GABAergic inputs shape responses to amplitude modulated stimuli in the inferior colliculus

The inferior colliculus (IC) is an important auditory processing center receiving inputs from lower brainstem nuclei, higher auditory and nonauditory structures, and contralateral IC. The IC, along with other auditory structures, is involved in coding information about the envelope of complex signals. Biologically relevant acoustic signals, including animal vocalizations and speech, are spectrally and temporally complex and display amplitude and frequency variations over time. Certain IC neurons respond selectively over a narrow range of modulation frequencies to sinusoidally amplitude modulated (SAM) stimuli. Responses to SAM stimuli can be measured in terms of discharge rate, with rate plotted against the modulation frequency to generate rate modulation transfer functions (rMTF). A role for the inhibitory neurotransmitter, gamma-aminobutyric acid (GABA), in shaping selective responses to SAM stimuli has been suggested. The present study examined the role of GABA in shaping responses to SAM stimuli in the IC of anesthetized chinchilla. Responses from 94 IC neurons were obtained before, during and after iontophoretic application of the GABA(A) receptor antagonist bicuculline methiodide. Complete responses to SAM stimuli were obtained from 55 extensively tested neurons, displaying band-pass (38) and low-pass rMTFs (17). For neurons showing band-pass rMTFs, GABA(A) receptor blockade selectively increased discharge rate at low modulation frequencies for 14 units, increased discharge near the best modulation frequency for 12 units. For neurons showing low-pass rMTFs, GABA(A) receptor blockade selectively increased discharge rate at low modulation frequencies for nine units. GABA(A) receptor blockade consistently reduced peak modulation gain, producing low-pass gain functions in a subset of IC neurons. In support of previous findings suggesting that selective temporal responses to SAM stimuli are coded in lower brainstem nuclei, temporal responses to SAM stimuli were relatively unaffected by GABA(A) receptor blockade. These findings support a role for GABA in shaping selective rate responses to SAM stimuli for a subset of chinchilla IC neurons.     

Response of the cochlear nucleus neurons of the cat to tone-burst-trains

Although both posteroventral cochlear nucleus (PVCN) and dorsal cochlear nucleus (DCN) are innervated by the descending branch of auditory nerve fibers, their intrinsic morphological organizations are so different that their physiological roles are expected to be different in signal processing. Temporal information coding of acoustic signals in the cochlear nucleus was examined by using stimuli of "tone-burst-trains (TBT)". Responses of cochlear nucleus neurons of anesthetized cats were recorded either intracellularly or extracellularly. Responses of the neurons to TBT stimuli were classified into "adaptive type" and "non-adaptive type". The "adaptive type" neurons were mainly recorded from PVCN. Responses of these neurons to TBT stimuli decayed exponentially, because of short-term adaptation, in the subsequent tone-bursts. These neurons faithfully preserve the adaptative behavior of auditory nerve fibers. On the contrary, the "non-adaptive type" neurons were mainly found in DCN. They showed variety of responses to TBT stimuli including facilitation, disinhibition and inhibition depending on duration and/or interval of tone-bursts. Our results suggest that some "non-adaptive type" neurons, showing facilitative and/or inhibitory responses to TBT stimuli, act as temporal filters that extract temporal information from acoustic signals.     

Modeling Responses in the Superior Paraolivary Nucleus: Implications for Forward Masking in the Inferior Colliculus

A phenomenological model of the responses of neurons in the superior paraolivary nucleus (SPON) of the rodent is presented in this study. Pure tones at the characteristic frequency (CF) and broadband noise stimuli evoke offset-type responses in these neurons. SPON neurons also phase-lock to the envelope of sinusoidally amplitude-modulated (SAM) stimuli for a range of modulation frequencies. Model SPON neuron received inhibitory input that was relayed by the ipsilateral medial nucleus of the trapezoid body from the contralateral model ventral cochlear nucleus neuron. The SPON model response was simulated by detecting the slope of its inhibitory postsynaptic potential. Responses of the proposed model to pure tones at CF and broadband noise were offset-type independent of the duration of the input stimulus. SPON model responses were also synchronized to the envelope of SAM stimuli with precise timing for a range of modulation frequencies. Modulation transfer functions (MTFs) obtained from the model response to SAM stimuli resemble the physiological MTFs. The output of the proposed SPON model provides an input for models of physiological responses at higher levels of the ascending auditory pathway and can also be utilized to infer possible mechanisms underlying gap detection and duration encoding as well as forward masking at the level of the auditory midbrain.     

