Temporal summation of pulsate brain stimulation in normal and deafened cats.

Behaviorally measured, electrical-stimulation thresholds were obtained from 11 electrodes permanently positioned in the auditory system and other brain loci. Number of pulses and interpulse intervals were varied to determine how detection thresholds were affected by stimulation parameters. Detection thresholds generally decreased with increased number of pulses and with shorter interpulse intervals. A method is presented to describe the parametric threshold data for each electrode in terms of three constants: a single-pulse threshold which characterizes the sensitivity of the placement; a time constant of temporal summation; and a compression factor which describes the range of threshold variation. For three placements in the vicinity of cochlear nucleus, bilateral cochlear destruction permanently altered parametric thresholds. In particular, single-pulse threshold was lowered by 9.2 dB; time constant of temporal summation was reduced by a factor of 100; and the compression factor was increased. Classic strength-duration time constants were determined using behavioral methods and were shown to be equal in magnitude to the greatly reduced time constants for temporal summation in the deafened animals. This implies that capacity for temporal integration may be substantially reduced or lost in at least the lower level of the auditory system following deafness.     

Perception of amplitude envelope variations of pulsatile electrotactile stimuli

Gross variations of the speech amplitude envelope, such as the duration of different segments and the gaps between them, carry information about prosody and some segmental features of vowels and consonants. The amplitude envelope is one parameter encoded by the Tickle Talker, an electrotactile speech processor for the hearing impaired which stimulates the digital nerve bundles with a pulsatile electric current. Psychophysical experiments measuring the duration discrimination and identification, gap detection, and integration times for pulsatile electrical stimulation are described and compared with similar auditory measures for normal and impaired hearing and electrical stimulation via a cochlear implant. The tactile duration limen of 15% for a 300-ms standard was similar to auditory measures. Tactile gap detection thresholds of 9 to 20 ms were larger than for normal-hearing but shorter than for some hearing-impaired listeners and cochlear implant users. The electrotactile integration time of about 250 ms was shorter than previously measured tactile values but longer than auditory integration times. The results indicate that the gross amplitude envelope variations should be conveyed well by the Tickle Talker. Short bursts of low amplitude are the features most likely to be poorly perceived.     

Place-pitch discrimination of single- versus dual-electrode stimuli by cochlear implant users (L)

Simultaneous or near-simultaneous activation of adjacent cochlear implant electrodes can produce pitch percepts intermediate to those produced by each electrode separately, thereby increasing the number of place-pitch steps available to cochlear implant listeners. To estimate how many distinct pitches could be generated with simultaneous dual-electrode stimulation, the present study measured place-pitch discrimination thresholds for single- versus dual-electrode stimuli in users of the Clarion CII device. Discrimination thresholds were expressed as the proportion of current directed to the secondary electrode of the dual-electrode pair. For 16 of 17 electrode pairs tested in six subjects, thresholds ranged from 0.11 to 0.64, suggesting that dual-electrode stimuli can produce 2-9 discriminable pitches between the pitches of single electrodes. Some subjects demonstrated a level effect, with better place-pitch discrimination at higher stimulus levels. Equal loudness was achieved with dual-electrode stimuli at net current levels that were similar to or slightly higher than those for single-electrode stimuli.     

Effects of modulation wave shape on modulation frequency discrimination with electrical hearing

Amplitude modulations of pulsitile stimulation can be used to convey pitch information to cochlear implant users. One variable in designing cochlear implant speech processors is the choice of modulation waveform used to convey pitch information. Modulation frequency discrimination thresholds were measured for 100 Hz modulations with four waveforms (sine, sawtooth, a sharpened sawtooth, and square). Just-noticeable differences (JNDs) were similar for all but the square waveform, which often produced larger JNDs. The results suggest that a sine, sawtooth, and sharpened sawtooth waveforms are likely to provide similar pitch discrimination within a speech processing strategy.     

Detection versus discrimination of brief tones by cats with auditory cortex lesions.

In recent years, a number of investigators have provided evidence that the auditory cortex has a critical role in both the detection and discrimination of brief sounds. Dogs and humans with lesions of the neocortical auditory centers have been reported to exhibit significantly elevated detection thresholds for signals shorter than 16 ms in duration. In tests of frequency discrimination, the same subjects also exhibited severe deficits whenever tonal signals were less than 20--40 mn in lengths. In the present report, we present evidence brief tones. Operated cats, while exhibiting normal difference limens for 1-kHz tones of 100-ms duration, have significantly elevated limens for discriminating tones of 8- and 2-ms duration. With further testing, the same operated cats can be shown to have normal absolute thresholds for detecting brief tones.     

