Excitation and suppression of primary auditory fibres in the pigeon

Spike potentials were recorded from single fibres in the auditory nerve of the pigeon. In fibres with recognizable responses to sound, spontaneous activity and properties of responses to tonal stimuli were studied in quiet background conditions. Mean spontaneous rate in the sample of fibres was 35 spikes/s. Tuning of spike response to tones was manifest as a single peak in rate at each sound pressure level (SPL) in the frequency-intensity plane. The majority of fibres showed only excitation of spike rate above spontaneous rate. Post stimulus time histograms (PSTs) in such cases were typical of excitatory responses, previously described in birds and mammals showing pronounced adaptation and post-stimulus suppression of spike rate. In most cases of excitation-only responses, however, slopes of rate functions depended on stimulus frequency. Close to characteristic frequency (CF), slopes tended to decrease with increasing SPL, whereas away from CF, slopes tended to increase with SPL. In a minority of excitation-only responses, slopes of rate functions were parallel. In some fibres, tones adjacent to the response area caused overt suppression of spontaneous firing. For these fibres, the slopes of rate functions were more-strongly frequency-dependent, being negative at low SPL when rate suppression occurred. Suppression of spontaneous activity at low SPL was non-monotonic and quite different from suppression of spike rate at stimulus intensities above rat saturation. In PSTs of suppressed spontaneous activity, rebound occurred at the termination of the tone. The results clarify previous observations of suppression of primary auditory responses in birds. We conclude that responses in the majority of auditory fibres in the pigeon are the product of opposing excitatory and suppressive influences in the cochlea, generated by single tones in quite.     

Temporal envelope processing in the human auditory cortex: response and interconnections of auditory cortical areas

Temporal envelope processing in the human auditory cortex has an important role in language analysis. In this paper, depth recordings of local field potentials in response to amplitude modulated white noises were used to design maps of activation in primary, secondary and associative auditory areas and to study the propagation of the cortical activity between them. The comparison of activations between auditory areas was based on a signal-to-noise ratio associated with the response to amplitude modulation (AM). The functional connectivity between cortical areas was quantified by the directed coherence (DCOH) applied to auditory evoked potentials. This study shows the following reproducible results on twenty subjects: (1) the primary auditory cortex (PAC), the secondary cortices (secondary auditory cortex (SAC) and planum temporale (PT)), the insular gyrus, the Brodmann area (BA) 22 and the posterior part of T1 gyrus (T1Post) respond to AM in both hemispheres. (2) A stronger response to AM was observed in SAC and T1Post of the left hemisphere independent of the modulation frequency (MF), and in the left BA22 for MFs 8 and 16Hz, compared to those in the right. (3) The activation and propagation features emphasized at least four different types of temporal processing. (4) A sequential activation of PAC, SAC and BA22 areas was clearly visible at all MFs, while other auditory areas may be more involved in parallel processing upon a stream originating from primary auditory area, which thus acts as a distribution hub. These results suggest that different psychological information is carried by the temporal envelope of sounds relative to the rate of amplitude modulation.     

Rate and synchronization measures of periodicity coding in cat primary auditory cortex.

Periodicity coding was studied in primary auditory cortex of the ketamine anesthetized cat by simultaneously recording with two electrodes from up to 6 neural units in response to one second long click trains presented once per 3 s. Trains with click rates of 1, 2, 4, 8, 16 and 32/s were used and the responses of the single units were quantified by both rate measures (entrainment and rate modulation transfer function, rMTF) and synchronization measures (vector strength VS and temporal modulation transfer functions, tMTF). The rate measures resulted in low-pass functions of click rate and the synchrony measures resulted in band-pass functions of click rate. Limiting rates (-6 dB point of maximum response) were in the range of 3-24 Hz depending on the measure used. Best modulating frequencies were in the range of 5-8 Hz again depending on the synchrony measure used. It appeared that especially the VS was highly sensitive to spontaneous firing rate, duration of the post click suppression and the size of the rebound response after the suppression. These factors were dominantly responsible for the band-pass character of the VS-rate function and the peak VS frequency was nearly identical to the inverse of the suppression period. It is concluded that the use of the VS and to a lesser extent also the tMTF as the sole measure for the characterization of periodicity coding is not recommended in cases where there is a strong suppression of spontaneous activity. The combination of entrainment and tMTF appeared to characterize the periodicity coding in an unambiguous way.     