Amino acid uptake and release in the guinea pig cochlear nucleus after inferior colliculus ablation

This study attempts to determine if the neurons in the guinea pig inferior colliculus that project to the cochlear nucleus could use certain amino acid transmitters. The left inferior colliculus was ablated surgically to destroy projections descending to the cochlear nuclei. Nissl and silver stained sections of the brain stem indicated that this procedure destroyed most of the left inferior colliculus, but spared a small amount of collicular tissue ventrally and rostrally. Six to seven days after the ablation, degenerated fibers were present in the right inferior colliculus, in the left lateral lemniscus, and in the cochlear nucleus, bilaterally. Three to five days after the ablation, the uptake and electrically-evoked release of exogenous, radiolabeled D-aspartate, gamma-aminobutyrate, and glycine were measured in the three major subdivisions of the cochlear nucleus, the anteroventral, posteroventral, and dorsal divisions. These activities were compared to those in unlesioned controls. The ablation did not alter the uptake and release of the amino acids in the dorsal and posteroventral divisions of the cochlear nucleus. However, it lowered slightly (by 10-18%) the uptake and release of gamma-aminobutyrate and glycine in the anteroventral division, although the difference from the control group was not statistically significant. These findings suggest that most of the neurons in the inferior colliculus that project to the cochlear nucleus probably do not use glutamate, aspartate, gamma-aminobutyrate, or glycine as a transmitter. However, the possibility remains that a small proportion of the collicular projections to the anteroventral cochlear nucleus might use gamma-aminobutyrate or glycine as a transmitter.     

Responses of DCN-PVCN neurons and auditory nerve fibers in unanesthetized decerebrate cats to AM and pure tones: analysis with autocorrelation/power-spectrum

We investigated amplitude-modulated (AM) tone encoding behavior of dorsal and posteroventral cochlear-nucleus (DCN and PVCN) neurons and auditory nerve (AN) fibers in decerebrate unanesthetized cats. Some of the modulation transfer functions (MTFs) were narrowly-tuned band-pass functions; these included responses at moderate and high stimulus levels of DCN pause/build-type-III neurons and the following types of DCN and PVCN chopper neurons: chop-S and/or chop-type-I/III. Other MTFs were broad low-pass or complex functions. Chop-T neurons of the DCN and PVCN tended to exhibit low-pass or flat MTFs. The band-pass MTF neurons exhibited intrinsic oscillations (IOs) in responses to AM or pure tones. The IOs, which were detected in autocorrelation functions and power spectra, were closely correlated (r = 0.863) with the best envelope frequency (BEF). All of the AN fibers showed broad low-pass MTFs with some showing a rudimentary peak in the MTF. The MTFs of DCN-PVCN neurons and AN fibers showed, respectively: (1) BEFs ranging 50-500 Hz, and 400-1300 Hz; (2) upper cut-off frequencies ranging 200-1200 Hz, and 1600-3200 Hz. At stimulus levels of 60-85 dB SPL, maximum modulation gains were as high as 12 dB for DCN-PVCN neurons but were limited to below about 0 dB for AN fibers. The median dynamic ranges of DCN and PVCN neurons (51 and 42 dB, respectively) were substantially wider than those of the low and high spontaneous rate AN fibers (30 and 31 dB, respectively). The observation of higher modulation gain, wider dynamic range, and more narrowly-tuned MTF of DCN-PVCN neurons than AN fibers supports the concept that the capabilities to encode dynamic signals are enhanced in DCN-PVCN neurons compared with AN fibers.     

