Response to best-frequency tone bursts in the ventral cochlear nucleus is governed by ordered inter-spike interval statistics.

The spike trains generated by short constant-amplitude constant-frequency tone bursts in the ventral cochlear nucleus of the anaesthetised guinea pig are examined. Spikes are grouped according to the order in which they occur following the onset of the stimulus. It is found that successive inter-spike intervals have low statistical dependence according to information-theoretic measures. This is in contrast to previous observations with long-duration tone bursts in the cat dorsal and posteroventral cochlear nuclei and lateral superior olive, where it was found that long intervals tended to be followed by shorter ones and vice versa. The interval distributions can also be reasonably modelled by a shifted Gamma distribution parameterised by the dead-time and the mean and coefficient of variation of the dead-time corrected ISI distribution. Knowledge of those three parameters for each interval is sufficient to determine the peri-stimulus time histogram and the regularity measures used to classify these neurons.

An exact method of regularity analysis for auditory brainstem neurons (L).

The standard regularity analysis for spike trains in cochlear nucleus neurons evoked by tonebursts first proposed by Bourk is widely used, primarily as one of the criteria for classification of such neurons. It is shown that this procedure does not estimate quite what it is supposed to, and introduces unnecessary noise to its results due to its use of bins. Instead the desired quantities (mean and coefficient of variation of the lengths of all inter-spike intervals in progress as a function of time since stimulus onset) can all be exactly calculated directly from the spike train without the need for data binning. The implications for classification and other studies are discussed.

Rebound depolarization in single units of the ventral cochlear nucleus: a contribution to grouping by common onset?

Simultaneous grouping by common onset time is believed to be a powerful cue in auditory perception; components that start or stop roughly at the same time are judged as far more likely to have originated from the same source. Here we report a simple experiment designed to simulate a complex psychophysical paradigm first described by Darwin and Sutherland [(1984) Grouping frequency components of vowels. When is a harmonic not a harmonic? Quarterly J of Experimental Psychology: Hum Exp Psychol 36(A):193-208]. It is possible to change the perception of the vowel /I/ to /epsilon/ by manipulating the harmonics around the first formant (F1). Increasing the amplitude of one harmonic around F1 caused the perception of the vowel to change from /I/ to /epsilon/. Extending the increased component before the vowel could, however, greatly reduce this change. The role of neural adaptation in this effect was questioned by repeating the experiment but this time using a 'captor' tone which was switched on with the asynchronous harmonic and off when the vowel started. This time the vowel percept did change in a fashion analogous to the effect of an increase in the amplitude of the fourth harmonic (which is close to F1). This effect was explained by assuming that the captor had grouped with the leading portion of the asynchronous component enabling the remainder of the asynchronous component to be grouped with the remainder of the components. We propose a relatively low-level neuronal explanation for this grouping effect: the captor reduces the neural response to the leading segment of the asynchronous component by activating across-frequency suppression, either from the cochlea, or acting via a wideband inhibitor in the ventral cochlear nucleus. The reduction in neural response results in a release from adaptation with the offset of the captor terminating the inhibition, such that the response to the continuation of that component is now enhanced. Using a simplified paradigm we show that both primary-like and chopper units in the ventral cochlear nucleus of the anesthetized guinea pig may show a rebound in excitation when a captor is positioned so as to stimulate the suppressive sidebands in its receptive field. The strength of the rebound was positively correlated with the strength of the suppression. These and other results are consistent with the view that low-level mechanisms underlie the psychophysical captor effect.

Human auditory nerve compound action potentials and long latency responses.