Temporal response properties of the auditory nerve: data from human cochlear-implant recipients

The primary goal of this study was to characterize the variability in auditory-nerve temporal response patterns obtained with the electrically evoked compound action potential (ECAP) within and across a relatively large group of cochlear-implant recipients. ECAPs were recorded in response to each of 21 pulses in a pulse train for five rates (900, 1200, 1800, 2400, and 3500 pps) and three cochlear regions (basal, middle, and apical). An alternating amplitude pattern was typically observed across the pulse train for slower rates, reflecting refractory properties of individual nerve fibers. For faster rates, the alternation ceased and overall amplitudes were substantially lower relative to the first pulse in the train, reflecting cross-fiber desynchronization. The following specific parameters were examined: (1) the rate at which the alternating pattern ceased (termed stochastic rate), (2) the alternation depth and the rate at which the maximum alternation occurred, and (3) the average normalized ECAP amplitude across the pulse train (measure of overall adaptation/desynchronization). Data from 29 ears showed that stochastic rates for the group spanned the entire range of rates tested. The majority of subjects (79%) had different stochastic rates across the three cochlear regions. The stochastic rate occurred most frequently at 2400 pps for basal and middle electrodes, and at 3500 pps for apical electrodes. Stimulus level was significantly correlated with stochastic rate, where higher levels yielded faster stochastic rates. The maximum alternation depth averaged 19% of the amplitude for the first pulse. Maximum alternation occurred most often at 1800 pps for basal and apical electrodes, and at 1200 pps for middle electrodes. These differences suggest some independence between alternation depth and stochastic rate. Finally, the overall amount of adaptation or desynchronization ranged from 63% (for 900 pps) to 23% (for 3500 pps) of the amplitude for the first pulse. Differences in temporal response properties across the cochlea within subjects may have implications for developing new speech-processing strategies that employ varied rates across the array.     

Divergent response properties of layer-V neurons in rat primary auditory cortex

Layer-V pyramidal cells comprise a major output of primary auditory cortex (A1). At least two cell types displaying different morphology, projections and in vitro physiology have been previously identified in layer-V. The focus of the present study was to characterize extracellular receptive field properties of layer-V neurons to determine whether a similar breakdown of responses can be found in vivo. Recordings from 105 layer-V neurons revealed two predominant receptive field types. Thirty-two percent displayed strong excitatory V/U-shaped receptive field maps and spiking patterns with shorter stimulus-driven interspike intervals (ISIs), reminiscent of the bursting cells discussed in the in vitro literature. V/U-shaped maps remained relatively unchanged across the three sequential repetitions of the map run on each neuron. Neurons with V/U-shaped maps were also easily depolarized with extracellular current pulse stimulation. In contrast, 47% of the neurons displayed Complex receptive field maps characterized by weak and/or inconsistent excitatory regions and were difficult to depolarize with current pulses. These findings suggest that V/U-shaped receptive fields could correspond to previously described intrinsic bursting (IB) cells with corticotectal projections, and that neurons with Complex receptive fields might represent the regular spiking (RS) cells with their greater inhibitory input and corticocortical/corticostriatal projection pattern.     

Binaural interactions shape binaural response structures and frequency response functions in primary auditory cortex

The overall purpose of this study is to examine the behavior of primary auditory cortex (AI) units in the three-dimensional stimulus space that resembles normal listening conditions, viz., level at the two ears and frequency. A binaural-level response area (LRA) is the response to a matrix of contralateral and ipsilateral stimuli presented at a single frequency. LRAs have been examined in the inferior colliculus and AI and found to be highly organized response patterns that are shaped by binaural interactions. The aggregate of LRAs across frequency is the binaural response structure (BRS), a new concept that captures unit behavior in this three-dimensional stimulus space. Since binaural interactions contribute greatly to configuring component LRAs, it is clear that binaural interactions help shape the aggregate BRS. The BRS contains the data required to generate binaural frequency response functions. The frequency range and magnitude of these functions depend on the level of the stimulus at each ear and the configuration of the BRS. Changing either level can greatly alter the binaural frequency response function. Thus, in addition to their classic role in localization, binaural interactions play a fundamentally important role in determining the frequency domain of units in AI.     

Temporal response patterns of auditory nerve fibers to electrical stimulation in deafened squirrel monkeys

Electroneural response patterns of single auditory-nerve neurons were studied in aminoglycoside-deafened squirrel monkeys. The electrical stimuli were delivered through bipolar electrodes implanted in the scala tympani. The effects of pulse width, shape, frequency, and intensity on neural adaptation, phase locking, and spectral content were evaluated. Our results did not demonstrate the characteristic adaptation seen in auditory-nerve neurons in response to acoustic stimulation. Phase locking to a broad stimulus pulse (3200 microseconds/phase) was found to a very restricted phase angle of the electrical stimulus which was broader for square wave than for sine wave stimulation. The latency of the phase locked response varied inversely with stimulus intensity with greater variation for square wave stimulation than for sine wave stimulation. Auditory neurons were capable of a very high degree of phase locking to a 200-microseconds/phase pulse presented at 156 pulses per second (PPS) and to the first pulse of a 2500-Hz pulse burst. Phase locking was much poorer for the subsequent 200-microseconds/phase pulses comprising the 2500-Hz pulse burst where the neuron's response was determined by its relative recovery status. These findings can be explained by an interaction between the neuron's relative refractory status and its integration of charge over the stimulatory half cycle of the electrical stimulus. These two factors also appear to determine the interspike interval of the neural response. This interval decreased monotonically with increasing stimulus intensity. The neural spike rate (150-500 Hz) producing this interval increased with intensity and may be a source of periodicity information which the central auditory nervous system could interpret as pitch. This may account for the proportional relationship between pitch and stimulus intensity seen in some cochlear implant patients. Our study demonstrates that auditory-nerve neurons comply with basic neurophysiological principles in their responses to electrical stimulation. These principles should be incorporated into the cochlear prosthesis stimulator if more normal neural response patterns are desired in the cochlear prosthesis patient.     