Two-channel recording of auditory-evoked potentials to detect age-related deficits in temporal processing

Auditory brainstem responses (ABRs), and envelope and frequency following responses (EFRs and FFRs) are widely used to study aberrant auditory processing in conditions such as aging. We have previously reported age-related deficits in auditory processing for rapid amplitude modulation (AM) frequencies using EFRs recorded from a single channel. However, sensitive testing of EFRs along a wide range of modulation frequencies is required to gain a more complete understanding of the auditory processing deficits. In this study, ABRs and EFRs were recorded simultaneously from two electrode configurations in young and old Fischer-344 rats, a common auditory aging model. Analysis shows that the two channels respond most sensitively to complementary AM frequencies. Channel 1, recorded from Fz to mastoid, responds better to faster AM frequencies in the 100-700 Hz range of frequencies, while Channel 2, recorded from the inter-aural line to the mastoid, responds better to slower AM frequencies in the 16-100 Hz range. Simultaneous recording of Channels 1 and 2 using AM stimuli with varying sound levels and modulation depths show that age-related deficits in temporal processing are not present at slower AM frequencies but only at more rapid ones, which would not have been apparent recording from either channel alone. Comparison of EFRs between un-anesthetized and isoflurane-anesthetized recordings in young animals, as well as comparison with previously published ABR waveforms, suggests that the generators of Channel 1 may emphasize more caudal brainstem structures while those of Channel 2 may emphasize more rostral auditory nuclei including the inferior colliculus and the forebrain, with the boundary of separation potentially along the cochlear nucleus/superior olivary complex. Simultaneous two-channel recording of EFRs help to give a more complete understanding of the properties of auditory temporal processing over a wide range of modulation frequencies which is useful in understanding neural representations of sound stimuli in normal, developmental or pathological conditions.     

Sensitivity of neurons in the dorsal medullary nucleus of the grassfrog to spectral and temporal characteristics of sound

The responses of 58 dorsal medullary nucleus units to a set of spectrally and temporally structured stimuli were investigated. Responses to tonepips and noise indicated monomodal spectral sensitivities, with diverse response patterns. Phase-locking was strong for frequencies from 0.1 to 0.2 kHz, and in one unit extended up to 0.6 kHz. To clicks, amplitude modulated tonebursts and natural and artificial versions of the mating call various responses were found. Most low-frequency units fired tonically. They showed a non-selective or low-pass rate response to increasing modulation frequency, and a low-pass synchronization behavior to the envelope. A group of mid-frequency units fired phasically and exhibited a band-pass rate characteristic of amplitude modulated tonebursts. Frequently this was combined with a low-pass rate characteristic of click trains. These units hardly responded to the time-reversed mating call, but fired in a time-locked fashion to the pulses of the original mating call, up to a signal-to-noise ratio of 0 dB. This suggests that aspects of pulse envelope and interpulse interval are coded in the dorsal medullary nucleus.     

Spectral time-course analysis of firing patterns in the dorsal cochlear nucleus

Many previous studies of central auditory neurons have involved independent analyses of spectral and temporal response properties. The spectral response analysis is useful for defining the frequency and intensity regions over which a neuron is excited or inhibited. However, the conventional spectral response analysis only defines this distribution for the synaptic polarity (excitation or inhibition) which dominates the duration of the response. PST histograms of dorsal cochlear nucleus neurons however, often exhibit both excitatory and inhibitory (i.e. pause) components. The distribution of these transient pause intervals may in turn be highly dependent on stimulus parameters suggesting that the spectral area of excitation and inhibition, when considered in terms of short time frames, may be time-dependent. We performed a temporal analysis of the spectral response areas of neurons in the rat dorsal cochlear nucleus and present here an example based on a neuron showing distinct pauser and buildup responses in its PST histograms. The resulting analysis yielded a time course of the spectral response area which indicates that the transient periods of inhibition may have the effect of narrowing the bandwidth of excitation during the early portion of stimulation. Possible implications of this time course are discussed in relation to the narrower tuning that cochlear nucleus neurons often display in response to frequency sweeps than to pure tones.     