CONCLUSION: The compound action potential (CAP) is followed by a long latency response (LLR), attributable to the post-auricular musculature. The LLR to one pulse may overlap with the CAP to a subsequent one, contributing to the clinically observed reduction in CAP at high pulse rates. OBJECTIVES: To measure refractory and other influences on CAPs in humans and guinea pigs. MATERIALS AND METHODS: CAPs were obtained from humans using trans-tympanic and extra-tympanic electrocochleography and from anaesthetized guinea pigs. Stimuli were single pulses presented at a slow rate, pairs of pulses, and 100 ms pulse trains where the inter-pulse interval alternated between 4 and 6 ms. RESULTS: For single pulses, the CAP shape was similar across species. For pairs of pulses, the CAP to the second pulse was smaller than that to the first, and decreased with increasing inter-pulse interval in a way that was similar across species. For pulse trains, CAPs were observed in response to each pulse in the train for the guinea pigs, but not for humans. For both filtered and unfiltered single pulses, there was a large LLR in humans, but not in guinea pigs, with peaks at latencies of 10-12 and 20-25 ms. Posture affected the LLR in a way consistent with the post-auricular response.

Using genetic algorithms to find the most effective stimulus for sensory neurons.

Genetic algorithms (GAs) can be used to find maxima in large search spaces in a relatively short period of time. We have used GAs in electrophysiological experiments to find the most effective stimulus (MES) for sensory neurons in the cochlear nucleus and inferior colliculus of anaesthetised guinea pigs. The MES is the stimulus that elicits the greatest number of spikes from a unit. We show that GAs provide an effective means of determining the best combination of up to four parameters for sinusoids with amplitude modulation. Using GAs, we have found tuning to modulation frequencies as a function of carrier frequency, sound level and temporal asymmetry. These results demonstrate the suitability of GAs in electrophysical experiments for estimating the position of the most effective stimulus in a specified parameter space.

Dual action of olivocochlear collaterals in the guinea pig cochlear nucleus.

Axons of olivocochlear neurones in the superior olivary complex terminate on hair cells of the cochlea, reducing the sensitivity to sound. These axons also have collateral branches to neurones in the cochlear nucleus, the first processing centre in the brainstem. Anatomical data show that these collaterals terminate mainly in the granule cell area but their precise neuronal targets and the effects they might have are unknown. We have studied the effects of these collaterals in guinea pigs, by electrically stimulating the olivocochlear axons at the floor of the IVth ventricle while recording single neurone responses in the cochlear nucleus. We eliminated the peripheral effects of olivocochlear stimulation either by destruction of the target receptor cells using chronic administration of kanamycin, or by acute perfusion of the cochlea with strychnine, a specific blocker of the postsynaptic receptors. Electrical stimulation of the olivocochlear axons in normal animals caused a variety of effects on cochlear nucleus neurones. In some neurones, there was suppression of spontaneous firing and a reduction in sensitivity to sound, while in others there was an excitatory effect of olivocochlear axon stimulation. When the peripheral olivocochlear action was eliminated, we still found both inhibition and excitation in the cochlear nucleus. These results show that the effects of olivocochlear stimulation on cochlear nucleus responses are not a simple passive reflection of peripheral changes but are a result of complex interactions between peripheral suppression of afferent input and collateral-mediated excitation and possibly also inhibition.

The effect of vestibular nerve section upon tinnitus.

This paper reviews the published evidence regarding the effect of vestibular nerve section upon tinnitus. This is of relevance not only for those performing and undergoing this procedure, but also for those considering the hypothesis that auditory efferent system dysfunction may be influential in tinnitus perception. The auditory medial efferent fibres within the internal auditory canal run within the inferior vestibular nerve, only joining the cochlear nerve at the anastomosis of Oort, a bundle of 1300 fibres running from the saccular branch of the inferior vestibular nerve to the cochlear nerve. Vestibular nerve section procedures therefore section this efferent olivocochlear pathway, and ablate efferent influence upon that cochlear. If auditory efferent dysfunction is involved in tinnitus perception, this ablation might influence the tinnitus status of that patient. A literature search identified 18 papers mentioning tinnitus status after vestibular nerve section, describing the experiences of a total of 1318 patients. The proportion of patients in whom tinnitus was said to be exacerbated postoperatively ranged from 0% to 60%, with a mean of 16.4% (standard deviation 14.0). The proportion of patients in whom tinnitus was unchanged was 17% to 72% (mean 38.5%, standard deviation 15.6), and in whom tinnitus was said to be improved was 6% to 61% (mean 37.2%, standard deviation 15.2). In the majority of patients undergoing this procedure, ablation of auditory efferent input (and thus total efferent dysfunction) to the cochlea was not associated with an exacerbation of tinnitus. The finding of this review is that efferent dysfunction after vestibular nerve section does not consistently worsen tinnitus.

