Search

Responses of neurons to binaural, harmonic complex stimuli in urethane-anesthetized guinea pig inferior colliculus (IC) are reported. To assess the binaural integration of harmonicity cues for sound segregation and grouping, responses were measured to harmonic complexes with different fundamental frequencies presented to each ear. Simultaneously gated harmonic stimuli with fundamental frequencies of 125 Hz and 145 Hz were presented to the left and right ears, respectively, and recordings made from 96 neurons with characteristic frequencies >2 kHz in the central nucleus of the IC. Of these units, 70 responded continuously throughout the stimulus and were excited by the stimulus at the contralateral ear. The stimulus at the ipsilateral ear excited (EE: 14%; 10/70), inhibited (EI: 33%; 23/70), or had no significant effect (EO: 53%; 37/70), defined by the effect on firing rate. The neurons phase locked to the temporal envelope at each ear to varying degrees depending on signal level. Many of the cells (predominantly EO) were dominated by the response to the contralateral stimulus. Another group (predominantly EI) synchronized to the contralateral stimulus and were suppressed by the ipsilateral stimulus in a phasic manner. A third group synchronized to the stimuli at both ears (predominantly EE). Finally, a group only responded when the waveform peaks from each ear coincided. We conclude that these groups of neurons represent different "streams" of information but exhibit modifications of the response rather than encoding a feature of the stimulus, like pitch., (Copyright © 2015 the American Physiological Society.)

Psychophysical studies show a slower response to changes in the specifically binaural input than to changes in the monaural input (binaural sluggishness). However, there is disagreement about the time course. Tracking changes in a target yields fast time constants, while detecting a constant target against a varying background yields the slowest. Changes in the binaural properties of a target are tracked up to high rates by cells in the midbrain. Indeed cells respond rapidly to a step change and then the firing rate slowly adapts. These experiments, though, are analogues of psychophysical experiments that give the faster time constants. Sluggishness should be more apparent physiologically in a binaural masking paradigm, detecting a short tone in a noise masker with a step change in masker correlation: the small change in firing rate due to the signal must be detected against the adapting firing rate change caused by the step change in the masker. However, in 40 inferior colliculus cells in the anesthetized guinea pig, in a direct analogue of the psychophysical masking paradigm, measuring thresholds for short tones across a transition in a binaural masker (e.g., from N0S0 to NpiS0) provided little evidence of sluggishness within individual cells despite masking level differences in these cells comparable with previous data. Previous studies of physiological correlates of binaural masking level difference suggested that different psychophysical thresholds arise from different populations of cells. This suggests the hypothesis that sluggishness may result from a change in focus between the different populations of cells signaling threshold in different binaural configurations rather than within the intrinsic properties of the cells themselves.

One of the fundamental questions of auditory research is how sounds are segregated because, in natural environments, multiple sounds tend to occur at the same time. Concurrent sounds, such as two talkers, physically add together and arrive at the ear as a single input sound wave. The auditory system easily segregates this input into a coherent perception of each of the multiple sources. A common feature of speech and communication calls is their harmonic structure and in this report we used two harmonic complexes to study the role of the corticofugal pathway in the processing of concurrent sounds. We demonstrate that, in the inferior colliculus (IC) of the anesthetized guinea pig, deactivation of the auditory cortex altered the temporal and/or the spike response to the concurrent, monaural harmonic complexes. More specifically, deactivating the auditory cortex altered the representation of the relative level of the complexes. This suggests that the auditory cortex modulates the representation of the level of two harmonic complexes in the IC. Since sound level is a cue used in the segregation of auditory input, the corticofugal pathway may play a role in this segregation.