Temporal Envelope Coding by Inferior Colliculus Neurons with Cochlear Implant Stimulation

Modulations in temporal envelopes are a ubiquitous property of natural sounds and are especially important for hearing with cochlear implants (CIs) because these devices typically discard temporal fine structure information. With few exceptions, neural temporal envelope processing has been studied in both normal hearing (NH) and CI animals using only pure sinusoidal amplitude modulation (SAM) which poorly represents the diversity of envelope shapes contained in natural sounds because it confounds repetition rate and the width of each modulation cycle. Here, we used stimuli that allow independent manipulation of the two parameters to characterize envelope processing by inferior colliculus (IC) neurons in barbiturate-anesthetized cats with CIs. Specifically, the stimuli were amplitude modulated, high rate pulse trains, where the envelope waveform interleaved single cycles (“bursts”) of a sinusoid with silent intervals. We found that IC neurons vary widely with respect to the envelope parameters that maximize their firing rates. In general, pure SAM was a relatively ineffective stimulus. The majority of neurons (60 %) preferred a combination of short bursts and low repetition rates (long silent intervals). Others preferred low repetition rates with minimal dependence on envelope width (17 %), while the remainder responded most strongly to brief bursts with lesser sensitivity to repetition rate (23 %). A simple phenomenological model suggests that a combination of inhibitory and intrinsic cellular mechanisms suffices to account for the wide variation in optimal envelope shapes. In contrast to the strong dependence of firing rate on envelope shape, neurons tended to phase lock precisely to the envelope regardless of shape. Most neurons tended to fire specifically near the peak of the modulation cycle, with little phase dispersion within or across neurons. Such consistently precise timing degrades envelope coding compared to NH processing of real-world sounds, because it effectively eliminates spike timing as a cue to envelope shape.     

Evaluation of two computational models of amplitude modulation coding in the inferior colliculus

Two computational models replicating amplitude-modulation encoding in the inferior colliculus (IC) are presented and compared. Neurons in this nucleus are modeled as point neurons using Mc Gregor equations, and receive depolarizing currents from action potentials delivered by stellate cells (chopper units) in the cochlear nucleus (CN). Stellate cells are modeled using modified Hodgkin-Huxley equations and receive inputs from a peripheral auditory model. The CN models of the two proposed architectures are characterized by an important dispersion of cellular characteristics, and therefore by various cellular best modulation frequencies (BMFs) ranging from 60 to 300 Hz. In contrast with the previous model proposed by [M.J. Hewitt, R. Meddis, A computer model of amplitude-modulation sensitivity of single units in the inferior colliculus, J. Acoust. Soc. Am. 95 (1994) 2145], each IC cell model receives convergent input from stellate cells with various BMFs. This approach assumes therefore minimal constraints on the model architecture and cell characteristics. The two models differ in terms of the neuronal structure of the IC, composed of 1 or 2 layers of point neurons acting as coincidence detectors. Each model is evaluated using two metrics: mean firing rate and modulation gain. Rate and temporal modulation transfer functions (r-MTFs and t-MTFs, respectively) are simulated and compared with physiological data. Simulations reveal that (i) an important dispersion of BMFs in the CN cells providing input to IC cells yields plausible IC cells responses to AM stimuli, (ii) the 2-layer IC structure yields the best approximation of IC responses measured in vivo.     

Neural encoding of amplitude modulation within the auditory midbrain of squirrel monkeys

The neuronal responses to amplitude modulated (AM) sounds were investigated in the auditory midbrain of the squirrel monkey. Sinusoidally modulated tones and noise served as acoustic stimuli. In order to describe the response properties of collicular neurons, Fast-Fourier-Transformation (FFT), a cross-correlation algorithm and spike-rate counts were applied to translate the neuronal reactions into modulation transfer functions. FFT and cross-correlation defined a measure for synchronic- ity of the neuronal discharges with the modulation cycles. All neurons (542) responded selectively to AM-sounds insofar as all displayed a best modulation frequency (BMF). Most of them furthermore had a band-pass-like modulation transfer function, whose center frequencies were mainly between 8 and 128 Hz. Transfer functions obtained by spike-rate showed less selectivity: a relatively great number of neurons did not change their spike rate as a function of modulation frequency.

