Temporal integration of electrical stimulation of auditory nuclei in normal-hearing and hearing-impaired cat

Temporal integration functions were measured, before and after a sound-induced hearing loss, in 5 cats using trains of electrical pulses applied to auditory nuclei in the brainstem. The 8 stimuli ranged from 1 pulse (0.25 ms duration) to 16 pulses (0.25 ms pulses spaced over 240 ms). The stimuli were applied to inferior colliculus or cochlear nucleus via permanently implanted electrodes. One electrode was tested extensively in each animal to obtain 10 sets of behaviorally-measured electrical detection thresholds counterbalanced across stimuli. The animal was then exposed to a 110 dB SPL, 2 kHz tone for 48 h and pre- and post-exposure audiograms were measured. The mean permanent threshold shift for acoustic stimuli was 48.5 dB. Another 10 thresholds for each of the 8 electrical stimuli were then measured. In the normal hearing animals, the mean slope of the temporal integration function for electrical stimulation was -7.6 dB per factor of 10 pulses. Alternatively, the mean time constant was 139 ms. In the hearing impaired animals, the slope was reduced to -1.5 dB per factor of 10 pulses, which corresponded to a mean time constant of 17 ms. In addition, the hearing impaired animals showed a decreased threshold for the electrical stimuli (stimulation hypersensitivity) as well as reduced variability across electrical stimulation thresholds. The results suggest that a major contribution to temporal integration occurs in inferior colliculus or higher. In addition, the results suggest that the reduction in temporal integration that follows hearing impairment is a peripherally-induced, central effect.     

Eighth nerve fiber firing features in normal-hearing rabbits

Neural activity of single eighth nerve fibers was recorded with glass microelectrodes in anesthetized normal-hearing rabbits. The units had a spontaneous rate ranging from 0 to approximately 120 spikes/s. In a large number of fibers this rate was below 2 spikes/s. The frequency tuning curves (FTCs) had a tip and a tail region for fibers with a high or medium characteristic frequency (CF). For low-frequency units the FTC was more symmetrically U-shaped. The tip threshold reached the behavioral threshold and units with thresholds of up to more than 60 dB above the mean behavioral threshold were found. There was a weak negative correlation between spontaneous rate and tip threshold. Frequency selectivity, Q10, was about 2 for units with CF below 2 kHz and about 5 for those with CF above 4 kHz. The peristimulus time (PST) histogram showed an initial peak, a plateau, and poststimulus inhibition. For the majority of fibers the dynamic range was 20-30 dB. Some fibers did not reach saturation within the stimulus intensity available. The tip-to-tail distance was 50 dB for high-frequency units at one octave below CF, a matter of potential interest for further studies of animals with inner ear lesions.     

Response patterns of auditory nerve fibers during temporary threshold shift

Temporary threshold shifts were studied in chinchillas exposed to noise (octave-band noise centered at 500 Hz, 95 dB SPL, 5 days duration) and the response properties of their auditory nerve fibers were measured. The threshold shifts of the fibers were approximately 35 to 65 dB; these values were equal to or slightly greater than those measured behaviorally. Most units had broad V-shaped tuning curves due to a greater loss in sensitivity near the characteristic frequency (CF) than in the low-frequency tail. In 17% of the units, the thresholds were actually lower in the tail than at CF, so that the tuning curves were W-shaped. The latencies of the fibers were within normal limits in terms of absolute intensity, but shorter than normal in terms of intensity relative to threshold. Other measures such as the spontaneous discharge rate, the discharge rate-intensity functions, and the firing patterns to tone bursts at CF appeared normal. These results indicate that neural response patterns during noise-induced temporary threshold shift are similar to those measured during permanent threshold shift.     