We report evidence of mode-locking to the envelope of a periodic stimulus in chopper units of the ventral cochlear nucleus (VCN). Mode-locking is a generalized description of how responses in periodically forced nonlinear systems can be closely linked to the input envelope, while showing temporal patterns of higher order than seen during pure phase-locking. Re-analyzing a previously unpublished dataset in response to amplitude modulated tones, we find that of 55% of cells (6/11) demonstrated stochastic mode-locking in response to sinusoidally amplitude modulated (SAM) pure tones at 50% modulation depth. At 100% modulation depth SAM, most units (3/4) showed mode-locking. We use interspike interval (ISI) scattergrams to unravel the temporal structure present in chopper mode-locked responses. These responses compared well to a leaky integrate-and-fire model (LIF) model of chopper units. Thus the timing of spikes in chopper unit responses to periodic stimuli can be understood in terms of the complex dynamics of periodically forced nonlinear systems. A larger set of onset (33) and chopper units (24) of the VCN also shows mode-locked responses to steady-state vowels and cosine-phase harmonic complexes. However, while 80% of chopper responses to complex stimuli meet our criterion for the presence of mode-locking, only 40% of onset cells show similar complex-modes of spike patterns. We found a correlation between a unit's regularity and its tendency to display mode-locked spike trains as well as a correlation in the number of spikes per cycle and the presence of complex-modes of spike patterns. These spiking patterns are sensitive to the envelope as well as the fundamental frequency of complex sounds, suggesting that complex cell dynamics may play a role in encoding periodic stimuli and envelopes in the VCN.

The function of the profuse descending innervation from the auditory cortex is largely unknown; however, recent studies have demonstrated that focal stimulation of auditory cortex effects frequency tuning curves, duration tuning, and other auditory parameters in the inferior colliculus. Here we demonstrate that, in an anesthetized guinea pig, nonfocal deactivation of the auditory cortex alters the sensitivity of populations of neurons in the inferior colliculus (IC) to one of the major cues for the localization of sound in space, interaural level differences (ILDs). Primary and secondary auditory cortical areas were inactivated by cooling. The ILD functions of 46% of IC cells changed when the cortex was inactivated. In extreme cases, the ILD functions changed from monotonic to nonmonotonic during cooling and vice versa. Eight percent of the cells became unresponsive after deactivation of the auditory cortex. Deactivation of the cortex has previously been shown to alter the maximum spike count of cells in the IC; the change in normalized ILD functions is shown to be separate from this effect. In some cases, the ILD function changed shape when there was no change in the maximum spike count and in other cases there was no change in the shape of the ILD function even though there was a large change in the maximum spike count. Overall, the sensitivity of the IC neural population to ILD is radically altered by the corticofugal pathway.

Accurate temporal coding of low-frequency tones by spikes that are locked to a particular phase of the sine wave (phase-locking), occurs among certain groups of neurons at various processing levels in the brain. Phase-locked responses have previously been studied in the inferior colliculus and neocortex of the guinea pig and we now describe the responses in the auditory thalamus. Recordings were made from 241 single units, 32 (13%) of which showed phase-locked responses. Units with phase-locked responses were mainly (82%) located in the ventral division of the medial geniculate body (MGB), and also the medial division (18%), but were not found in the dorsal or shell divisions. The upper limiting frequency of phase-locking varied greatly between units (60-1,100 Hz) and between anatomical divisions. The upper limit in the ventral division was 520 Hz and in the medial was 1,100 Hz. The range of steady-state delays calculated from phase plots also varied: ventral division, 8.6-14 ms (mean 11.1 ms; SD 1.56); medial division, 7.5-11 ms (mean 9.3 ms; SD 1.5). Taken together, these measurements are consistent with the medial division receiving a phase-locked input directly from the brain stem, without an obligatory relay in the inferior colliculus. Cells in both the ventral and medial divisions of the MGB showed a response that phase-locked to the fundamental frequency of a guinea pig purr and may be involved in analyzing communication calls.

In the auditory system, some ascending pathways preserve the precise timing information present in a temporal code of frequency. This can be measured by studying responses that are phase-locked to the stimulus waveform. At each stage along a pathway, there is a reduction in the upper frequency limit of the phase-locking and an increase in the steady-state latency. In the guinea pig, phase-locked responses to pure tones have been described at various levels from auditory nerve to neocortex but not in the inferior colliculus (IC). Therefore we made recordings from 161 single units in guinea pig IC. Of these single units, 68% (110/161) showed phase-locked responses. Cells that phase-locked were mainly located in the central nucleus but also occurred in the dorsal cortex and external nucleus. The upper limiting frequency of phase-locking varied greatly between units (80-1,034 Hz) and between anatomical divisions. The upper limits in the three divisions were central nucleus, >1,000 Hz; dorsal cortex, 700 Hz; external nucleus, 320 Hz. The mean latencies also varied and were central nucleus, 8.2 +/- 2.8 (SD) ms; dorsal cortex, 17.2 ms; external nucleus, 13.3 ms. We conclude that many cells in the central nucleus receive direct inputs from the brain stem, whereas cells in the external and dorsal divisions receive input from other structures that may include the forebrain.