The results show that encoding of amplitude-modulated sounds occurs to a greater extent via phase locking of discharges than via changes in spike number. In the same way, changing modulation depth is processed: whereas spike rate on average remains constant between 100% and 0% modulation, there is a drastic reduction in synchronicity. No clear relationship was found between a unit's characteristic frequency and BMF; the same applied to BMF and recording place. The results furthermore show that amplitude modulations are encoded selectively in a band pass function in a non-human primate. The midbrain thereby occupies an intermediate position within the pathway from the periphery to the cortex. This form of temporal resolution probably underlies mechanisms caused by the increasing synaptic activity in the course of the pathway. This may indicate adaptation since those modulation frequencies embedded in this species' vocal repertoire fit quite well with the system's tuning properties for amplitude modulation.     

Responses of young and aged rat inferior colliculus neurons to sinusoidally amplitude modulated stimuli

The inferior colliculus (IC) is a processing center for monaural and binaural auditory signals. Many units in the central nucleus of the inferior colliculus (CIC) respond to amplitude and frequency modulated tones, features found in communication signals. The present study examined potential effects of age on responses to sinusoidally amplitude modulated (SAM) tones in CIC and external cortex of the inferior colliculus (ECIC) units in young and aged F344 rats. Extracellular recordings from 154 localized single units of aged (24 month) rats were compared to recordings from 135 IC units from young adult (3 month) animals. SAM tones were presented at 30 dB above threshold. Comparisons were made between CIC and ECIC regarding the percentage of units responding to SAM stimuli, the relationship between SAM responsiveness and temporal response patterns, maximum discharge rates and maximum modulation gains, shapes of rate transfer functions and synchronization modulation transfer functions (MTFs) in response to SAM tones. Sixty percent of units in young and aged rat IC were selectively responsive to SAM stimuli. Eighty-one percent of units classified as onset temporal response patterns were not tonically responsive to SAM stimuli. Median maximum discharge rate in response to SAM tones was 17.6/s in young F344 rats; median maximum modulation gain was 3.85 dB. These measurements did not change significantly with age. Thirty-seven percent of young rat units displayed bandpass MTFs and 53% had lowpass MTFs. There was a significant age-related shift in the distribution of MTF shapes in both the CIC and ECIC. Aged animals showed a lower percentage of bandpass functions and a higher percentage of lowpass functions. Age-related changes observed in SAM coding may reflect an altered balance between excitatory/inhibitory neurotransmitter efficacy in the aged rat IC, and/or possibly a change in the functional dynamic range of IC neurons.     

Glycine immunoreactivity in the brainstem auditory and vestibular nuclei of the guinea pig

An immunohistochemical study was performed on the brainstem of the guinea pig, using a specific antibody against glycine. Glycine-like immunoreactivity was observed in stellate and multipolar neurons in the cochlear nucleus, in the medial and lateral nuclei of the trapezoid body and in the ventromedial periolivary cell group. No immunoreactive neurons were found in the vestibular nuclei. Positive fibre tracts were observed mainly in dorsal acoustic stria and lateral lemniscus. The results are consistent with electrophysiological and anatomical data from the literature concerning the response pattern in the fusiform layer of the dorsal cochlear nucleus and the phenomenon of binaural inhibition in the superior olivary complex.     