Contribution of Cochlear Compression to Discrimination of Rippled Spectra in On- and Low-frequency Noise

The goal of the study was to assess cochlear compression when rippled-spectrum signals are perceived in noise assuming that the noise might produce both masking and confounding effects. In normal listeners, discrimination between rippled signals with and without ripple phase reversals was assessed in background noise. The signals were band-limited (0.5 oct at a ??6-dB level) rippled noise centered at 2 kHz, with a ripple density of 3.5 oct^?1. The noise (masker) was band-limited nonrippled noise centered at either 2 kHz (on-frequency masker) or 1 kHz (low-frequency masker). The masker was simultaneously presented with the signals. Masker levels at the discrimination threshold were measured as a function of the signal level using the adaptive (staircase) two-alternative forced-choice procedure. For the on-frequency masker, the searched-for function had a slope of 0.98 dB/dB. For the low-frequency masker, the function had a slope of 1.19 dB/dB within a signal level range of 30 to 40 dB sound pressure level (SPL) and as low as 0.15 dB/dB within a signal level range of 70 to 80 dB SPL. These results were interpreted as indicating compression of responses to both the signal and on-frequency masker and no compression of the effect of the low-frequency masker. In conditions when above-threshold signals are presented in simultaneous noise (the masker), cochlear compression manifests to a substantial degree despite possible confounding effects.     

Frequency dependence of synchronization of cochlear nerve fibers in the alligator lizard: evidence for a cochlear origin of timing and non-timing neural pathways

The dependence of synchronization of spike discharges on tone frequency was measured in cochlear nerve fibers of anesthetized alligator lizards at 21 degrees C. Synchronization measures were based on the fundamental component of a Fourier analysis of the instantaneous discharge rate in response to tone bursts. Measurements were obtained from fibers innervating hair cells in both the region of the cochlea that contains a tectorial membrane (tectorial fibers) and the region where hair-cell stereocilia are free-standing in scala media (free-standing fibers). Both rate and synchronization tuning-curves were measured automatically as a function of tone frequency. For tectorial fibers, the shapes of synchronization tuning-curves are roughly similar to the shapes of rate tuning-curves: the characteristic frequencies (CF's) of both curves are approximately equal. For free-standing fibers, the shape of synchronization tuning-curves differ markedly from those of rate tuning-curves. The CF's of synchronization and rate tuning-curves differ - the ranges are 0.2-0.6 kHz and 1-4 kHz, respectively - and the two CF's are uncorrelated. Synchronization filter-functions, which are contours of synchronization index at constant average discharge rate, were measured as a function of tone frequency for both tectorial and free-standing fibers. These synchronization filter-functions have the shapes of lowpass filters. For the populations of tectorial fibers and of free-standing fibers taken separately, these functions are independent of CF. The corner frequency of these functions is 0.50 +/- 0.038 kHz for tectorial fibers and 0.37 +/- 0.037 kHz for free-standing fibers. We conclude that these populations are characterized by different synchronization filters. For free-standing fibers, synchronization filter-functions measured at average driven discharge rates of about 20 and 40 spikes/s do not differ appreciably, and the high-frequency slope is -80 to -115 dB/decade. The results show that tectorial fibers encode timing information for low-level stimuli, whereas free-standing fibers do not. It is proposed that in the alligator lizard, neural pathways that encode timing information originate in the tectorial region and those that encode non-timing information originate in the free-standing region.     

Neural mechanisms in sound detection and temporal summation

The psychophysics and neurophysiology of sound detection in quiet and under noise masking were studied in goldfish. Psychophysical masking is a linear function of masker level. For long duration signals, signal-to-noise ratios (S/N) at threshold are 15.5, 19. and 22.5 dB for 200, 400 and 800 Hz signals, respectively, and is ? 5 dB for a noise signal. Threshold declines with signal duration to about 700 ms. The slopes of the masked temporal summation functions are about unity, indicating that energy is constant at threshold. In quiet however, the slopes are generally less than 0.5, indicating that shorter signals are detected at lower energy. Neural correlates of the masked S/Ns and the slopes of temporal summation functions were sought in the response patterns of single saccular neurons. Rate- and synchronization-intensity functions were obtained for tone and noise signals in quiet and in noise. S/Ns at behavioral threshold correspond closely to those required to raise spike rate just above that evoked by the masker alone, but are well above those required to cause clear synchronization. Therefore, sound detection is probably based on spike rate and not synchronization criteria. The equivalence of behavioral and neural thresholds indicates that the filters used in behavioral sound detection are simply the bandwidths of saccular fibers. A model outlined by Zwislocki which predicts the rate of temporal summation from the rate of growth of neural activity with intensity accounts quite well for the observed slopes of temporal summation functions both in quiet and in noise.     