The temporal representation of the delay of iterated rippled noise in the ventral cochlear nucleus of the guinea-pig.

1. We have examined the temporal discharge patterns of single units from the ventral cochlear nucleus (VCN) of anaesthetized guinea-pigs in response to iterated rippled noise (IRN). The pitch range evoked by the stimuli was from 32 to 1000 Hz. 2. Single units were classified into four groups using existing classification schemes: primary-like (PL), onset (O), sustained chopper (CS) and transient chopper (CT). For all unit types the delay of the IRN stimuli was well represented in the all-order interspike interval histograms (ISIHs). 3. A subset of the onset units (onset-chopper, OC) showed a clear preference for some delays of the IRN in their first-order interval statistics. We describe this delay preference as 'periodicity tuning'. The delay at which the pitch estimate was at its maximum was designated its best periodicity. The range of best periodicities for OC units was 3.75-13 ms (between 77 and 267 Hz). 4. The other unit types also showed enhancement of the first-order interval statistics at the delay of the IRN. The range of best periodicities was 1.4-8.8 ms (113-714 Hz) for the CT group, 2.25-10.8 ms (93-444 Hz) for the CS group and 0.5-4.6 ms (217-2000 Hz) for the PL group. 5. The correlation between the maximum interval enhancement observed in response to the IRN stimuli and the peak in the first-order ISIH in response to white noise was 0.81 for OC units, 0.72 for CS units, 0.44 for CT units and -0.15 for PL units. 6. These results demonstrate that all unit types in the VCN can enhance the representation of the delay of IRN using first-order interspike intervals (ISIs) over a range of periodicities. CS and OC units show the greatest range of best periodicities and they are well-suited to encode the delay of IRN in their first-order ISIs for a wide range of pitches.

Physiological correlates of comodulation masking release in the mammalian ventral cochlear nucleus.

Comodulation masking release (CMR) enhances the detection of signals embedded in wideband, amplitude-modulated maskers. At least part of the CMR is attributable to across-frequency processing, however, the relative contribution of different stages in the auditory system to across-frequency processing is unknown. We have measured the responses of single units from one of the earliest stages in the ascending auditory pathway, the ventral cochlear nucleus, where across frequency processing may take place. A sinusoidally amplitude-modulated tone at the best frequency of each unit was used as a masker. A pure tone signal was added in the dips of the masker modulation (reference condition). Flanking components (FCs) were then added at frequencies remote from the unit best frequency. The FCs were pure tones amplitude modulated either in phase (comodulated) or out of phase (codeviant) with the on-frequency component. Psychophysically, this CMR paradigm reduces within-channel cues while producing an advantage of approximately 10 dB for the comodulated condition in comparison with the reference condition. Some of the recorded units showed responses consistent with perceptual CMR. The addition of the comodulated FCs produced a strong reduction in the response to the masker modulation, making the signal more salient in the poststimulus time histograms. A decision statistic based on d' showed that threshold was reached at lower signal levels for the comodulated condition than for reference or codeviant conditions. The neurons that exhibited such a behavior were mainly transient chopper or primary-like units. The results obtained from a subpopulation of transient chopper units are consistent with a possible circuit in the cochlear nucleus consisting of a wideband inhibitor contacting a narrowband cell. A computational model was used to confirm the feasibility of such a circuit.

The responses of single units in the inferior colliculus of the guinea pig to damped and ramped sinusoids.