Differential Group Delay of the Frequency Following Response Measured Vertically and Horizontally

The frequency following response (FFR) arises from the sustained neural activity of a population of neurons that are phase locked to periodic acoustic stimuli. Determining the source of the FFR noninvasively may be useful for understanding the function of phase locking in the auditory pathway to the temporal envelope and fine structure of sounds. The current study compared the FFR recorded with a horizontally aligned (mastoid-to-mastoid) electrode montage and a vertically aligned (forehead-to-neck) electrode montage. Unlike previous studies, envelope and fine structure latencies were derived simultaneously from the same narrowband stimuli to minimize differences in cochlear delay. Stimuli were five amplitude-modulated tones centered at 576 Hz, each with a different modulation rate, resulting in different side-band frequencies across stimulus conditions. Changes in response phase across modulation frequency and side-band frequency (group delay) were used to determine the latency of the FFR reflecting phase locking to the envelope and temporal fine structure, respectively. For the FFR reflecting phase locking to the temporal fine structure, the horizontal montage had a shorter group delay than the vertical montage, suggesting an earlier generation source within the auditory pathway. For the FFR reflecting phase locking to the envelope, group delay was longer than that for the fine structure FFR, and no significant difference in group delay was found between montages. However, it is possible that multiple sources of FFR (including the cochlear microphonic) were recorded by each montage, complicating interpretations of the group delay.     

Onset Neurones in the Anteroventral Cochlear Nucleus Project to the Dorsal Cochlear Nucleus

Considerable circumstantial evidence suggests that cells in the ventral cochlear nucleus, that respond predominantly to the onset of pure tone bursts, have a stellate morphology and project, among other places, to the dorsal cochlear nucleus. The characteristics of such cells make them leading candidates for providing the so-called wideband inhibitory input which is an essential part of the processing machinery of the dorsal cochlear nucleus. Here we use juxtacellular labeling with biocytin to demonstrate directly that large stellate cells, with onset responses, terminate profusely in the dorsal cochlear nucleus. They also provide widespread local innervation of the anteroventral cochlear nucleus and a small innervation of the posteroventral cochlear nucleus. In addition, some onset cells project to the contralateral dorsal cochlear nucleus.     

Models of Brainstem Responses to Bilateral Electrical Stimulation

A simple, biophysically specified cell model is used to predict responses of binaurally sensitive neurons to patterns of input spikes that represent stimulation by acoustic and electric waveforms. Specifically, the effects of changes in parameters of input spike trains on model responses to interaural time difference (ITD) were studied for low-frequency periodic stimuli, with or without amplitude modulation. Simulations were limited to purely excitatory, bilaterally driven cell models with basic ionic currents and multiple input fibers. Parameters explored include average firing rate, synchrony index, modulation frequency, and latency dispersion of the input trains as well as the excitatory conductance and time constant of individual synapses in the cell model. Results are compared to physiological recordings from the inferior colliculus (IC) and discussed in terms of ITD-discrimination abilities of listeners with cochlear implants. Several empirically observed aspects of ITD sensitivity were simulated without evoking complex neural processing. Specifically, our results show saturation effects in rate–ITD curves, the absence of sustained responses to high-rate unmodulated pulse trains, the renewal of sensitivity to ITD in high-rate trains when inputs are amplitude-modulated, and interactions between envelope and fine-structure delays for some modulation frequencies.     

Auditory cortical projections to the cochlear nucleus in guinea pigs

We used anterograde tracing techniques to examine projections from auditory cortex to the cochlear nucleus in guinea pigs. Following injection of dextrans into the temporal cortex, labeled axons were present bilaterally in the cochlear nucleus. The distribution of boutons within the cochlear nucleus was similar on the two sides. The majority of boutons was usually located on the ipsilateral side. Most of the boutons were located in the granule cell areas, where many small boutons and a few larger, mossy-type endings were labeled. Additional small, labeled boutons were found in all layers of the dorsal cochlear nucleus, with the majority located in the fusiform cell layer. Labeled boutons were also present in the ventral cochlear nucleus, where they were located in the small cell cap as well as magnocellular parts of both posteroventral and anteroventral cochlear nucleus. Similar results were obtained with injections restricted to primary auditory cortex or to the dorsocaudal auditory field. The results illustrate direct cortical projections to the cochlear nucleus that are likely to modulate the activity in a number of ascending auditory pathways.     