Differential electrical excitation of the auditory nerve

The multichannel cochlear prosthesis requires an electrode stimulus configuration which produces a stimulus field spatially localized to each electrode. In this paper, a three-dimensional discrete resistance model of the cochlea was developed which exhibits electrical response properties similar to those observed during electrical stimulation of the cochlea. The model results suggest that the spatial attenuation of current within the cochlea varies greatly in magnitude, depending on the stimulus configuration. In addition, the model suggests that the spatial attenuation of current in both the auditory nerve fiber endings in the organ of Corti and in the myelinated fibers within the cochlear ground paths is different from the voltage attenuation in the scalar fluids. Therefore the efficacy with which a particular stimulus configuration differentially excites local terminal auditory nerve fiber populations cannot be deduced from scalar voltage measurements which have previously been recorded in the literature. Consequently physiological experiments were performed in the cat to measure the current distributions in the terminal nerve fiber region for monopolar and bipolar stimulation of the scala tympani, and also for stimulation between the scala tympani and the scala vestibuli. The mean length constants measured in the basal turn for these stimuli were found to be 12, 3, and 7.5 mm, respectively.     

Binaural unmasking with multiple adjacent masking electrodes in bilateral cochlear implant users

Bilateral cochlear implant (BiCI) users gain an advantage in noisy situations from a second implant, but their bilateral performance falls short of normal hearing listeners. Channel interactions due to overlapping electrical fields between electrodes can impair speech perception, but its role in limiting binaural hearing performance has not been well characterized. To address the issue, binaural masking level differences (BMLD) for a 125 Hz tone in narrowband noise were measured using a pair of pitch-matched electrodes while simultaneously presenting the same masking noise to adjacent electrodes, representing a more realistic stimulation condition compared to prior studies that used only a single electrode pair. For five subjects, BMLDs averaged 8.9 ± 1.0 dB (mean ± s.e.) in single electrode pairs but dropped to 2.1 ± 0.4 dB when presenting noise on adjacent masking electrodes, demonstrating a negative impact of the additional maskers. Removing the masking noise from only the pitch-matched electrode pair not only lowered thresholds but also resulted in smaller BMLDs. The degree of channel interaction estimated from auditory nerve evoked potentials in three subjects was significantly and negatively correlated with BMLD. The data suggest that if the amount of channel interactions can be reduced, BiCI users may experience some performance improvements related to binaural hearing.     

Psychophysical correlates of contralateral efferent suppression. I. The role of the medial olivocochlear system in "central masking" in nonhuman primates

An extensive physiological literature, including experimental and clinical studies in humans, demonstrates that activation of the medial olivocochlear (MOC) efferent system, by either contralateral sound or electrical stimulation, can produce significant alterations in cochlear function and suggests a role for the MOC system in influencing the auditory behavior of binaural hearing. The present data are from psychophysical studies in nonhuman primates which seek to determine if the noted physiological changes in response to contralateral acoustic stimulation have a perceptual counterpart. Four juvenile Japanese macaques were trained to respond to the presence of 1-s sinusoids, presented to the test ear, in an operant reinforcement paradigm. Thresholds were compared for frequencies ranging from 1.0 to 4.0 kHz in quiet, with thresholds measured when continuous, two octave-band noise, centered on the test tone frequency, was presented in the contralateral ear. Contralateral noise was presented at levels of 10-60 dB above detection threshold for the test-tone frequency. While some variability was evident across subjects, both in the frequency distribution and magnitude (as a function of contralateral noise level), all subjects exhibited an increase, or suppression of thresholds in the presence of contralateral noise. On average, thresholds increased systematically with contralateral noise level, to a peak of 7 dB. In one subject, the threshold increase seen with contralateral noise was significantly reduced when the MOC was surgically sectioned on the floor of the IVth ventricle. The characteristics of the measured shifts in behavioral thresholds, in the presence of contralateral noise reported here, are qualitatively and quantitatively similar to both efferent physiological suppression effects and psychophysical central masking threshold shifts which have been reported previously. These data suggest that at least some aspects of "central masking" are efferent-mediated peripheral processes, and that the term "central masking" may be incorrect.     