We have measured the responses of inferior colliculus neurons in the anesthetized guinea pig to signals which in human psychophysical experiments reveal a release of masking as a result of binaural processing (the binaural masking level difference: BMLD). More specifically we have used diotic tones at 500 Hz (So) masked by noise that is either identical at the two ears (No) or inverted in one ear (Npi). This combination of signals and noise maskers produces a prominent masking release in humans such that the So signal is about 6-12 dB more detectable in the presence of the Npi noise than the No noise. Low-frequency inferior colliculus neurons are sensitive to the interaural delay of the masking noise and generally respond most to the components nearest their best frequency. Since most inferior colliculus neurons have peaks in their delay functions close to zero interaural time delay this means that while No noise is effective in driving the unit, Npi noise is much less effective. As the level of an So tone was progressively increased in the presence of No and Npi noises, the first response could be either an increase or a decrease in the activity due to the noise. However, because Npi generated little or no activity itself, the predominant response to the So tone was an increase in discharge in this condition. Masked thresholds were defined as the point at which the standard separation D (related to the d' of signal detection theory) = 1 in either direction. BMLDs were measured in single neurons and in the majority of units were in a direction consistent with the psychophysical observations irrespective of the direction of the discharge rate change that occurred at threshold. The lowest masked thresholds always occurred at or near the signal frequency of 500 Hz. An average value of the single unit BMLD around 500 Hz was 3.6 dB (NoSo vs. NpiSo) compared with 6.6 dB for the NoSo versus NoSpi BMLD we had previously reported. This lower magnitude is consistent with the hierarchy of human psychophysical BMLDs.

Responses to sound stimuli that humans perceive as moving were obtained for 89 neurons in the inferior colliculus (IC) of urethan-anesthetized guinea pigs. Triangular and sinusoidal interaural phase modulation (IPM), which produced dynamically varying interaural phase disparities (IPDs), was used to present stimuli with different depths, directions, centers, and rates of apparent motion. Many neurons appeared sensitive to dynamic IPDs, with responses at any given IPD depending strongly on the IPDs the stimulus had just passed through. However, it was the temporal pattern of the response, rather than the motion cues in the IPM, that determined sensitivity to features such as motion depth, direction, and center locus. IPM restricted only to the center of the IPD responsive area, evoked lower discharge rates than when the stimulus either moved through the IPD responsive area from outside, or up and down its flanks. When the stimulus was moved through the response area first in one direction and then back in the other, and the same IPDs evoked different responses, the response to the motion away from the center of the IPD responsive area was always lower than the response to the motion toward the center. When the IPD was closer at which the direction of motion reversed was to the center, the response to the following motion was lower. In no case did we find any evidence for neurons that under all conditions preferred one direction of motion to the other. We conclude that responses of IC neurons to IPM stimuli depend not on the history of stimulation, per se, but on the history of their response to stimulation, irrespective of the specific motion cues that evoke those responses. These data are consistent with the involvement of an adaptation mechanism that resides at or above the level of binaural integration. We conclude that our data provide no evidence for specialized motion detection involving dynamic IPD cues in the auditory midbrain of the mammal.

