Place and Time Coding of Frequency in the Peripheral Auditory System: Some Physiological Pros and Cons

Evidence for and against classical theories of ‘place’ and ‘period’ mechanisms for the coding of frequency, and the modifications of the theories invoked to account for the pitch of ‘residue’ and other types of stimuli, are examined in the light of physiological data. These include new data on the temporal discharge patterns of cochlear nerve fibres under stimulation with two-tone complexes, harmonic and inharmonic three-tone complexes, and five-tone complexes of differing relative phase. They show, in particular, that certain arguments against ‘period’ coding of ‘residue’ pitch are invalid. The interspike intervals in the discharge patterns of cochlear fibres under these conditions are consistent with the pitches heard. On the other hand, the classical ‘period’ theory needs to be modified to take into account the normally relatively sharp frequency selectivity of cochlear fibres, and requires certain inefficiencies on the part of the central processor for pitch.

Comparison of measures of cochlear fibre frequency selectivity with analogous psychophysical data in man, including those on the ‘existence region’ of ‘residue’ pitch, suggests that ‘residue’-type stimuli judged to be tonal in quality could both: (a) be sufficiently resolved spectrally at the cochlear fibre level to serve as input to any of the current spectral ‘pattern recognition’ mechanisms proposed for the pitch extraction of complex signals, and also, (b) could generate patterns of temporal discharge reflecting enough waveform interaction between the harmonics to convey the pitch heard, because of the shape of the cochlear filters. (This conclusion might have to be qualified in the light of further physiological experiments on the ‘second effect’ of pitch shift.)

The present evidence, both psychophysical and physiological, suggests the following synthesis: musical interval recognition and relatively crude frequency discrimination can be accomplished by trained observers on signals where the frequency appears to be coded exclusively in terms of temporal information. However, the pitch quality of these signals is judged to be poor or absent. Likewise, signals, apparently coded exclusively by ‘place’ mechanisms, while having tonality, allow relatively crude frequency discrimination and judgment of musical intervals. With the possible exception of psychophysical data on the phenomenon of diplacusis, the present evidence cannot exclude the possibility that the central pitch extractor mechanism utilizes both the ‘place’ and ‘period’ cues produced by pure-tone signals (below 5 kHz) and ‘residue’-type signals, both signals evoking strong pitch and fine acuity of frequency discrimination. The degree of salience of a signal's pitch could well depend on the coherence of the two types of cue.

However, the greatest obstacle to the acceptance of ‘place’ coding mechanisms for frequency, particularly of the frequency components of a complex sound, is the restricted dynamic range of the peripheral elements of the auditory nervous system. Because of this, it is not clear how differences in the spectral energy distribution in signals at medium to high sound levels can be established in terms of patterns of mean discharge rate across the cochlear fibre array. At high sound levels, physiological evidence suggests that the discharge rates of the majority of stimulated fibres will be saturated, whereas psychophysical evidence suggests that the coding of the frequency and the relative level of even single-component signals can be carried out over a wide dynamic range in the absence of cues derived from spread of activity across the fibre array. Some new data, however, indicate that this problem may be circumvented at the cochlear nucleus level, but the coding mechanisms involved at the primary neurone level are obscure. One intriguing possibility exists that the auditory nervous system may utilize the fine temporal structure of cochlear fibre discharge patterns for the transmission of ‘place’ information.     

Neural coding of repetitive clicks in the medial geniculate body of cat

The activity of 418 medial geniculate body (MGB) units was studied in response to repetitive acoustic pulses in 35 nitrous oxide anaesthetized cats. The proportion of MGB neurons insensitive to repetitive clicks was close to 30%. On the basis of their pattern of discharge, the responsive units were divided into three categories. The majority of them (71%), classified as ‘lockers’, showed discharges precisely time-locked to the individual clicks of the train. A few units (8%), called ‘groupers’, had discharges loosely synchronized to low-rate repetitive clicks. When the spikes were not synchronized, the cell had transient or sustained responses for a limited frequency range and was classified as a ‘special responder’ (21%). Responses of ‘lockers’ were time-locked up to a limiting rate, which varied between 10 and 800 Hz; half of the ‘lockers’ had a limiting rate of locking equal to or higher than 100 Hz. The degree of entrainment, defined as the probability that each click evokes at least one spike, regularly decreases for increasing rates; on the other hand, the precision of locking increases with frequency. The time jitter observed at 100 Hz might be as small as 0.2 ms and was 1.2 ms on average. The population of ‘lockers’ can mark with precision the transients of complex sounds and has response properties still compatible with a temporal coding of the fundamental frequency of most animal vocalizations.     