An analysis of the effects of electrical field interaction with an acoustic model of cochlear implants

Electrical field interaction caused by current spread in a cochlear implant was modeled in an explicit way in an acoustic model (the SPREAD model) presented to six listeners with normal hearing. The typical processing of cochlear implants was modeled more closely than in traditional acoustic models by careful selection of parameters related to current spread or parameters that could amplify the electrical field interactions caused by current spread. These parameters were the insertion depth, electrode spacing, electrical dynamic range, and dynamic range compression function. The hypothesis was that current spread could account for the asymptote in performance in speech intelligibility experiments observed at around seven stimulation channels in a number of cochlear implant studies. Speech intelligibility for sentences, vowels, and consonants at three noise levels (SNR of +15 dB, +10 dB, and +5 dB) was measured as a function of the number of spectral channels (4, 7, and 16). The SPREAD model appears to explain the asymptote in speech intelligibility at seven channels for all noise levels for all speech material used in this study. It is shown that the compressive amplitude mapping used in cochlear implants can have a detrimental effect on the number of effective channels.     

Distortion product otoacoustic emission input/output functions in normal-hearing and hearing-impaired human ears.

DPOAE input/output (I/O) functions were measured at 7f2 frequencies (1 to 8 kHz; f2/f1 = 1.22) over a range of levels (-5 to 95 dB SPL) in normal-hearing and hearing-impaired human ears. L1-L2 was level dependent in order to produce the largest 2f1-f2 responses in normal ears. System distortion was determined by collecting DP data in six different acoustic cavities. These data were used to derive a multiple linear regression model to predict system distortion levels. The model was tested on cochlear-implant users and used to estimate system distortion in all other ears. At most but not all f2's, measurements in cochlear implant ears were consistent with model predictions. At all f2 frequencies, the ears with normal auditory thresholds produced I/O functions characterized by compressive nonlinear regions at moderate levels, with more rapid growth at low and high stimulus levels. As auditory threshold increased, DPOAE threshold increased, accompanied by DPOAE amplitude reductions, notably over the range of levels where normal ears showed compression. The slope of the I/O function was steeper in impaired ears. The data from normal-hearing ears resembled direct measurements of basilar membrane displacement in lower animals. Data from ears with hearing loss showed that the compressive region was affected by cochlear damage; however, responses at high levels of stimulation resembled those observed in normal ears.     

Characterizing auditory processing and perception in individual listeners with sensorineural hearing loss

This study considered consequences of sensorineural hearing loss in ten listeners. The characterization of individual hearing loss was based on psychoacoustic data addressing audiometric pure-tone sensitivity, cochlear compression, frequency selectivity, temporal resolution, and intensity discrimination. In the experiments it was found that listeners with comparable audiograms can show very different results in the supra-threshold measures. In an attempt to account for the observed individual data, a model of auditory signal processing and perception [Jepsen et al., J. Acoust. Soc. Am. 124, 422-438 (2008)] was used as a framework. The parameters of the cochlear processing stage of the model were adjusted to account for behaviorally estimated individual basilar-membrane input-output functions and the audiogram, from which the amounts of inner hair-cell and outer hair-cell losses were estimated as a function of frequency. All other model parameters were left unchanged. The predictions showed a reasonably good agreement with the measured individual data in the frequency selectivity and forward masking conditions while the variation of intensity discrimination thresholds across listeners was underestimated by the model. The model and the associated parameters for individual hearing-impaired listeners might be useful for investigating effects of individual hearing impairment in more complex conditions, such as speech intelligibility in noise.     

Relationship of neural and otoacoustic emission thresholds during endocochlear potential development in the gerbil

Distortion product otoacoustic emissions and auditory brainstem responses (ABRs) were measured in neonatal gerbils at three ages: at 15-16 days after birth (dab), near the onset of hearing when the endocochlear potential (EP) is known to be still immature; at 22 dab, when the EP first reaches mature levels; and at 30 dab. Comparing individual 15-16 dab animals to the 22 dab group, ABR threshold changes were typically larger than those for cubic distortion tone (CDT, 2f1-f2) emission thresholds which were, in turn, larger than those for the simple difference tone (DT, f2-f1). In contrast, from 22 to 30 dab there were no important changes in CDT or DT emission thresholds. Observed threshold-change relationships were very similar to those found in differential diagnosis investigations, where the EP was experimentally decreased using a chronic furosemide application. Therefore, most of the change in cochlear function over the two week period studied could be attributed to the maturation of EP during the first week. Model calculations further show that relative changes in CDT and DT emission thresholds are compatible with a movement of the operating point of the cochlear amplifier toward its symmetrical "central" point as the EP reaches mature levels.     