We examined the adequacy of decorrelation of the responses to dichotic noise as an explanation for the binaural masking level difference (BMLD). The responses of 48 low-frequency neurons in the inferior colliculus of anesthetized guinea pigs were recorded to binaurally presented noise with various degrees of interaural correlation and to interaurally correlated noise in the presence of 500-Hz tones in either zero or pi interaural phase. In response to fully correlated noise, neurons' responses were modulated with interaural delay, showing quasiperiodic noise delay functions (NDFs) with a central peak and side peaks, separated by intervals roughly equivalent to the period of the neuron's best frequency. For noise with zero interaural correlation (independent noises presented to each ear), neurons were insensitive to the interaural delay. Their NDFs were unmodulated, with the majority showing a level of activity approximately equal to the mean of the peaks and troughs of the NDF obtained with fully correlated noise. Partial decorrelation of the noise resulted in NDFs that were, in general, intermediate between the fully correlated and fully decorrelated noise. Presenting 500-Hz tones simultaneously with fully correlated noise also had the effect of demodulating the NDFs. In the case of tones with zero interaural phase, this demodulation appeared to be a saturation process, raising the discharge at all noise delays to that at the largest peak in the NDF. In the majority of neurons, presenting the tones in pi phase had a similar effect on the NDFs to decorrelating the noise; the response was demodulated toward the mean of the peaks and troughs of the NDF. Thus the effect of added tones on the responses of delay-sensitive inferior colliculus neurons to noise could be accounted for by a desynchronizing effect. This result is entirely consistent with cross-correlation models of the BMLD. However, in some neurons, the effects of an added tone on the NDF appeared more extreme than the effect of decorrelating the noise, suggesting the possibility of additional inhibitory influences.

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

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

1. The responses of units in the ventral cochlear nucleus in anesthetized guinea pigs have been measured to best-frequency tones, noise bands geometrically centered around the unit best frequency, and noise bands asymmetrically positioned around the best frequency. 2. Each unit isolated was characterized using peristimulus time histograms (PSTHs) to best-frequency tones at 20 and 50 dB suprathreshold, frequency-intensity response areas and rate-versus-level functions in response to best-frequency tones and wideband noise. The data reported here are derived from full analyses of 5 chopper units and 17 onset units. The onsets were divided into onset-I (OnI), onset-L (OnL), and onset-C (OnC) by the criteria described by Winter and Palmer: the PSTHs of OnI units show only an onset response, OnL units respond with a single spike at onset followed by a low level of sustained activity, and OnC units have PSTHs with one to four onset peaks and low levels of sustained discharge. 3. In response to geometrically centered noise bands of constant spectral density, the discharge of chopper units and one OnI unit increased over a relatively narrow range of bandwidths, corresponding to the equivalent rectangular bandwidth calculated from their response area, and then became constant. In contrast, OnL and OnC units showed increases in discharge rate with noise bandwidth over very wide ranges of bandwidth. The growth of the discharge rate with noise bandwidth was approximately linear on double logarithmic axes and therefore could be described by a power function with an exponent of 0.37. This relation held even for noise levels near threshold. 4. When noise bands with constant spectral density (at the input to the earphone) were presented with one edge fixed at the unit's best frequency, the discharge rate of most chopper units and the one OnI unit increased over a narrow range of bandwidths and then became constant. This pattern was observed irrespective of whether the second edge of the noise was progressively increased above, or decreased below, the best frequency. For two of the chopper units, in which lateral inhibitory sidebands could be demonstrated, increasing the noise bandwidth led first to increases and then to decreases in the discharge rate as the noise energy impinged upon the sideband. The chopper units act like energy detectors with a filter corresponding to their single tone response area, but, for some units, with the addition of inhibitory sidebands. 5. For the OnL and OnC units, increasing the noise bandwidth above or below best frequency caused progressive increases in the discharge rate over wide ranges of bandwidth. These increases occurred even for low noise spectral densities. The growth in discharge rate for these onset units was well fitted at all spectral density levels by power functions: one above best frequency and one below. At levels of the noise 40 dB above the unit threshold, the point at which the discharge rate reached 90% of its maximum was, on average, about 2 octaves below best frequency and 1 octave above. For some onset units, changes in the discharge rate were seen as the noise bandwidth was varied over about 14 kHz, which is about one-third of the total frequency hearing range of the guinea pig. 6. The data for onset units is consistent with the hypothesis that onset units in the ventral cochlear nucleus achieve their precision in the temporal domain by integration of the inputs from auditory nerve fibers with a wide range of best frequencies. The range of frequency over which onset units integrate frequency matches that of the inhibitory input to dorsal cochlear nucleus neurons, suggesting a possible role as an inhibitory interneuron.

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.

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)