Temporal asymmetry can have a pronounced effect on the perception of a sinusoid. For instance, if a sinusoid is amplitude modulated by a decaying exponential that restarts every 50 ms, (a damped sinusoid) listeners report a two-component percept: a tonal component corresponding to the carrier and a drumming component corresponding to the envelope modulation period. When the amplitude modulation is reversed in time (a ramped sinusoid) the perception changes markedly; the tonal component increases while the drumming component decreases. The long-term Fourier energy spectra are identical for damped and ramped sinusoids with the same exponential half-life. Modelling studies suggest that this perceptual asymmetry must occur central to the peripheral stages of auditory processing (Patterson and Irino, 1998). To test this hypothesis, we have recorded the responses of single units in the inferior colliculus of the anaesthetised guinea pig. We divided single units into three groups: onset, on-sustained and sustained, based on their temporal adaptation properties to suprathreshold tone bursts at the unit's best frequency. The asymmetry observed in the neural responses of single units was quantified in two ways: a simple total spike count measure and a ratio of the tallest bin of the modulation period histogram to the total number of spikes. Responses were more diverse than those observed with similar stimuli in a previous study in the ventral cochlear nucleus (Pressnitzer et al., 2000). The main results were: (1) The shape of the responses of on-sustained units to ramped sinusoids resembled the shape of the responses to damped sinusoids. This is in contrast to the response shapes in the VCN, which were always similar to the stimulating sinusoid. (2) Units classified as onsets often responded only to the damped stimuli. (3) All units display significant asymmetry in discharge rate for at least one of the half-lives tested and 20% showed significant response asymmetry over all of the half-lives tested. (4) A summary population measure of temporal asymmetry based on total spike count reveals the same pattern of results as that obtained psychophysically using the same stimuli (Patterson, 1994a).

Temporal representation of iterated rippled noise as a function of delay and sound level in the ventral cochlear nucleus.

The discharge patterns of single units in the ventral cochlear nucleus (VCN) of anesthetized guinea pigs were examined in response to iterated rippled noise (IRN) as a function of the IRN delay (which determines the IRN pitch) and the IRN sound level. Delays were varied over five octaves in half-octave steps, and sound levels were varied over a 30- or 50-dB range in steps of 5 dB. Neural responses were analyzed in terms of first-order and all-order inter-spike intervals (ISIs). The IRN quasi-periodicity was preserved in the all-order ISIs for most units independent of unit type or best frequency (BF). A deterioration of the temporal all-order code was found, however, when the neural response was influenced by inhibition. The IRN quasi-periodicity was also preserved in first-order ISIs for a limited range of IRN delays and levels. Sustained Chopper units (CS) in the VCN responded with very regular ISIs when the IRN delay corresponded to the unit's chopping period; i.e., the unit showed an increased proportion of intervals corresponding to the IRN delay (interval enhancement) relative to an equal-level, white-noise stimulation. This interval enhancement has a band-pass characteristic with a peak corresponding to the chopping period. Moreover, for CS units in rate saturation, the chopping period, and thus the interval enhancement to the IRN, did not vary with level. Units classified as onset-chopper also show a band-pass interval enhancement to the IRN stimuli; however, they show more level-dependent changes than CS units. Primary-like (PL) units also show level-dependent changes in their ability to code the IRN pitch in first-order intervals. The range of delays where PL units showed interval enhancement was broader and extended to shorter delays. Based on these findings, it is suggested that CS units may play an important role in pitch processing in that they transform a higher-order interval code into a first-order interval place code. Their limited dynamic range together with the preservation of the temporal stimulus features in saturation may serve as a physiological basis for the perceived level independence of pitch.

The responses of single units in the ventral cochlear nucleus of the guinea pig to damped and ramped sinusoids.