Vestibular signal processing by separate sets of neuronal filters

Second-order vestibular neurons (2°VN) are the central element for the transformation of body motion-related sensory signals into extraocular motor commands for retinal image stabilization during locomotion. The wide range of motion dynamics necessitates sensory signal transformation in parallel, frequency-tuned channels. Accordingly, in various vertebrates, 2°VN have been shown to form differently tuned functional subgroups. In frog, these neurons subdivide into two separate populations with distinctly different intrinsic membrane properties, discharge dynamics and synaptic response characteristics. Frog tonic 2°VN exhibit low-pass filter characteristics and membrane properties that cause amplification of synaptic inputs, whereas phasic 2°VN form band-pass filters that allow frequency-dependent shunting of repetitive inputs. The differential, yet complementary membrane properties render tonic 2°VN particularly suitable for synaptic integration and phasic 2°VN for differentiation and event detection. Differential insertion of the two cell types into local circuits reinforces the functional consequences of the intrinsic membrane properties, respectively. As a consequence, the synergy of cellular and network properties creates sets of neuronal elements with particular filter characteristics that form flexible, frequency-tuned components for optimal transformation of all dynamic aspects of body motion-related multisensory signals.     

Single motor unit activity of human intrinsic laryngeal muscles during respiration.

Individual motor units in the thyroarytenoid (TA) and cricothyroid (CT) muscles were studied in 10 normal human volunteers during quiet respiration. Both tonic and phasic firing patterns were found in both TA and CT units. The rate of firing was higher during inhalation than during exhalation in phasic TA units and in tonic CT units. Tonically active units had a higher firing frequency than phasically active units in both TA and CT muscles. Phasically active units corresponded with the respiratory cycle, with firing associated with inhalation in both the TA and CT muscles. A variety of firing patterns were found between units in both the TA and CT muscles, and in one subject, units recorded from the same muscle had very different firing patterns. The results suggest that although laryngeal motoneurons are modulated by the respiratory cycle, they do not respond uniformly to respiration.     

A model of anuran auditory periphery reveals frequency-dependent adaptation to be a contributing mechanism for two-tone suppression and amplitude modulation coding

Anuran auditory nerve fibers (ANF) tuned to low frequencies display unusual frequency-dependent adaptation which results in a more phasic response to signals above best frequency (BF) and a more tonic response to signals below. A network model of the first two layers of the anuran auditory system was used to test the contribution of this dynamic peripheral adaptation on two-tone suppression and amplitude modulation (AM) tuning. The model included a peripheral sandwich component, leaky-integrate-and-fire cells and adaptation was implemented by means of a non-linear increase in threshold weighted by the signal frequency. The results of simulations showed that frequency-dependent adaptation was both necessary and sufficient to produce high-frequency-side two-tone suppression for the ANF and cells of the dorsal medullary nucleus (DMN). It seems likely that both suppression and this dynamic adaptation share a common mechanism. The response of ANFs to AM signals was influenced by adaptation and carrier frequency. Vector strength synchronization to an AM signal improved with increased adaptation. The spike rate response to a carrier at BF was the expected flat function with AM rate. However, for non-BF carrier frequencies the response showed a weak band-pass pattern due to the influence of signal sidebands and adaptation. The DMN received inputs from three ANFs and when the frequency tuning of inputs was near the carrier, then the rate response was a low-pass or all-pass shape. When most of the inputs were biased above or below the carrier, then band-pass responses were observed. Frequency-dependent adaptation enhanced the band-pass tuning for AM rate, particularly when the response of the inputs was predominantly phasic for a given carrier. Different combinations of inputs can therefore bias a DMN cell to be especially well suited to detect specific ranges of AM rates for a particular carrier frequency. Such selection of inputs would clearly be advantageous to the frog in recognizing distinct spectral and temporal parameters in communication calls.     

Superposition models of the discharge patterns of units in the lower auditory system

Superposition of point processes has often been suggested as an abstract model for the generation of neural discharge patterns due to its simplicity (input spike trains are simply merged and then retransmitted by the model neuron). The properties of the superposition of two renewal processes are examined in relation to the properties of its components. A satisfactory condition is found so that the superposition of two renewal processes possesses negative serial dependence of intervent intervals. However, the imposition of a dead time of approximately the same duration as that of the components on their superposition process is shown to remove much of this dependence. Thus, the adequacy of the simple superposition of renewal processes as a model for discharge patterns from regions that are typified by negative serial dependence (such as the lateral superior olive) may be limited, but may suffice for the discharges of primary-like units.     