Frequency specificity of ABR evaluated by tuning curves

A tone on tone simultaneous masking paradigm was used to determine tuning curves of ABR both from the normal and hearing-impaired subjects. ABR tuning curves were constructed to define masker intensity that resulted in a 50% reduction in probe elicited wave V amplitude. The frequency specificity of each probe stimulus was evaluated by Q10, low cut-off slope, high cut-off slope and the maximum masker frequency calculated for the tuning curves. The results were as follows; 1) Q10, low cut-off slope and high cut-off slope increased gradually with the increase in rise time. However, prolongations of the rise time beyond 3 cycles of probe frequency yielded little improvement in Q10, low cut-off slope and high cut-off slope. 2) Q10, low cut-off slope and high cut-off slope for normal-hearing subjects increased gradually with the increase in stimulus frequency or the decrease in stimulus pressure. Maximum masker frequency of the tuning curves was not always equal to the frequency of probe without the 2-kHz. For the 0.5, 1kHz probe, the maximum masker frequency of the tuning curves showed higher values than the frequency of probe. For the 4kHz probe, the maximum masker frequency of the tuning curves showed lower values than the frequency of probe. The results indicate that the tone pip stimuli will allow to assess certain conditions of auditory function at different frequencies, and they show wider frequencies' spread in the cochlea area near stimulus frequencies. 3) For subject with abrupt high-frequency hearing loss (30dB/oct), a pronounced down-ward shift of maximum masker frequency, down-ward shift of high cut-off slope and up-ward shift of low cut-off slope were found when the probe was placed in the region of elevated threshold. For subject with low-frequency hearing loss (25dB/oct), a pronounced up-ward shift of maximum masker frequency, down-ward shift of low cut-off slope were found. Maximum masker frequency, low and high cut-off slope of hearing-impaired subjects were not always equal to those of normal subjects for same probe. Especially the value of the maximum masker frequency shifted to the direction in which the most sensitive frequency was observed in audiogram. The threshold of ABR reflected the cochlea function of the most sensitive area near stimulus frequencies. Greatest predictive error was observed in steeply sloping audiograms.     

Detection of Tones and Their Modification by Noise in Nonhuman Primates

A fundamental function of the auditory system is to detect important sounds in the presence of other competing environmental sounds. This paper describes behavioral performance in a tone detection task by nonhuman primates (Macaca mulatta) and the modification of the performance by continuous background noise and by sinusoidally amplitude modulating signals or noise. Two monkeys were trained to report detection of tones in a reaction time Go/No-Go task using the method of constant stimuli. The tones spanned a wide range of frequencies and sound levels, and were presented alone or in continuous broadband noise (40 kHz bandwidth). Signal detection theoretic analysis revealed that thresholds to tones were lowest between 8 and 16 kHz, and were higher outside this range. At each frequency, reaction times decreased with increases in tone sound pressure level. The slope of this relationship was higher at frequencies below 1 kHz and was lower for higher frequencies. In continuous broadband noise, tone thresholds increased at the rate of 1 dB/dB of noise for frequencies above 1 kHz. Noise did not change either the reaction times for a given tone sound pressure level or the slopes of the reaction time vs. tone level relationship. Amplitude modulation of tones resulted in reduced threshold for nearly all the frequencies tested. Amplitude modulation of the tone caused thresholds for detection in continuous broadband noise to be changed by smaller amounts relative to the detection of steady-state tones in noise. Amplitude modulation of background noise resulted in reduction of detection thresholds of steady-state tones by an average of 11 dB relative to thresholds in steady-state noise of equivalent mean amplitude. In all cases, the slopes of the reaction time vs. sound level relationship were not modified. These results show that macaques have hearing functions similar to those measured in humans. These studies form the basis for ongoing studies of neural mechanisms of hearing in noisy backgrounds.     

