthetized cat. Acoustic stimuli consisted of two tones, one of which was at characteristic frequency (CF), the other (the suppressor) was at considerably lower frequency. Tones were presented in simultaneous and sequential configurations. For simultaneous presentations, well-known response properties were observed. The rising limb of the two-tone rate-intensity function closely matched that of the appropriately adapted response to the suppressor tone presented alone. Also, whether strongly suppressed relative to CF-driven rate, or equal to CF-driven rate, rate responses to the two-tone stimuli persisted unchanged when the CF tone was terminated and the suppressor tone continued alone. These results support the hypothesis that the suppressor tone has dual influences, suppressive and excitatory, that are distinct and additive. Peristimulus response histograms confirm in the cat that depression and slow recovery of sensitivity to CF may follow termination of the suppressor tone, as reported for the guinea pig [Hill, K.G. and Palmer, A.R. (1991) Hear. Res. 55, 167-176]. This delay in recovery of normal sensitivity to CF appeared to be directly related to the amount of excitation of the fibre that is attributable to the suppressor tone. A similar, delayed re-establishment of sensitivity also occurred in the response to a tone at CF, presented immediately following excitation by a suppressor tone. However, no delay occurred in the onset of response to the suppressor when preceded by the CF tone.(ABSTRACT TRUNCATED AT 250 WORDS)     

Furosemide has no effect on endocochlear potential and tuning properties of primary afferent fibres in the pigeon inner ear

In the pigeon no influence of high doses (135 mg/kg) of furosemide on endocochlear potential and sound evoked activity in single auditory nerve fibres was found. This finding contrasts strongly to results in mammals.     

Shapes of rate-versus-level functions of primary auditory nerve fibres: test of the basilar membrane mechanical hypothesis.

Rate-versus level functions (RI functions) for characteristic frequency (CF) stimulation were measured from primary auditory nerve fibres from different spontaneous rate categories in the guinea pig cochlea. Attention was focussed on those fibres that showed clear breakpoints in their RI functions (sloping-saturation fibres). A statistical curve fitting procedure to an empirical equation was used to provide a quantitative estimate of the breakpoint position in individual fibres. It was found that, within the limits of reliability of the curve fitting procedure, the breakpoint position was the same in fibres from the same CF regions in any given animal. This result is consistent with the notion that the breakpoint position is determined by global basilar membrane mechanics and not by processes private to each nerve fibre. However, a subgroup of fibres not easily classifiable as sloping-saturation, showed features of their RI functions suggesting that factors other than basilar membrane mechanics could lead to fibre-to-fibre differences in rate-versus-level behaviour.     

Changes in cochlear sensitivity do not alter relative thresholds of different spontaneous rate categories of primary auditory nerve fibres

The CF thresholds of three different spontaneous firing rate categories of single auditory nerve fibres were measured in guinea pigs with widely differing cochlear sensitivities. The CF thresholds for all fibres were compared to the CAP thresholds at the same frequencies. The relationship between single fibre CF threshold and CAP thresholds for the three different fibre categories remained unchanged despite the very wide range of absolute CAP thresholds. The widely ranging CAP thresholds in these animals were probably due to varying degrees of outer hair cell pathology and these data therefore support the notion that in normal animals, the origin of differences in threshold between fibres with different spontaneous rates, lies in the properties of the inner hair cells and/or their synapses with afferent dendrites.     

Local auditory evoked potentials and effects of pure tones on click evoked potentials in the pigeon

Local auditory evoked potentials (AEPs) in the pigeon were recorded from the nucleus magnocellularis (NM), nucleus angularis (NA) and Field L with tungsten microelectrodes. In the NM and NA, AEPs in response to clicks were always suppressed by application of continuous pure tones at specific frequencies as is usual for simultaneous masking. In the NA, frequencies of continuous pure tones which produced maximum suppression and frequencies of tone bursts which elicited maximum response both centered around 0.8 kHz. The NM tended to respond similarly. In Field L, however, amplitudes of the AEPs to clicks were suppressed, enhanced, both suppressed and enhanced, or unaffected by presentation of continuous pure tones at specific frequencies. The frequencies of tone bursts which caused maximum AEP were vaguely related to the frequencies of continuous pure tones which elicited maximum suppression of the AEPs to clicks. On the other hand, enhancement was produced by 1-2 kHz continuous pure tones independent of the frequency of tone bursts that produced maximum AEP. It was concluded that enhancement, suppression and lack of effect in Field L were due to some central neural mechanism.     