Human listeners hear an asymmetry in the perception of damped and ramped sinusoids; the partial loudness of the envelope component is greater than the partial loudness of the carrier component for damped sinusoids. Here we show that an asymmetry also occurs in the physiological responses of most units in the ventral cochlear nucleus to these same sounds. The activity elicited by damped sinusoids is mainly restricted to the beginning of each envelope period, which is not the case for ramped sinusoids. This can be quantified by computing the ratio of the tallest bin of the modulation period histogram to the total number of spikes (the peak-to-total ratio, p/t). Damped sinusoids produce a higher p/t than ramped sinusoids, which demonstrates physiological temporal asymmetry. It is also the case that ramped sinusoids typically elicit more spikes than damped sinusoids. The physiological asymmetry occurs where the perceptual asymmetry is present. It is maximal at modulation half-lives of 4 and 16 ms, greatly reduced at 1 ms and absent at 64 ms. Different unit types exhibit differing degrees of temporal asymmetry. Onset units produce the greatest p/t asymmetry, primary-like units produce the least asymmetry and chopper units are in-between. With regard to total spike count, the maximal asymmetry occurs with chopper units. If primary-like units are assumed to reflect the activity in primary auditory nerve fibres, then there is enhancement of temporal asymmetry in the ventral cochlear nucleus by both onset and chopper units.

Temporal and mean rate discharge patterns of single units in the dorsal cochlear nucleus of the anesthetized guinea pig.

1. We examined the temporal and mean rate discharge characteristics of 514 single units recorded extracellularly from the dorsal cochlear nucleus (DCN) of anesthetized guinea pigs. A mean rate response area (receptive field) was measured for the majority of units in this study. Each response area was placed in one of seven categories (type I to type V and the intermediate types I/III and IV-T) as defined by previous workers. The shape of the best frequency (BF) rate-level function has been used to aid in the distinction between type IV and type IV-T units, and the classification of type II units is based on their relative response to noise and tone bursts. 2. The threshold of single units was normalized to the cochlear action potential (CAP) threshold (a negative relative threshold indicates that the unit's threshold was more sensitive than the corresponding CAP threshold). There were significant differences (P < 0.05; 1-way analysis of variance--Duncan test) between the mean relative thresholds of type IV units (-12 dB) and those of type I (-6.52 dB), type II (-3 dB), and type I/III units (-4.25 dB). There were also significant differences between the relative thresholds of types III and IV-T and those of types I/III and II. 3. Rate-level functions at a unit's BF were divided into groups according to shape and degree of nonmonotonicity. Six units responded with a decrease in firing rate at all suprathreshold sound levels. However, most units increased their discharge rate over approximately the first 20 dB above BF threshold. Units were further subdivided by the change in slope 20 dB above BF threshold. The majority of units (60%) showed monotonic increases in discharge rate with sound level: some rate-level functions clearly resembled the sloping saturation rate-level functions observed in intermediate-threshold auditory nerve fibers. An unexpected finding was the relatively large number of nonmonotonic rate-level functions (40%). Among a relatively homogenous group of projection neurons (predominantly type IV and pause/build units) with nonmonotonic rate-level functions, the range of "best intensities" (the sound level evoking the highest discharge rate) was < 50 dB. This range of best intensities is narrower than found in higher auditory nuclei. 4. Units were also classified by their temporal activity pattern in response to suprathreshold BF tones. The most common pattern identified is the pause/build pattern (n = 294). This temporal activity pattern has been associated with the principal output neuron of the DCN, the fusiform cell. Our definition of pause/build units includes units with an almost constant steady-state discharge rate. Nonmonotonic rate-level functions were observed in 42% (99 of 233) of pause/build units. A measure of discharge regularity (the SD of the interspike interval/mean interspike interval: coefficient of variation, CV) revealed that the majority (82%) of units classified as pause/build and with steady-state discharge rates > 75 spikes/s (n = 142) were characterized by regular discharge patterns (CV = 0.41 +/- 0.15, mean +/- SD). 5. Units characterized by chopper or onset-type discharges were the next most frequently encountered units. The chopper units (n = 75) showed a regular discharge (CV = 0.39 +/- 0.17) similar to that found in recordings from the ventral division of the cochlear nucleus (VCN). One difference between many chopper units in the DCN compared with those recorded in the VCN was the relatively high value (> 5 ms) of the mean interspike interval (and thus the low steady-state discharge rate). The majority (44 of 59; 75%) of chopper units had monotonic rate-level functions. Onset units (n = 47) may represent several response types, linked by the predominance of discharges in response to stimulus onset, and the majority of onset units reported here bear little resemblance to onset units recorded in the VCN of the guinea pig. Approximately 10% of units did not fit easily into any of th

The temporal window of two-tone facilitation in onset units of the ventral cochlear nucleus.