Temporal effects in simultaneous pure-tone masking: effects of signal frequency, masker/signal frequency ratio, and masker level

The effect of the temporal relationship between a pure-tone masker and a pure-tone signal in simultaneous masking was investigated in three experiments. The experiments extend previous work by: studying the temporal effect over a wide range of signal frequencies, studying the change in masking over time for several masker/signal frequency ratios, and studying the growth of masking for a brief signal at different temporal positions within a longer duration masker. In the first experiment, threshold was measured for a 20-ms signal temporally centered in a masker whose duration ranged from 20 ms to continuous. Signal frequency (fs) was 0.5, 1.0, 2.0, 4.0, or 8.0 kHz; masker frequency (fm) was 1.2 fs. For all signal frequencies, the amount of masking decreased as masker duration increased. In the second experiment, threshold was measured for a 20-ms, 1.0-kHz signal as a function of the signal's temporal position within a 400-ms masker whose frequency ranged from 1.0 to 1.25 kHz. For all but the 1.0-kHz masker, for which threshold was almost independent of the signal's temporal position, threshold decreased as signal onset was delayed relative to masker onset, but then increased slightly as the signal approached masker offset. In the final experiment, growth-of-masking functions were measured for a 20-ms, 1.0-kHz signal positioned at the beginning, at the temporal center, or at the end of a 400-ms masker whose frequency was 1.20 or 1.25 kHz. The masking functions generally were steepest for a signal at the onset of the masker and, for a given temporal position, steepest for the 1.20-kHz masker.     

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.     

Effects of contralateral noise on measurement of the psychophysical tuning curve

The effects of the addition of contralateral noise on the psychophysical tuning curve (PTC) were examined in subjects with normal hearing. The masking threshold of the tail part of the PTC tended to decrease with the addition of contralateral noise, although the threshold reduction was usually less than 5 dB. On the other hand, the effects of contralateral noise were relatively small around the tip of the PTC contour. Focusing on the effects of contralateral noise on the masking threshold at the tail part of the PTC, the effects of changing the time between initiation of masking the tone and the presentation of the masked probe tone on the threshold reduction at the tail part of the PTC were also observed. The results indicate that the reduction of the masking threshold by the addition of contralateral noise tended to be larger when the presentation of the signal tone was delayed after the onset of the masker. Usually, when the signal tone was presented under conditions of the forward masking paradigm, the reduction of the threshold was most remarkable. Results obtained in the present study are discussed based on the known characteristics of the olivocochlear (OC)-efferent fibers activated by contralateral noise.     

Comparison between local field potentials and unit cluster activity in primary auditory cortex and anterior auditory field in the cat

Multi-unit (MU) activity and local field potentials (LFP) were simultaneously recorded from 161 sites in the middle cortical layers of the primary auditory cortex (AI) and the anterior auditory field (AAF) in 51 cats. Responses were obtained for frequencies between 625 Hz and 40 kHz, at intensities from 75 dB SPL to threshold. We compared the response properties of MU activity and LFP triggers, in terms of characteristic frequency (CF), threshold at CF, minimum latency and frequency tuning-curve bandwidth 20 dB above threshold. On average, thresholds at CF were significantly lower for LFP events than those for MU spikes (4.6 dB for AI, and 3 dB for AAF). Minimum latencies were significantly shorter for LFP events than for MU spikes (1.5 ms in AI, and 1.7 ms in AAF). Frequency tuning curves were significantly broader for LFP events than those for MU spikes (1.0 octave in AI, and 1.3 octaves in AAF). In contrast, the CF was not significantly different between LFP events and MU spikes. The LFP results indicate that cortical neurons receive convergent sub-cortical inputs from a broad frequency range. The sharper tuning curves for MU activity compared to those of LFP events are likely the result of intracortical inhibitory processes.     