Central auditory deficits associated with compromise of the primary auditory cortex

The subject of this study was a 46-year-old female who had suffered a cerebrovascular accident (CVA). Magnetic resonance imaging revealed damage in the area of the distribution of the middle cerebral artery involving most, if not all, of the primary auditory area of the left hemisphere. No auditory problems were noted prior to the CVA; however, following the CVA, the subject reported a number of auditory difficulties. Pure-tone thresholds were normal post-CVA, and performance on speech recognition testing was good in both ears if ample time was provided between a response and the presentation of the next test item. Duration pattern, intensity discrimination, and middle latency response test results were abnormal for both ears, and right ear deficits were evident on an auditory fusion test and two dichotic speech tests (digits and rhymes). This case is significant in that it demonstrates a good correlation between damage to known key auditory regions and central auditory test results.     

Neural correlates of auditory stream segregation in primary auditory cortex of the awake monkey

An important feature of auditory scene analysis is the perceptual organization of sequential sound components, or 'auditory stream segregation'. Auditory stream segregation can be demonstrated by presenting a sequence of high and low frequency tones in an alternating pattern, ABAB. When the tone presentation rate (PR) is slow or the frequency separation (DeltaF) between the tones is small (<10%), a connected alternating sequence ABAB is perceived. When the PR is fast or the DeltaF is large, however, the alternating sequence perceptually splits into two parallel auditory streams, one composed of interrupted 'A' tones, and the other of interrupted 'B' tones. The neurophysiological basis of this perceptual phenomenon is unknown. Neural correlates of auditory stream segregation were examined in A1 of the awake monkey using neuronal ensemble techniques (multiunit activity and current source density). Responses evoked by alternating frequency sequences of tones, ABAB, were studied as a function of PR (5, 10, 20 and 40 Hz). 'A' tones corresponded to the best frequency (BF) of the cortical site, while 'B' tones were situated away from the BF by an amount DeltaF. At slow PRs, 'A' and 'B' tones evoked responses that generated an overall pattern of activity at the stimulus PR. In contrast, at fast PRs, 'B' tone responses were differentially suppressed, resulting in a pattern of activity consisting predominantly of 'A' tone responses at half the PR. The magnitude of 'B' tone response suppression increased with DeltaF. Differential suppression of BF and non-BF tone responses at high PRs can be explained by physiological principles of forward masking. The effect of DeltaF is explained by the hypothesis that responses to tones distant from the BF are more susceptible to suppression by BF tones than responses to tones near the BF. These results parallel human psychoacoustics of auditory stream segregation and suggest a cortical basis for the perceptual phenomenon.     

Masking and suppression in auditory nerve fibers of the goldfish, Carassius auratus.

The responses of single fibers of the auditory nerve of the goldfish (Carassius auratus) were recorded in response to two tones of different duration (20 ms 'signals' and 200 ms 'maskers') presented simultaneously or non-simultaneously. A single tone may produce excitation, adaptation, and suppression in auditory nerve fibers. For fibers with characteristic frequencies (CF) in the 200 to 400 Hz range, frequencies well above CF tend to produce suppression. If the net response to the masker tone is excitation, an added excitatory signal tone tends to increment the response in a way predictable from the rate-level function for the masker. A masker can attenuate the response to a signal as a result of a compressive and saturating response to the masker, and as a result of a low signal-to-masker ratio. If the net response to a masker tone is suppression, it effectively subtracts from signal excitation, causing 'suppressive masking.' In non-spontaneous fibers, suppression, additive excitatory effects, and adaptation can be revealed by responses to the signal in the absence of spike responses to the masker. In general, the ability of one tone (the masker) to reduce the response to a second tone (the signal) is greater in non-spontaneous fibers than in spontaneous fibers. These results also show that estimates of the frequency selectivity of many goldfish auditory nerve fibers will depend on whether the response of the fiber is defined by excitation, suppression, or both. The response of many fibers with CF in the 200-400 Hz region, as defined by excitation, can be masked or suppressed by a broad range of frequencies covering the effective hearing range of the goldfish.     

Diversity of characteristic frequency rate-intensity functions in guinea pig auditory nerve fibres

Rate-intensity functions at characteristic frequency (CF) were recorded from single fibres in the auditory nerve of anaesthetised guinea pigs. Within the same animal, CF rate-intensity functions, although probably forming a continuum, could be conveniently divided into three groups; (1) Saturating; reach maximum discharge rate within 30 dB of threshold, (2) Sloping-saturation; initially rapid growth in discharge rate leading to a slower growth in discharge rate but not saturating and (3) Straight; approximately constant increase in firing rate per decibel increase in sound pressure up to the maximum sound pressures used. Thresholds for individual fibres were plotted relative to compound action potential thresholds at the appropriate frequency. Fibres with straight CF rate-intensity functions had the highest thresholds. Fibres of the saturating CF sloping-saturation CF rate-intensity type had thresholds intermediate between saturating and straight. There was a close relationship between the type of CF rate-intensity function exhibited by a fibre and its spontaneous discharge rate. Fibres with saturating CF rate-intensity functions generally had high spontaneous discharge rates (greater than 18/s), whereas those with straight CF rate-intensity functions generally had low spontaneous discharge rates (less than 0.5/s). The majority of fibres with sloping-saturation CF rate-intensity functions had spontaneous rates between 0.5/s and 18/s. There was a negative correlation (r = -0.59) between the logarithm of the spontaneous discharge rate and relative threshold at CF with the lowest spontaneous rate fibres having the highest thresholds and vice-versa. This diversity of CF rate-intensity functions has functional implications for both frequency and intensity coding at high sound pressures in the mammalian auditory system.     