Detection of gaps in sinusoids and pulse trains by patients with cochlear implants

Gap detection thresholds were measured in patients with the Nucleus and Symbion cochlear implants as a function of several current waveform parameters. Detection of gaps in an electrical sinusoidal stimulus or in a train of biphasic pulses by implanted patients was similar to detection of gaps in comparable acoustic stimuli by normal listeners. Threshold gaps were 20-50 ms for low-level stimuli and improved with stimulus level to 2-5 ms for high-level stimuli. Gap detection performance was not affected by the electrode position in the cochlea or by the distance between stimulating electrodes. The data from most patients were well fitted by a trading relation between the duration of the gap and the square of stimulus intensity, indicating energy detection. The similarity of gap thresholds for normal subjects and implant patients suggests that many details of the peripheral neural activity are probably not important for this task, and that there is no retrocochlear loss of auditory temporal resolution with sensorineural hearing loss.     

Acoustic distortion in the ear canal. I. Cubic difference tones: effects of acute noise injury

Acoustic emissions in the form of cubic difference tones (CDT's), 2f1-f2, were measured in the ear canals of gerbils and cats. The state of the cochlea was manipulated by means of acute exposure to noise and was monitored with the aid of the whole-nerve response to tone pips. The resulting shifts in the levels of emissions generated by pairs of primary tones of equal intensity were then compared to the corresponding threshold shifts of the whole-nerve response across frequency. Data obtained from normal ears before injury indicate that the absolute thresholds of the whole-nerve responses across frequency are not necessarily good predictors of the absolute levels of CDT emissions generated by 70- and 80-dB SPL primaries. While high emission levels were often linked to low whole-nerve thresholds in pre-exposed ears, instances of animals with sensitive whole-nerve thresholds coupled with very weak emissions were also found. Conversely, animals with poor whole-nerve thresholds (shifted by up to 30 dB) could occasionally have high levels of emissions. After acute noise injury, however, the shifts of emission levels as a function of the center frequency of the primary-tone pair largely corresponded to the threshold shifts seen in the whole-nerve response. In other words, the temporary level shift of an acoustic emission largely reflected the acute change to a specific cochlear region associated with the primary frequencies.     

Short-term effects of sound exposure on the 2f1-f2 acoustic emission

The 2f1-f2 acoustic emission (AE) was recorded in the ear canals of cats following exposure to tone bursts of 200-ms duration. Exposures known to result in short-term adaptation (i.e., adaptation lasting under 1 s) at the level of the auditory nerve failed to produce significant post-exposure changes in AE amplitude. Given the apparent cochlear origin of the 2f1-f2 AE, this result is consistent with the view that short-term adaptation in the auditory periphery does not involve substantial changes in cochlear mechanics.     

Distortion product otoacoustic emissions in hearing-impaired mutant mice

The acoustic intermodulation distortion product (2f1-f2) was recorded in the ear canal of two different types of normally hearing mice and in four different types of hearing-impaired mutant mice. In the normally hearing animals, primary tones at levels of 60- to 100-dB SPL evoked distortion product emissions (DP's) at 20-50 dB below the primary levels. In the hearing-impaired mutants the level was dependent on the particular type of auditory dysfunction associated with the mutation. In both the deafness and the viable dominant spotting mutants, where either the whole organ of Corti or the stria vascularis is affected by the mutation, no DP's could be detected. The quivering mutant has a central auditory dysfunction associated with the nuclei of the superior olivary complex and the lateral lemniscus, with apparently normal cochlear function. DP's at levels and thresholds similar to those in normally hearing animals were recorded in quivering mice. The Bronx Waltzer mutant has a full complement of outer hair cells but only about of 20%-25% inner hair cells. DP's of small amplitude were recorded but the thresholds were raised by about 30 dB. The data suggest that the 2f1-f2 emission can be used as a noninvasive monitor of cochlear function.     

Frequency difference limens in normal and sensorineural hearing impaired chinchillas

This study assessed normal frequency discrimination ability in the chinchilla and determined how this ability changes as a function of an experimentally induced sensorineural hearing loss. Four chinchillas were trained by the methods of positive reinforcement to report absolute thresholds and frequency difference limens (FDLs). Subjects were then treated with the aminoglycosidic antibiotic amikacin until a 30-dB hearing loss was measured at 10.0 kHz. Absolute and frequency difference thresholds were determined during and after drug treatment. When post-drug thresholds had stabilized, subjects were sacrificed and their cochleas stained, embedded in plastic, microdissected, and viewed with phase contrast microscopy to permit examination of the cochlear tissue. Post-drug data suggest that frequency discrimination at a high frequency is unaffected by a 40- to 45-dB sensorineural hearing loss, considerable hair cell damage, and the resultant disruption of the cochlear micromechanics. The data, in concert with previously published reports, suggest that FDLs may be less affected by a high-frequency sensorineural hearing loss than by a low-frequency sensorineural hearing loss.     