Temporal integration in the gerbil: the effects of age, hearing loss and temporally unmodulated and modulated speech-like masker noises

We characterized temporal integration for 2k Hz pure tones with durations between 10 and 1000 ms in young, normal hearing old and old gerbils with a small hearing loss. Thresholds determined in silence increased for durations below 300 ms and were on average more than 10 dB higher for the 10 ms signal than asymptotic thresholds for the long signals. The amount of temporal integration tended to be less in gerbils with hearing loss. Threshold determination was repeated in the same individuals in the presence of speech-like unmodulated and modulated masking noises. Threshold shift due to the maskers was inversely related to the threshold in silence resulting in a reduced inter-individual variability of thresholds in both masking conditions. Thresholds differed systematically between both masker types in a duration dependent fashion. For long signal durations (300 and 1000 ms) thresholds were on average 2dB lower and for the 10 ms signal 1.9 dB higher in the presence of the modulated masker. These differences in threshold obtained with the two maskers were significant. One hypothesis is that long signals can be detected in the troughs of the modulated masker, while peaks interfere with the detection of short signals.     

Effects of submarine engine room steady noise on the compound action potential tuning curves and its relation to cochlear pathology in guinea pigs

Compound Action Potential Tuning Curves (CAP-TC) for tone pip of 2k, 4 kHz were examined in 8 guinea pigs before and after exposure to noise with main energy centered in the range of 0.25-4.0 kHz. CAP-TC was measured with the pure tone simultaneous masking profiles. AP was evoked by tone pip with an intensity of 10 dB above threshold. Masker level producing 40% reduction in AP amplitude was used. Relations between changes in CAP-TC and AP threshold shifts and the pathology of the stereocilia of hair cells were evaluated by surface preparation and SEM observation in 13 ears. After noise exposure, animals with damaged stereocilia showed AP threshold shift of 20-50 dB, deteriorations of CAP-TC, decrease of Q10 dB value, threshold shift of characteristic frequencies (CF) and displacement of CF towards higher frequencies. It showed that stereocilia damage may affect the susceptibility and frequency selectivity of the cochlea. We consider the CAP-TC may be an useful and sensitive index for detecting physiological and pathological conditions of the cochlea.     

The relative role of beats and combination tones in determining the shapes of masking patterns: II. Hearing-impaired listeners

Masking patterns were measured for hearing-impaired subjects with varying degrees of hearing loss. In one set of conditions, three subjects were tested using narrowband noise ('noise') and sinusoidal ('tone') maskers and narrowband noise signals. The maskers had centre frequencies of 0.25, 0.5, 1.0 and 4.0 kHz and levels of 60, 80 and 100 dB SPL. Masking patterns for both the noise and tone maskers showed irregularities ('dips'), especially for signal frequencies up to 500 Hz above the masker frequency. The irregularities occurred for all masker levels and for all subjects for at least one masker frequency and they occurred for a relatively constant range of masker-signal frequency separations, suggesting that they were the result of beat detection. In another set of conditions, masking patterns were measured using two subjects, for a 2.0-kHz tone masker with a level of 100 dB SPL and tone and noise signals. For the tone masker alone (baseline condition), the masking patterns again exhibited prominent dips above, and sometimes below, the masker frequency. The addition of a lowpass noise to the masker, intended to mask combination tones, had little effect for one subject. For the other subject, who had near-normal absolute thresholds at low frequencies, the noise elevated thresholds for masker-signal frequency separations between 500 and 1500 Hz. For this subject, an extra tone with a frequency equal to the masker-signal frequency separation, added in place of the lowpass noise, had a very similar effect to that produced by the lowpass noise, suggesting that he was detecting a simple difference tone in the baseline condition. The addition of a pair of high-frequency tones (MDI tones - intended to reduce the detectability of beats) to the masker elevated thresholds for signal frequencies from 1500 to 2500 Hz for one subject and from 1500 to 3500 Hz for another subject. The addition of lowpass noise and MDI tones to the masker produced masking patterns very similar to those observed when the MDI tones alone were added to the masker. Overall, the results suggest that the irregularities in the masking patterns were caused mainly by the detection of beats and not by the detection of combination tones.     