Coding of amplitude-modulated tones in the central auditory system of catfish

In the catfish central acoustic system information is coded by two subsets of units. Type I units show little or no adaptation, type II units adapt rapidly, and some units are transitional, showing moderate adaptation. The two groups of units also respond differently when exposed to sinusoidal amplitude modulation of the signal's carrier frequency. The tonic, less readily adapting type I units code over an intensity range of about 30 dB, are fairly insensitive to intensity changes, and follow stimulus envelopes of 60 Hz and less. They apparently discharge in response to the actual intensity of the signal rather than in response to something in its temporal pattern. The onset-sensitive, fast-adapting type II units on the other hand are restricted to an intensity range of only 10 dB, show greater sensitivity to intensity changes, and are capable of following the temporal pattern of amplitude-modulated stimuli exceeding 100 Hz. These units appear to code the temporal changes in the stimulus intensity irrespective of the absolute intensity of the signal.     

Intensity coding in the auditory periphery of the cat: Responses of cochlear nerve and cochlear nucleus neurons to signals in the presence of bandstop masking noise

The dynamic range over which fine intensity discrimination is possible has been reported to be largely unaffected by limitation of the spread of neuronal activity to neighbouring frequency regions by bandstop noise masking.We have therefore examined the responses of cochlear nerve and nucleus neurons to tone and noise signals in the presence of a bandstop masking noise designed to be comparable to that employed in the psychophysical experiments. Under these conditions, the vast majority of cochlear nerve fibres were saturated by sound levels at which some 50% of our sample of cochlear nucleus neurons still responded to signal level differences. The extended dynamic ranges of these cochlear nucleus neurons was shown to be a result of activation, by the masking noise, of the lateral inhibitory side-bands ‘biassing’ the neuron's discharge. A small proportion of cochlear fibres, having low spontaneous discharge rates and showing strong two-tone suppression effects, demonstrated analogous but not so pronounced effects. It is unclear in what form information on the level of stimuli under these conditions is transmitted by the majority of apparently saturated cochlear nerve fibres, but several possible mechanisms are discussed.     

Relations between frequency selectivity and two-tone rate suppression in lizard cochlear-nerve fibers

Cochlear-nerve fibers innervating the apicial region of the alligator lizard basilar papilla show sharp frequency selectivity in response to single tones (measured with the frequency threshold contour, or FTC), and the phenomenon of two-tone rate suppression (TTRS) in response to two simultaneously presented tones (measured with the iso-TTRS contour, or ITC). The gross shapes of the FTCs, as characterized by the slopes of the sides and Q_10dB, vary systematically with the fiber's characteristic frequency (CF). ‘Fine-structural’ features are also found: below CF, notches (frequency regions of relatively high threshold) occur in the FTC at frequencies related to CF. Above CF, a break frequency, which varies with CF, divides the FTC into segments of different slope. Features of the ITC also vary with CF. The detailed shapes of the FTCs and ITCs are related: lobes of the ITC interdigitate with notches in the FTC; the side of the FTC with steepest slope is closely associated with the side of the ITC with steepest slope. The close relation that is observed between sharp frequency selectivity and TTRS suggests that both phenomena arise from a common cochlear mechanism.     

Cortical Processing Related to Intensity of a Modulated Noise Stimulus—a Functional Near-Infrared Study

Sound intensity is a key feature of auditory signals. A profound understanding of cortical processing of this feature is therefore highly desirable. This study investigates whether cortical functional near-infrared spectroscopy (fNIRS) signals reflect sound intensity changes and where on the brain cortex maximal intensity-dependent activations are located. The fNIRS technique is particularly suitable for this kind of hearing study, as it runs silently. Twenty-three normal hearing subjects were included and actively participated in a counterbalanced block design task. Four intensity levels of a modulated noise stimulus with long-term spectrum and modulation characteristics similar to speech were applied, evenly spaced from 15 to 90 dB SPL. Signals from auditory processing cortical fields were derived from a montage of 16 optodes on each side of the head. Results showed that fNIRS responses originating from auditory processing areas are highly dependent on sound intensity level: higher stimulation levels led to higher concentration changes. Caudal and rostral channels showed different waveform morphologies, reflecting specific cortical signal processing of the stimulus. Channels overlying the supramarginal and caudal superior temporal gyrus evoked a phasic response, whereas channels over Broca’s area showed a broad tonic pattern. This data set can serve as a foundation for future auditory fNIRS research to develop the technique as a hearing assessment tool in the normal hearing and hearing-impaired populations.     