Temporal response patterns of single auditory nerve fibers elicited by periodic electrical stimuli

Single auditory nerve fibers exhibit firing synchronized to one or both phases of periodic AC stimulus currents. Responses to biphasic pulses depend on order and excitation sites of the two phases. Sine and triangle stimuli between 100 Hz and 500 Hz elicit similar response patterns. Responses to square waves are sometimes more synchronized and generally shifted in phase with respect to sine wave responses. Preferred firing phase(s): (1) are largely independent of stimulus intensity; (2) vary among fibers; (3) may shift continuously or discontinuously over several seconds before steady state is achieved. Responses to an unprocessed synthetic vowel stimulus were dominated by pitch period, first formant, and 'spurious' components.     

The auditory steady-state response: comparisons with the auditory brainstem response

Two studies are reported in which the threshold estimates from auditory steady-state response (ASSR) tests are compared to those of click- or toneburst-evoked auditory brainstem responses (ABRs). The first, a retrospective review of 51 cases, demonstrated that both the click-evoked ABR and the ASSR threshold estimates in infants and children could be used to predict the pure-tone threshold. The second, a prospective study of normal-hearing adults, provided evidence that the toneburst-evoked ABR and the modulated tone-evoked ASSR thresholds were similar when both were detected with an automatic detection algorithm and that threshold estimates varied with frequency, stimulus rate, and detection method. The lowest thresholds were obtained with visual detection of the ABR. The studies illustrate that ASSRs can be used to estimate pure-tone threshold in infants and children at risk for hearing loss and also in normal-hearing adults.     

Temporal integration in an anuran auditory nerve

We determined temporal integration in individual auditory nerve fibers of the arboreal frog, Eleutherodactylus coqui by measuring the change in rate threshold to tonal stimulation for tone burst durations from 20 to 450 ms. Temporal integration was quantified in two ways: (1) we calculated the temporal integration time constant of the fiber (tau), and (2) we calculated the shift in threshold per decade increment of the tone stimulus (dB/decade). In some cases, the procedure was repeated using a continuous, broadband noise masker (20-50 dB/Hz). Our results indicate that low frequency fibers (CF less than 0.50 kHz) have the longest mean integration time (274 ms), mid-frequency fibers (0.50 to 1.30 kHz) have the shortest mean integration time (183 ms) and high frequency fibers (CF greater than 1.3 kHz) have an intermediate mean integration time (235 ms). Continuous noise increased the integration times of some, but not all fibers, and caused some fibers which did not display temporal integration to do so. We investigated the possibility that these changes may be caused by a decrease in the slope of the rate-intensity function (measured between +5 and +15 dB re threshold) with the addition of the continuous noise masker. The slope of the rate-intensity function decreased (from 73 spikes/s/dB to 49 spikes/s/dB) with the addition of the continuous noise for those fibers (N = 28) that showed an increase in temporal integration with the addition of noise. However, the slopes of the rate-intensity functions also decreased by 30% for those fibers (N = 8) that did not show increasing temporal integration.     

Comparison of auditory steady-state response and auditory brainstem response thresholds in children

Recently, auditory steady-state responses (ASSRs) have been proposed as an alternative to the auditory brainstem response (ABR) for threshold estimation. The goal of this study was to investigate the degree to which ASSR thresholds correlate with ABR thresholds for a group of sedated children with a range of hearing losses. Thirty-two children from the University of Iowa Hospitals and Clinics ranging in age from 2 months to 3 years and presenting with a range of ABR thresholds participated. Strong correlations were found between the 2000-Hz ASSR thresholds and click ABR thresholds (r = .96), the average of the 2000- and 4000-Hz ASSR thresholds and click ABR thresholds (r = .97), and the 500-Hz ASSR and 500-Hz toneburst ABR thresholds (r = .86). Additionally, it was possible to measure ASSR thresholds for several children with hearing loss that was great enough to result in no ABR at the limits of the equipment. The results of this study indicate that the ASSR may provide a reasonable alternative to the ABR for estimating audiometric thresholds in very young children.     