正常人中枢掩蔽测试结果分析

目的 :通过对一组耳科正常青年人进行中枢掩蔽测试 ,探讨其发生机制及其对听力学检查所产生的影响。方法 :分别测试有无对侧掩蔽声 (CAS)时的纯音气导听阈 ,比较其在相同频率不同强度的CAS时听阈的变化 ,即中枢掩蔽效应 (CME)的大小 ;同时测试相同强度不同频率CAS时CME的大小。结果 :CME具有频率特异性及强度特异性 ,中频 1kHz、2kHz时 ,其CME较低频及高频大 ,其中以频率为 2kHz时最明显 ;CAS强度在6 0dBHL以下时 ,CME随强度增加而增大 ,在 6 0dBHL时 ,2kHz处CME达 (11.5 3± 4 .38)dB ,CAS >70dBHL时 ,则出现过度掩蔽。结论 :因CME的存在 ,纯音听阈测试中 ,如所用CAS >4 0dBHL时 ,则需对测试结果进行修正 ;当CAS强度为 6 0dBHL时 ,CAS在 2kHz处产生大于 10dB的CME ,可用于对伪聋的鉴别     

Masking of short probe sounds by tone bursts with a sweeping frequency

The maske threshold of short ($\text{? 40}\text{ms}$) probe tone or a short one-third octave probe noise appears to increase if the frequency of a tonal masker is swept. Frequency sweeps were exponential (octaves/s) and unidirectional. Probe sounds were presented in the time center of the masker at the center frequency of the masker. The sweep speed, S, appeared to be an important parameter; masker duration was much less important. For 10-ms tonal probes, in-phase with the masker in their common time center, 100-ms maskers, and upward sweeps, increase of the masked threshold appeared to be maximal at a sweep speed of 30 oct/s and the masked threshold was 21 dB higher than the masked threshold found for the stationary masker (S = 0 oct/s). Above 30 oct/s the masked threshold decreased. For downward sweeps masking was maximal at S = 20 oct/s and the threshold was 15 dB higher. Sweeping upward the increase in masking was 12 dB for both noise probes and tonal probes with phase differing by 90° from the masker in their common time center. The results are inconsistent with current models of masking: sweeping the frequency the masked threshold increases whereas the energy within the critical band at the probe frequency decreases.     

Asymmetry of masking in the European starling: behavioural auditory thresholds

Psychophysical studies of simultaneous masking with human observers exhibit an asymmetry in the amount of masking that depends on the relative bandwidths of signals and maskers. For noise bands up to the bandwidth of one auditory filter, masked auditory thresholds are considerably lower when the bandwidth of the signal exceeds that of the masker compared to the reversed condition. We investigate asymmetry of masking in an animal model, that will allow to study the mechanisms associated with the asymmetry of masking effect. European starlings (Sturnus vulgaris) were trained in a Go/NoGo paradigm to report the detection of a 500 ms noise signal centred in a 700 ms noise masker. Signals and maskers with centre frequencies of 2 kHz had bandwidths of 4 Hz or 256 Hz. Thresholds for detecting the 256 Hz wide-band signal in a 4 Hz narrow-band masker were considerably lower compared to detecting the 4 Hz narrow-band signal in a 256 Hz wide masker and compared to all other conditions. The asymmetry of masking in starlings was on average 15 and 17 dB for 40 and 70 dB SPL overall masker level, respectively. Our animal model thus proved perceptual abilities similar to human subjects. The results are discussed with respect to the importance of both intensity and temporal cues for signal detection.     