Stimulus properties influencing the responses of inferior colliculus neurons to amplitude-modulated sounds

The temporal pattern of the responses of neurons in the inferior colliculus of the anesthetized rat were studied using continuous tone or noise carrier signals, amplitude modulated by pseudorandom noise. Period histograms of the responses, cross-correlated with the pseudorandom noise, gave an estimate of the unit's impulse responses to modulation. The amplitude-modulation rate transfer function (MTF) was obtained by Fourier transforming the correlograms. At sound levels within approximately 15 dB of the unit threshold, the MTFs were near lowpass functions between 6 and 200 Hz but became more bandpass-like as the intensity was increased. There was a steep decline in the response to modulation at modulation frequencies above 200 Hz for all stimulus intensities. For the bandpass-type MTFs the greatest modulation of the discharge pattern occurred at modulation frequencies between 10 and 200 Hz with a maximum in the distribution of MTF peak values between 100 and 120 Hz. There was no consistent relationship with characteristic frequency of either the position of the MTF peak or the high-frequency cutoff of the MTF. The cross-correlograms obtained at high stimulus intensities (30-60 dB above threshold) often showed a negative peak, representing a decrease in the probability of firing in response to intensity increments in the stimulus, and denoting a nonmonotonic rate-intensity function. The MTFs for units responding to amplitude-modulated broadband noise were often flatter in the low frequency region than those generated with tone carriers at corresponding intensities. For some units addition of a broadband noise background to the modulated tone changed the response characteristic of the MTF from bandpass to lowpass and shifted the MTF peak to a lower modulation frequency. The results demonstrate that although neurons in the inferior colliculus are selectively sensitive to the modulation frequency of dynamic stimuli, the response characteristics are not invariant, but instead are closely dependent on the conditions under which the modulation is presented.     

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.     

Relation of binaural interaction and spectro-temporal characteristics in the auditory midbrain of the grassfrog

The relation between binaural interaction type and spectro-temporal characteristics was studied for single units in the auditory midbrain of the grassfrog. Tonal and continuous wideband noise ensembles have been used as stimuli. Spectro-temporal sensitivities were determined for ipsi-, contra- and bilateral stimulus presentation by a closed sound system. Binaural interaction was classified in monaural EO (one ear excitatory), binaural EE (both ears excitatory) and EI (one ear excitatory, the other inhibitory) and purely inhibitory categories. Binaural interaction appeared to be rather invariant to alterations in stimulus intensity and type. A very clear correlation was observed between best frequency and binaural interaction type: EE units are predominantly of high best frequency, whereas EI units are predominantly of low best frequency. The correlation with latency was less significant: EE units tended to have somewhat shorter latencies that EI units. EO units take an intermediate position. Comparisons of ipsi-, contra- and bilateral spectro-temporal sensitivities, revealed differences in best frequency, latency and temporal discharge pattern. In some units a complex interplay of excitatory and inhibitory monaural influences was demonstrated. A number of units was recorded, which were characterized by multiple activation or suppression areas. The majority of these units exhibited frequency-dependent binaural interaction types. In some units it was noticed that binaural interaction type can be dependent on state of adaptation. A comparison of binaural interaction types of neighbouring units provided only weak evidence for a binaural organization in the anuran auditory midbrain, since simultaneously recorded pairs shared the same binaural interaction type only slightly more than expected by mere chance (chi 2-test, P less than 0.10).     