The effect of superior auditory skills on vocal accuracy

The relationship between auditory perception and vocal production has been typically investigated by evaluating the effect of either altered or degraded auditory feedback on speech production in either normal hearing or hearing-impaired individuals. Our goal in the present study was to examine this relationship in individuals with superior auditory abilities. Thirteen professional musicians and thirteen nonmusicians, with no vocal or singing training, participated in this study. For vocal production accuracy, subjects were presented with three tones. They were asked to reproduce the pitch using the vowel /a/. This procedure was repeated three times. The fundamental frequency of each production was measured using an autocorrelation pitch detection algorithm designed for this study. The musicians' superior auditory abilities (compared to the nonmusicians) were established in a frequency discrimination task reported elsewhere. Results indicate that (a) musicians had better vocal production accuracy than nonmusicians (production errors of 1/2 a semitone compared to 1.3 semitones, respectively); (b) frequency discrimination thresholds explain 43% of the variance of the production data, and (c) all subjects with superior frequency discrimination thresholds showed accurate vocal production; the reverse relationship, however, does not hold true. In this study we provide empirical evidence to the importance of auditory feedback on vocal production in listeners with superior auditory skills.     

Hearing loss from simulated work-week exposure to impulse noise.

Six monaural chinchillas were exposed to a repetitive, reverberant, impulse noise for a total of five days, 8 h per day. The average peak overpressure within the holding cage was 113 dB. The reverberation time (pressure fluctuation envelope within 20 dB of peak) was 160 ms. Auditory thresholds were measured at 0.25, 0.5, 1, 2, 4, and 8 kHz before and after each day's exposure using either the average-evoked response technique or shock avoidance conditioning. After the last exposure, recovery was monitored for five successive days. Final thresholds were obtained starting at 30 days postexposure after which the animals were sacrificed for cochlear histology. The high frequencies (4, 8 kHz) showed a daily median shift of 40 dB and a 27 dB recovery before the following day's exposure. The low frequencies (0.25, 0.5 kHz) were shifted 35 dB after each day's exposure with a 15 dB recovery overnight. Final median audiograms showed little permanent threshold shift. The cochleagrams for two test animals were found to be normal while the remaining four displayed 10%--40% losses in hair cells at specific cochlear sites.     

Postnatal functional development of the dorsal and posteroventral cochlear nuclei of the cat

The postnatal development of firing patterns and response areas was determined for single neurons in the dorsal (DCN) and posteroventral ( PVCN ) cochlear nuclei of the kitten. Extracellular, single-unit responses to pure-tone stimulation were recorded in ketamine and sodium pentobarbital anesthetized kittens between the ages of 5 and 52 days. Within the first two weeks of postnatal life threshold is high, first-spike latency is long, and maximal discharge rate is low as compared to older kittens and adult cats. Prior to the end of the second postnatal week the tone-evoked temporal discharge patterns that characterize neurons of the DCN and PVCN in the adult cat are routinely recorded. These patterns, which appear within the first 50 ms of tonal stimulation, include the so-called " primarylike ," "chopper," " pauser ," "buildup," and "onset" types and their variants. In animals younger than about 10-12 days of age, the driven activity that occurs later than about 50 ms after stimulus onset often is not sustained, but breaks up during the stimulus into bursts that are separated by intervals of about 100-150 ms. Also within the first two weeks of postnatal life, many of the response-area properties of DCN and PVCN neurons are similar to those recorded in adult cats. The excitation and inhibition found within the so-called type II/III, type IV, and type V response areas of the adult occur in this early postnatal period. We conclude that many of the cellular mechanisms that underlie the temporal firing patterns and the organization of the response areas of DCN and PVCN neurons are active in the growing, differentiating cochlear nuclei and that the emergence of these mechanisms does not depend on afferent activity generated in the cochlear and auditory nerve by the animal's acoustic environment. Furthermore, if temporal firing patterns and response-area profiles remain relatively constant over the life span of the animal, then so must the spatial and temporal relationships of the inputs that produce and maintain them as these neurons, and the circuits of which they are a part, grow in size and complexity.