Physiology of the young adult Fischer 344 rat inferior colliculus: responses to contralateral monaural stimuli

This study was designed to establish the young adult (3 month) Fischer 344 (F344) rat as a model of inferior colliculus (IC) physiology, providing a baseline for analysis of changes in single unit responses as the animals age and for the study of noise induced hearing loss. The response properties of units localized to the central nucleus of the IC (CIC) and those localized to the external cortex of the IC (ECIC) were compared in order to better characterize differences between these two subnuclei in the processing of simple auditory stimuli. In vivo extracellular single unit recordings were made from IC neurons in ketamine/xylazine anesthetized young adult F344 rats. When a unit was electrically isolated, the spontaneous activity level, characteristic frequency (CF) and CF threshold were determined. Rate/intensity functions (RIFs) in response to contralateral CF tones and to contralateral noise bursts were obtained as were tone isointensity functions. The recording site was marked by ejecting horseradish peroxidase (HRP) from an electrode. Locations of recorded units were determined from electrode track marks and HRP marks in serial brain sections. Recordings were made from 320 neurons in the IC; 176 were localized to the CIC and 87 to the ECIC. Thirteen percent of the units in each subdivision were found to be poorly responsive to auditory stimulation (clicks, tones or noise), and spontaneous activity was generally low. Characteristic frequencies representative of the full rat audiogram were found in each subdivision with the mean threshold significantly higher in the ECIC (28.7 dB SPL) than in the CIC (22.3 dB SPL). The mean maximum discharge rate to CF tone bursts was near 24 spikes/s in each subdivision. Dynamic range tended to be higher in the ECIC (28.3 dB) than in the CIC (23.2 dB), reflecting the lower percentage of nonmonotonic units found in the ECIC. Most units responded more robustly with a slower tone presentation rate, displayed lower levels of discharge to noise bursts than to tone bursts, and had differently shaped tone and noise RIFs. Most units were classified as onset responders to CF tone bursts in both subdivisions, with the percentage of onset responders higher in the ECIC (68.9%) than in the CIC (57.8%). First spike latency did not differ significantly between the subdivisions, but tended to be shorter in the CIC. The breadth of the excitatory receptive fields did not differ significantly between subdivisions, although the mean was slightly larger in the ECIC. These results are generally consistent with the results of CIC studies from other species, establishing the F344 rat as a model of CIC physiology. Differences between CIC and ECIC units included a higher percentage of nonmonotonic RIFs and lower percentage of onset temporal response patterns in the CIC than in the ECIC. Some properties which have been previously used as hallmarks for differentiation between CIC and ECIC units, namely broader tuning and longer first spike latencies in the ECIC, did not reach statistical significance in this study. These may reflect species differences and/or the highly variable and largely overlapping sets of responses evident in the large sample size used in this study.     

Effects of masker frequency and duration in forward masking: further evidence for the influence of peripheral nonlinearity

Forward masking has often been thought of in terms of neural adaptation, with nonlinearities in the growth and decay of forward masking being accounted for by the nonlinearities inherent in adaptation. In contrast, this study presents further evidence for the hypothesis that forward masking can be described as a linear process, once peripheral, mechanical nonlinearities are taken into account. The first experiment compares the growth of masking for on- and off-frequency maskers. Signal thresholds were measured as a function of masker level for three masker-signal intervals of 0, 10, and 30 ms. The brief 4-kHz sinusoidal signal was masked by a 200-ms sinusoidal forward masker which had a frequency of either 2.4 kHz (off-frequency) or 4 kHz (on-frequency). As in previous studies, for the on-frequency condition, the slope of the function relating signal threshold to masker level became shallower as the delay between the masker and signal was increased. In contrast, the slopes for the off-frequency condition were independent of masker-signal delay and had a value of around unity, indicating linear growth of masking for all masker-signal delays. In the second experiment, a broadband Gaussian noise forward masker was used to mask a brief 6-kHz sinusoidal signal. The spectrum level of the masker was either 0 or 40 dB (re: 20 μPa). The gap between the masker and signal was either 0 or 20 ms. Signal thresholds were measured for masker durations from 5 to 200 ms. The effect of masker duration was found to depend more on signal level than on gap duration or masker level. Overall, the results support the idea that forward masking can be modeled as a linear process, preceded by a static nonlinearity resembling that found on the basilar membrane.