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

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

The alteration of the low frequency response of primary auditory afferents by cochlear trauma

The response phase of primary auditory afferents in the first turn of the guinea pig cochlea to low frequency sinusoidal stimuli is altered by trauma to the cochlea. Loud sounds of sufficient intensity to produce temporary shifts in the thresholds of these cells at their characteristic frequency produce a reversal of the neural response from increased firing for basilar membrane displacements towards scala tympani before trauma to an increased firing towards scala vestibuli following trauma. These changes can be reversible.     

A comparison of synchronization filters in different auditory receptor organs

Measurements of the frequency dependence of synchronization of cochlear nerve fibers obtained in different auditory receptor organs are compared. These synchronization filter-functions are lowpass filter-functions and differ primarily in corner frequencies which we estimate to be (in kHz): 2.5 (cat), 1.1 (guinea pig), 0.48 (alligator-lizard tectorial fibers), 0.42 (tree frog), and 0.34 (alligator-lizard free-standing fibers). Some of this variation in corner frequency can be explained by temperature-dependent lowpass-filter mechanisms with a temperature factor of 2.6-3.3 for a change in temperature of 10 degrees C. However, factors in addition to temperature must be involved in producing the differences in corner frequency between cat and guinea pig fibers and between tectorial and free-standing fibers in the alligator lizard.     

正常豚鼠听性脑干反应与听性稳态反应的对比

目的 比较正常豚鼠听性脑干反应(ABR)和听性稳态反应(ASSR)阈值的差异,为利用豚鼠进行听力学研究提供理论依据.方法 选正常听力豚鼠12只(24耳),在戊巴比妥钠镇静状态下,分别行ABR和ASSR测试.ABR为click刺激声,刺激率为11.1次/s,记录ABR的Ⅱ渡反应阈值.ASSR载波频率(CF)为0.5、1、2、3、4、6 kHz,调制频率(MF)为154 Hz,记录各载频的反应阅值.结果 正常豚鼠ASSR反应阈值高于ABR反应阈值,CF:0.5.4 kHz时.ABR与ASSR阈值间有统计学差异(P<0.01);CF=6 kHz时,两阈值间无统计学差异(P>0.05).结论 正常豚鼠ABR与ASSR阈值间存在较大差值,但ABR与6 kHz的ASSR阈值间无显著差异.故对豚鼠进行听阈评估时,要注意两者间由于ASSR载波频率不同所引起的差异.     

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

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

Effects of neonatal partial deafness and chronic intracochlear electrical stimulation on auditory and electrical response characteristics in primary auditory cortex

The use of cochlear implants in patients with severe hearing losses but residual low-frequency hearing raises questions concerning the effects of chronic intracochlear electrical stimulation (ICES) on cortical responses to auditory and electrical stimuli. We investigated these questions by studying responses to tonal and electrical stimuli in primary auditory cortex (AI) of two groups of neonatally deafened cats with residual high-threshold, low-frequency hearing. One group were implanted with a multi-channel intracochlear electrode at 8 weeks of age, and received chronic ICES for up to 9 months before cortical recording. Cats in the other group were implanted immediately prior to cortical recording as adults. In all cats in both groups, multi-neuron responses throughout the rostro-caudal extent of AI had low characteristic frequencies (CFs), in the frequency range of the residual hearing, and high-thresholds. Threshold and minimum latency at CF did not differ between the groups, but in the chronic ICES animals there was a higher proportion of electrically but not acoustically excited recording sites. Electrical response thresholds were higher and latencies shorter in the chronically stimulated animals. Thus, chronic implantation and ICES affected the extent of AI that could be activated by acoustic stimuli and resulted in changes in electrical response characteristics.     

Temporal relationship between the auditory brainstem response and focal responses of auditory nerve root and cochlear nucleus during development in the tammar wallaby (Macropus eugenii)

Thirty-two pouch-young tammar wallabies were used to discover the generators of the auditory brainstem response (ABR) during development by the use of simultaneous ABR and focal brainstem recordings. A click response from the auditory nerve root (ANR) in the wallaby was recorded from postnatal day (PND) 101, when no central auditory station was functional, and coincided with the ABR, a simple positive wave. The response of the cochlear nucleus (CN) was detected from PND 110, when the ABR had developed 1 positive and 1 negative peak. The dominant component of the focal ANR response, the N1 wave, coincided with the first half of the ABR P wave, and that of the focal CN response, the N1 wave, coincided with the later two thirds. In older animals, the ANR response coincided with the ABR's N1 wave, while the CN response coincided with the ABR's P2, N2 and P3 waves, with its contribution to the ABR P2 dominant. The protracted development of the marsupial auditory system which facilitated these correlations makes the tammar wallaby a particularly suitable model.