Frequency threshold curves and simultaneous masking functions in single fibres of the guinea pig auditory nerve

Tuning curves for simultaneous masking were measured electrophysiologically, in single fibres of the guinea pig auditory nerve. The masking paradigm used was an analogy of that used in the determination of psychophysical tuning curves in man. The resulting tuning curves ran nearly parallel to the corresponding neural frequency threshold curve, over all except the high-frequency portion of the tuning curve. There, the masking functions had a shallower slope than the excitatory frequency threshold curve. The frequency at which the slope became shallower had a close and consistent dependence on fibre characteristic frequency, reaching a value of 1.2 times fibre characteristic frequency in high-frequency fibres. Analysis of firing rates during the threshold determination gave information about the mechanism of the masking, and the results were supported by theoretical analysis. The results give information useful for the interpretation of psychophysical tuning curves, determined by simultaneous masking, in man.     

Interhemispheric suppression: a test of central auditory function

Two groups of brain damaged adults, those with cerebrocranial injury (CCI) and victims of cerebrovascular accident (CVA) were tested with the competing sentences test of Willeford (In: Central Auditory Dysfunction. New York: Grune and Stratton, 1977: Chap 2). The main purpose was the exposure of functional disorders of communication in such patients who test within normal or near-normal limits on routine audiological testing and also on tests for aphasia. Of 142 serially admitted CCI patients, 43%, many with diffuse lesions, gave abnormal responses, mostly at the left ear, with 19 of them showing total extinction at that ear. On two sound field competing message tests the CCI patients were significantly poorer than normal controls. CVA patients with confirmed right hemisphere lesions tended strongly toward left ear suppression, while left hemisphere damaged patients often showed bilateral drop in scores. It is clear that brain damaged patients can have central dysfunction of auditory processing despite normal findings on routine audiological tests and tests for aphasia. The implications for activities of daily living and for habilitation are discussed.     

Distortion product otoacoustic emission suppression in subjects with auditory neuropathy

OBJECTIVE: The objective of this experiment was to address: 1) whether normal efferent system function is required for normal cochlear tuning as measured by distortion product otoacoustic emission (DPOAE) suppression in humans and 2) whether cochlear function, assessed by DPOAE suppression tuning, is normal in a small group of patients with auditory neuropathy. DESIGN: DPOAE suppression tuning curves (STCs) are similar to other physiologic measures of tuning. They are generated by evoking a DPOAE with two simultaneously presented pure tones and then suppressing the distortion product with a third tone of varying frequency and level. In this study, DPOAE STCs were generated with f2 frequencies of 1500, 3000, and 6000 Hz in 15 normal-hearing adults and four subjects with documented auditory neuropathy. Tuning curve width, slope and tip characteristics, as well as rate of suppression growth were measured in each group. Contralateral suppression of otoacoustic emissions (OAEs) was also recorded as an index of medial efferent function. RESULTS: Results show that the four subjects with auditory neuropathy lacked efferent suppression of OAEs. However, these four subjects showed normal estimates of cochlear tuning as measured by DPOAE suppression results. CONCLUSIONS: This finding suggests that normal efferent system function is not required at the time of test for normal DPOAE suppression tuning. It also suggests that cochlear function as evaluated by detailed measures of DPOAE suppression, is normal in these "typical" patients with auditory neuropathy.     

Auditory associative memory and representational plasticity in the primary auditory cortex

Historically, the primary auditory cortex has been largely ignored as a substrate of auditory memory, perhaps because studies of associative learning could not reveal the plasticity of receptive fields (RFs). The use of a unified experimental design, in which RFs are obtained before and after standard training (e.g., classical and instrumental conditioning) revealed associative representational plasticity, characterized by facilitation of responses to tonal conditioned stimuli (CSs) at the expense of other frequencies, producing CS-specific tuning shifts. Associative representational plasticity (ARP) possesses the major attributes of associative memory: it is highly specific, discriminative, rapidly acquired, consolidates over hours and days and can be retained indefinitely. The nucleus basalis cholinergic system is sufficient both for the induction of ARP and for the induction of specific auditory memory, including control of the amount of remembered acoustic details. Extant controversies regarding the form, function and neural substrates of ARP appear largely to reflect different assumptions, which are explicitly discussed. The view that the forms of plasticity are task dependent is supported by ongoing studies in which auditory learning involves CS-specific decreases in threshold or bandwidth without affecting frequency tuning. Future research needs to focus on the factors that determine ARP and their functions in hearing and in auditory memory.