Intensity functions of single unit responses to tone in the medial geniculate body of cat

Extracellular spike activity was recorded from single units in the medial geniculate body (MGB) of nitrous oxide anaesthetized cats. The responses of 291 units to tone bursts at the characteristic frequency (CF) were studied as a function of stimulus intensity, covering a range from 10 to 100 dB SPL. The proportion of MGB units characterized by a monotonie or a non-monotonic discharge rate-intensity function was 26% and 74%, respectively. In addition, changes of response latency as a function of tone levels were demonstrated to be either monotonie (38% of units) or non-monotonic (62% of units). One third of MGB units showed a change of response pattern with increasing intensities, in similar proportion towards either prevailing excitatory or inhibitory components. The monotonie units tended to differ from non-monotonic ones in addition to their intensity function by showing shorter response latencies, a higher response probability to broad-band stimuli and simpler response patterns. The mean dynamic range of the monotonie unit population was 60 dB, with thresholds ranging from 10 to 90 dB SPL; most discharge rate-intensity functions did not saturate at sound levels of 100 dB SPL. In the population of non-monotonic units, the ‘best’ intensity, defined as the intensity giving the strongest response, ranged between 10 and 100 dB SPL. The present results suggest that the intensity could be signaled by the mean firing rate of a restricted population of monotonie units or place coded by the distribution of maximally activated non-monotonic units which are broadly tuned to different intensities.     

Temporal suppression and augmentation of click-evoked otoacoustic emissions

This study investigates temporal suppression of click-evoked otoacoustic emissions (CEOAEs), occurring when a suppressor-click is presented close in time to a test-click (e.g. 0-8ms). Various temporal suppression methods for examining temporal changes in cochlear compression were evaluated and measured here for seven subjects, both for short- and long-latency CEOAEs. Long-latency CEOAEs (duration >20ms) typically indicate the presence of synchronised spontaneous otoacoustic emissions (SSOAEs). Temporal suppression can only be linked to changes in CEOAE-compression if the suppressor-click affects the CEOAE magnitude. Phase changes induced by the suppressor-click were shown to bias suppression in two ways: (i) when a specific asymmetric measurement method was used and (ii) when synchronisation between the CEOAE and the click-stimuli was incomplete. When such biases were eliminated, temporal suppression and augmentation (the opposite effect) were observed and shown to be subject-dependent. This indicates that the nonlinearity underlying temporal suppression can work in a more (i.e., suppressed) or less (i.e., augmented) compressive state, depending on the inter-click interval and the subject under test. Temporal suppression was shown to be comparable for CEOAEs and SSOAEs, indicating similar underlying cochlear nonlinear mechanisms. This study contributes to a better understanding of the temporal properties of cochlear dynamics.     

Scalp thickness in the temporal region: its relevance to the development of cochlear implants

Abstract

The thickness of the scalp in the temporal region was measured at two representative points: point 1, over the most laterally prominent part of the mastoid process: and point 2, at the horizontal level of the upper border of the orbit, vertically in line with point 1. Measurements were made directly at autopsy in four patients, then in 50 live patients by magnetic resonance imaging (MRI). The relevance of the methodology and of these measurements to cochlear implant development is discussed. As the term ‘scalp’ usually includes the periosteum of the skull, the authors have used the term ‘scalp’ to include the range from the surface of the scalp to the bone shadow as seen on MRI, and the term ‘skin’ where, at autopsy, the skin is lifted leaving the periosteum intact.     

Tonic cervical stimulation: does it influence eye position and eye movements in man?

In healthy subjects eye movements were analysed during body rotation, during trunk torsion either with the head passively held stationary in space or with the head voluntarily stabilized in space, and during voluntary head movements. Trapezoidal movements around the vertical axis were performed (+/- 40 degrees, plateau 10 s, duration of ramp 1 or 4 s). Moreover the influence of a tonic head deviation up to 40 degrees on optokinetic nystagmus and on vestibulo-ocular reflex during sinusoidal turning was examined. Eye movements were recorded by DC-electrooculography. Saccadic and slow components of eye movements and the shift of eye position during the plateau of the trapezoidal stimulus were analysed. For all modes of stimulation during the plateau no nystagmus occurred. At the end of the dynamic phase of the stimulus relatively frequent eye deviations--mostly in the direction of the head deviation--were observed, not only after turning the trunk with the head stabilized in space (cervical stimulation) but also after turning head and trunk together. The fact that such eye deviations are thus observed even in the absence of any tonic, especially cervical stimulus, supports the assumption that they cannot be attributed to a tonic stimulus but merely to an effect of the preceding phasic stimulus which outlasts them. Also amplitude and direction of eye shifts during the plateau do not depend on a tonic stimulus, but merely on the eye deviation reached at the end of the dynamic phase of stimulation. Optokinetic nystagmus and vestibulo-ocular reflex are not influenced by an additional tonic cervical stimulus.     