Effects of a tone, not at the best frequency (BF), on the responses of onset units to BF tones were recorded in the ventral cochlear nucleus of the guinea pig. The off-BF tone was at a fixed non-excitatory sound level. When the two tones were gated simultaneously, a marked threshold facilitation was observed; however, no facilitation was observed if the off-BF tone burst was delayed by 5-10 ms. Facilitation in some units declined and was either absent or only just detectable, when the start of the off-BF tone preceded that of the BF by more than 10 ms. However, the responses of the majority of onset units were facilitated when the off-BF tone preceded the BF tone by as much as 20 ms. Similar results were obtained when the roles of the BF and off-BF tones were reversed. These data suggest that depolarization due to BF inputs is of shorter duration than that due to off-BF inputs.

Frequency extent of two-tone facilitation in onset units in the ventral cochlear nucleus.

1. The frequency threshold curves (FTCs) of 91 single units in the cochlear nucleus of the anesthetized guinea pig were measured using a conventional single-tone paradigm and a two-tone paradigm designed to elucidate the frequency extent of two-tone facilitation in onset units (On). Units were classified according to existing classification schemes into primary-like (n = 3), chopper (n = 23), and three onset groups: OnI (n = 12), OnC (n = 29), and OnL (n = 24). Histological reconstructions show onset units to be widely distributed within the ventral cochlear nucleus in a manner generally consistent with its tonotopic organization. 2. The FTCs of onset units differed in their minimum thresholds, the steepness of their high- and low-frequency cutoffs, and their sharpness of tuning as quantified by the quality factor at 10 dB (Q10dB) above best frequency (BF) threshold values. There was considerable overlap in the sharpness of tuning between onset units and auditory nerve fibers, as indicated by the distribution of Q10dB values in the octave around 10 kHz: onset units had Q10dB values of 3.56 +/- 1.38 (SD), compared with 6.3 +/- 2.48 for auditory nerve fibers. The tuning of chopper units was similar to that of auditory nerve fibers (5.52 +/- 1.46). 3. Seventy-five percent of onset units showed some degree of facilitation (a threshold reduction) when their FTCs were measured in the presence of BF tones 4 dB below BF threshold. The frequency extent of such facilitation was variable, with a maximum of 6 octaves around the BF. In extreme cases facilitation could be measured when the BF tone was as low as 30 dB below BF threshold. 4. In 17% of onset units, suppressive effects were evident, as shown by noncontiguous frequency regions of facilitation. These suppressive effects might be a reflection either of suppression in the auditory nerve input or of a direct inhibitory input to the onset units. The strength of this effect suggests that inhibition is a likely explanation, consistent with the finding in previous morphological studies of profuse synapses with pleomorphic vesicles on multipolar cells. 5. FTCs of chopper and primary-like units measured in the presence of BF tones showed little facilitation. The facilitation that was observed in chopper units was confined to a narrow region around BF and disappeared when the facilitatory tone was lowered to 4 dB below BF threshold. 6. These data support the hypothesis that onset units, but not chopper or primary-like units, receive excitatory inputs from auditory nerve fibers with a wide range of BFs. However, the frequency range of facilitation and the magnitude of the threshold facilitation varied from unit to unit, suggesting that the off-BF inputs from auditory nerve fibers are not evenly distributed or equally effective in all units.

Level dependence of cochlear nucleus onset unit responses and facilitation by second tones or broadband noise.