Subcortical neural coding mechanisms for auditory temporal processing

Biologically relevant sounds such as speech, animal vocalizations and music have distinguishing temporal features that are utilized for effective auditory perception. Common temporal features include sound envelope fluctuations, often modeled in the laboratory by amplitude modulation (AM), and starts and stops in ongoing sounds, which are frequently approximated by hearing researchers as gaps between two sounds or are investigated in forward masking experiments. The auditory system has evolved many neural processing mechanisms for encoding important temporal features of sound. Due to rapid progress made in the field of auditory neuroscience in the past three decades, it is not possible to review all progress in this field in a single article. The goal of the present report is to focus on single-unit mechanisms in the mammalian brainstem auditory system for encoding AM and gaps as illustrative examples of how the system encodes key temporal features of sound. This report, following a systems analysis approach, starts with findings in the auditory nerve and proceeds centrally through the cochlear nucleus, superior olivary complex and inferior colliculus. Some general principles can be seen when reviewing this entire field. For example, as one ascends the central auditory system, a neural encoding shift occurs. An emphasis on synchronous responses for temporal coding exists in the auditory periphery, and more reliance on rate coding occurs as one moves centrally. In addition, for AM, modulation transfer functions become more bandpass as the sound level of the signal is raised, but become more lowpass in shape as background noise is added. In many cases, AM coding can actually increase in the presence of background noise. For gap processing or forward masking, coding for gaps changes from a decrease in spike firing rate for neurons of the peripheral auditory system that have sustained response patterns, to an increase in firing rate for more central neurons with transient responses. Lastly, for gaps and forward masking, as one ascends the auditory system, some suppression effects become quite long (echo suppression), and in some stimulus configurations enhancement to a second sound can take place.     

Response properties of cochlear nucleus neurons in monkeys

Much of what is known about how the cochlear nuclei participate in mammalian hearing comes from studies of non-primate mammalian species. To determine to what extent the cochlear nuclei of primates resemble those of other mammalian orders, we have recorded responses to sound in three primate species: marmosets, cynomolgus macaques, and squirrel monkeys. These recordings show that the same types of temporal firing patterns are found in primates that have been described in other mammals. Responses to tones of neurons in the ventral cochlear nucleus have similar tuning, latencies, post-stimulus time and interspike interval histograms as those recorded in non-primate cochlear nucleus neurons. In the dorsal cochlear nucleus, too, responses were similar. From these results it is evident that insights gained from non-primate studies can be applied to the peripheral auditory system of primates.     

Neural and Behavioral Sensitivity to Interaural Time Differences Using Amplitude Modulated Tones with Mismatched Carrier Frequencies

Bilateral cochlear implantation is intended to provide the advantages of binaural hearing, including sound localization and better speech recognition in noise. In most modern implants, temporal information is carried by the envelope of pulsatile stimulation, and thresholds to interaural time differences (ITDs) are generally high compared to those obtained in normal hearing observers. One factor thought to influence ITD sensitivity is the overlap of neural populations stimulated on each side. The present study investigated the effects of acoustically stimulating bilaterally mismatched neural populations in two related paradigms: rabbit neural recordings and human psychophysical testing. The neural coding of interaural envelope timing information was measured in recordings from neurons in the inferior colliculus of the unanesthetized rabbit. Binaural beat stimuli with a 1-Hz difference in modulation frequency were presented at the best modulation frequency and intensity as the carrier frequencies at each ear were varied. Some neurons encoded envelope ITDs with carrier frequency mismatches as great as several octaves. The synchronization strength was typically nonmonotonically related to intensity. Psychophysical data showed that human listeners could also make use of binaural envelope cues for carrier mismatches of up to 2–3 octaves. Thus, the physiological and psychophysical data were broadly consistent, and suggest that bilateral cochlear implants should provide information sufficient to detect envelope ITDs even in the face of bilateral mismatch in the neural populations responding to stimulation. However, the strongly nonmonotonic synchronization to envelope ITDs suggests that the limited dynamic range with electrical stimulation may be an important consideration for ITD encoding.