Paired-Tone Stimuli Reveal Reductions and Alterations in Temporal Processing in Inferior Colliculus Neurons of Aged Animals

Temporal processing deficits in the central auditory system of aged subjects are apparent from animal and human studies but could be due to peripheral hearing loss. Sequential paired tone stimuli reveal age-related changes in temporal processing properties of neurons in the central nucleus of the inferior colliculus (CIC). A greater proportion of CIC neurons exhibit suppression of excitability following pure tone stimulation in 20 month old ("aged") compared with 3–6 month "young" Long Evans rats. The duration (time constant of exponential curve fit to recovery of excitability) of suppression is also increased in aged compared with young rats, with more neurons exhibiting suppression with time constants over 100 ms. The time course of post-stimulatory suppression is not dependent on the duration or intensity of preceding stimuli and is not correlated with either initial magnitude of suppression or best frequency of IC neurons. Although the increase in unit thresholds is greater for high-frequency units in old animals, the largest post-stimulatory suppression changes occur in neurons with best frequencies of less than 10 kHz. Since the increase in duration of post-stimulatory suppression is not correlated with peripheral hearing loss, the difference is likely attributed to central auditory neuron changes in aging. In addition, the proportion of IC neurons exhibiting other temporal patterns of excitability (post-stimulatory facilitation and delayed-maximum excitability) is reduced in aged animals. Therefore, temporal processing of acoustic information is significantly altered in aged animals. The greater post-stimulatory suppression of excitability, reduced facilitation, and delayed facilitation is expected to reduce and alter the encoded information passing from the brainstem through the IC to higher structures. These changes correlate with reduced speech understanding in noise, elevated thresholds in noisy conditions, and reduced temporal processing capabilities in the elderly.     

Developing an assessment approach for perceptual changes to tinnitus sound characteristics for adult cochlear implant recipients

Objective: To investigate the impact of cochlear implantation on tinnitus suppression, characteristics, localization, and duration. Design: A cochlear implant (CI) recipient-focused postal questionnaire survey. Study sample: The questionnaire was posted, with consent, to 100 adults who had received a unilateral CI at the RNTNEH between 1988 and 1999. All adults spoke English as their first language and were postlingually deafened. Sixty-eight adults (38 female, 29 male, one unspecified) aged 31–80 years (mean 61 years) completed and returned the questionnaire without interview. Results: With the processor ‘ON’, CI recipients experienced total or partial suppression of tinnitus ipsilateral to their CI in 57% of cases, and in 43% where tinnitus was perceived contralateral to the CI. The percentage of CI recipients who experienced high tone tinnitus was reduced from 60% pre-implant to 29% post-implant with the processor ‘ON’ while pulsatile tinnitus was reduced from 38% pre-implant to 13% post-implant. CIs were also found to reduce the tonal complexity and duration, and change the source localization of tinnitus post-implantation. Conclusions: Perceptual changes to tinnitus can take place post-implantation. Changes can occur within the four categories explored: tinnitus suppression, characteristics, localization, and duration of awareness per day.     

Temporal and spatial coding of periodicity information in the inferior colliculus of awake chinchilla (Chinchilla laniger)

Amplitude modulation responses and onset latencies of multi-unit recordings and evoked potentials were investigated in the central nucleus of inferior colliculus (ICC) in the awake chinchilla. Nine hundred and one recording sites with best frequencies between 60 and 30 kHz showed either phasic (18%), tonic (25%), or phasic-tonic (57%) responses. Of 554 sites tested for responses to modulation frequencies 73% were responsive and 57% showed clear preference for a narrow range of modulation frequencies. Well defined bandpass characteristics were found for 32% of rate modulation transfer functions (rate-MTFs) and 36% of synchronization MTFs (sync-MTFs). The highest best modulation frequency (BMF) of a bandpass rate-MTF was 600 Hz. Neurons with phasic responses to best-frequency tones showed strong phase coupling to modulation frequencies and were dominated by bandpass rate-MTFs and sync-MTFs. Most neurons with tonic responses showed bandpass tuning only for sync-MTFs. Both BMFs and onset latencies changed systematically across frequency-band laminae of the ICC. Low BMFs and long latencies were located medially and high BMFs and short latencies laterally. Latency distributions obtained with evoked potentials to clicks showed a similar gradient to the multi-unit data. These findings are in line with previous findings in different animals including humans and support the hypothesis that temporal processing results in a topographic arrangement orthogonal to the spectral processing axis, thus forming a second neural axis of the auditory system.