1. The responses of onset units in the cochlear nucleus of the anesthetized guinea pig have been measured to single tones, two-tone complexes, and broadband noise (BBN; 20-kHz bandwidth). The onset units were subdivided into three groups, onset-I (OnI), onset-L (OnL), and onset-C (OnC), on the basis of a decision tree using their peristimulus time histogram (PSTH) shape and discharge rate in response to suprathreshold best-frequency (BF) tone bursts. 2. PSTHs were constructed from responses either to single tones at a unit's BF or to BBN as a function of level. When sufficient sustained activity could be elicited from the unit, arbitrarily defined as > 100 spikes/s, a coefficient of variation (CV) was calculated; the majority were characterized by a CV that was similar to transient chopper units (0.35 < CV < 0.5). First spike latency decreased monotonically with increasing sound level. For the majority of onset units, the first spike timing was very precise. 3. BF rate-level functions recorded from OnL and OnC units did not show any signs of discharge rate saturation at the highest sound levels we have used (100-115 dB SPL). No systematic relationship was observed between the threshold at BF and the shape of the rate-level function. BBN rate-level functions were typically characterized by higher discharge rates than in response to BF tones. However, for OnI units and a minority of other onset units, there was little difference in the shape of their rate-level functions in response to BF tones or BBN. 4. The threshold of most onset units to BBN was similar to the threshold to a BF tone that had similar overall root-mean-square (RMS) energy. The BBN threshold was, on average, 5.5 dB greater than the BF threshold. This result contrasts with that found in auditory-nerve fibers recorded in the same species, with the use of an identical sound system, where the threshold to BBN was, on average, 19.4 dB higher. The mean threshold difference between BBN and BF tones for a population of chopper units recorded in the same series of experiments was 17.7 dB. The relative thresholds to BBN and BF tones indicated that the bandwidths near the onset units' BF threshold were broader than could be estimated with the use of single tones. Ten units were characterized by bimodal response areas.(ABSTRACT TRUNCATED AT 400 WORDS)

The response of guinea pig auditory-nerve fibers with high spontaneous discharge rates to increments in intensity.

We have re-examined the response of auditory-nerve fibres with high spontaneous discharge rates to increments in intensity as a function of the delay of the increment. In agreement with previous studies, the response measured over a relatively long time window (10 ms), emphasising the properties of short-term adaptation, did not decrease with the delay of the increment. However, the response to an increment in intensity, measured over a short time window (0.64 ms), was significantly larger when the increment was coincident with the stimulus onset than when it was delayed by either 5, 10 or 15 ms.

Intensity coding in low-frequency auditory-nerve fibers of the guinea pig.

Rate-level functions from 130 auditory-nerve fibers with characteristic frequencies below 3.5 kHz have been recorded. The relationship between rate-level function type and other properties of auditory-nerve fibers was similar to that previously reported for a high-frequency population in the same species [characteristic frequencies greater than 8 kHz-Winter et al., Hear. Res. 45, 191-202 (1990)]. To estimate whether there is sufficient information in the changes in discharge rate from auditory-nerve fibers to account for the intensity discrimination performance observed in humans (for 1-kHz tone bursts), the Sachs-Abbas model was modified to produce the variation in rate-level functions found in guinea pig auditory-nerve fibers. The ability of modeled, single auditory-nerve fibers and populations of these fibers to signal differences in intensity has been calculated. Optimal combination of information from those fibers assumed to synapse beneath just one inner hair cell resulted in a discrimination performance that exceeded human psychophysical performance up to sound levels of 113 dB SL. It is concluded from these results that there is more than sufficient information present in the changes in discharge rate of a localized population of guinea pig auditory-nerve fibers to account for the intensity discrimination performance of humans as measured psychophysically.

Temporal responses of primarylike anteroventral cochlear nucleus units to the steady-state vowel /i/.

It has previously been shown that a population of units classified as "primarylike" and located in the ventral cochlear nucleus of the anesthetized guinea pig was unable to signal the position of the higher formant-related peaks (greater than 1.5 kHz) of steady-state vowels in terms of a temporal-place representation [Palmer et al., J. Acoust. Soc. Am. 79, 100-113 (1986)]. In this paper, it is demonstrated that units characterized by a prepotential in their spike waveform and a primarylike post-stimulus time histogram shape can encode the relative position of the formant peaks present in the spectra of steady-state vowels in terms of a temporal-place code. The possible reasons for the differences in the present results and the previous report are discussed.