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Amphibious auditory responses of the American alligator (Alligator mississipiensis).

Animals that thrive both on land and underwater are faced with the task of interpreting stimuli in different media. This becomes a challenge to the sensory receptors in that stimuli (e.g., sound, motion) may convey the same type of information but are transmitted with different physical characteristics. We used auditory brainstem responses to examine hearing abilities of a species that makes full use of these two environments, the American alligator (Alligator mississipiensis). In water, alligators responded to tones from 100 Hz to 2,000 Hz, with peak sensitivity at 800 Hz. In air, they responded to tones from 100 Hz to 8,000 Hz, with peak sensitivity around 1,000 Hz. We also examined the contribution to hearing of an air bubble that becomes trapped in the middle ear as the animal submerges. This bubble has been previously implicated in underwater hearing. Our studies show that the trapped air bubble has no affect on auditory thresholds, suggesting the bubble is not an important adaptation for underwater hearing in this species.

Air↗

Low-frequency distortion product otoacoustic emissions in two species of kangaroo rats: implications for auditory sensitivity.

Low-frequency distortion-product otoacoustic emissions were measured in two species of kangaroo rats to test the prediction that a large footdrumming species (e.g., Dipodomys spectabilis) would have greater distortion-product otoacoustic emission amplitude than a small non-footdrumming species (e.g., Dipodomys merriami), indicating better hearing sensitivity at low frequencies. Equal-level (65 dB SPL) stimulus tones ( f(1), f(2)), presented over a ( f(1)) range of 200-1000 Hz, were used to evoke the 2 f(1)- f(2) distortion-product otoacoustic emission. Mean 2 f(1)- f(2) levels for D. merriami showed good correspondence to previously published audiograms for that species. Mean 2 f(1)- f(2) levels and 95% confidence intervals indicated species differences below 400 Hz, supporting the theory that low-frequency hearing sensitivity is better in large kangaroo rat species. These results suggest that the size-related divergence in footdrumming behavior may be related to differential auditory sensitivity.

Animals↗

Auditory response characteristics of the piebald odorous frog and their implications.

The piebald odorous frog (Odorrana schmackeri), the large odorous frog (Odorrana livida) and the concave-eared torrent frog (Amolops tormotus) are sympatric species living near the same torrent streams in the vicinity of Mt. Huangshan, China. A recent study demonstrated that A. tormotus can use sound signals involving ultrasonic components for communication in a noisy environment, and another sympatric species, O. livida, can also perceive ultrasonic sound. Here we report data on the hearing range of O. schmackeri by studying auditory evoked potentials and single-unit data from the torus semicircularis. This frog exhibits its two most sensitive peaks at 2 kHz and 3.5-4.0 kHz with thresholds <42 dB SPL, with an upper frequency limit of hearing at 8.5 kHz with threshold of 87 dB SPL. The upper limit is much lower than those of O. livida and A. tormotus, at 22 and 34 kHz, respectively. It suggests that sympatric species may respond differently to similar environmental selection pressures sculpting auditory communication systems.

Acoustic Stimulation↗

Integration and recovery processes contribute to the temporal selectivity of neurons in the midbrain of the northern leopard frog, Rana pipiens.

This study examined the mechanisms underlying amplitude modulation selectivity in the anuran auditory midbrain. Single units were recorded extracellularly in the torus semicircularis of the northern leopard frog, Rana pipiens. Two physiologically distinct classes of neurons were identified, based on their response latencies and their selectivities to pulse repetition rates. Cells in one group had short response latencies (median = 31 ms) and responded best to pulse repetition rates below 40 Hz. Tuning to low amplitude modulation rates was largely determined by recovery processes and phasic response properties. Cells in the second group had much longer latencies (median=81 ms) and were generally selective for pulse repetition rates greater than 40-50 Hz. Sensitivity to higher amplitude modulation rates resulted from integration processes; these units only responded when a threshold number of pulses were presented at a minimum pulse density (amplitude modulation rate). At amplitude modulation rates above their best rate, their responses decreased, apparently due to inadequate recovery time between pulses.

Acoustic Stimulation↗

Coding of concurrent vocal signals by the auditory midbrain: effects of duration.

Neural selectivity to signal duration within the auditory midbrain has been observed in several species and is thought to play a role in signal recognition. Here we examine the effects of signal duration on the coding of individual and concurrent vocal signals in a teleost fish with exceptionally long duration vocalizations, the plainfin midshipman, Porichthys notatus. Nesting males produce long-duration, multi-harmonic signals known as hums to attract females to their nests; overlapping hums produce acoustic beats at the difference frequency of their spectral components. Our data show that all midbrain neurons have sustained responses to long-duration hum-like tones and beats. Overall spike counts increase linearly with signal duration, although spike rates decrease dramatically. Neurons show varying degrees of spike rate decline and hence, differential changes in spike rate across the neuron population may code signal duration. Spike synchronization to beat difference frequency progressively increases throughout long-duration beats such that significant difference frequency coding is maintained in most neurons. The significance level of difference frequency synchronization coding increases by an order of magnitude when integrated over the entirety of long-duration signals. Thus, spike synchronization remains a reliable difference frequency code and improves with integration over longer time spans.

Animals↗

Multidimensional assessment of functional outcomes of medialization thyroplasty.

Medialization thyroplasty (MT) is the most widely used laryngeal framework phonosurgical procedure for managing glottic incompetence secondary to unilateral vocal fold paralysis (UVFP). The aim of the study was to evaluate the functional outcomes of MT in 32 UVFP patients, comparing multidimensional perceptual and instrumental measures of voice before and after surgery, and to evaluate how close to normal that postoperative voice measure became. Video laryngostroboscopy (VLS), auditory perceptive evaluation of voice, the patients' self-evaluation of hoarseness on the Visual Analogue Scale (VAS) and calculation of the Voice Handicap Index (VHI), analysis of objective acoustic voice parameters, quantitative assessment of phonetograms and measurement of maximum phonation time were conducted. Vocal function was evaluated before the surgery and for the period from 1 month to 3 years (Mo 1.0 month; Me 2.0 months) after MT. VLS confirmed remarkable medialization of the paralyzed vocal fold. As a consequence, hoarseness and breathiness were found to be significantly decreased after MT. Pitch and intensity range and phonetogram area were significantly increased. A significant decrease of jitter, shimmer and normalized noise energy reflected improvement of the stability of acoustic signal and a more efficient pattern of phonation. Thus, the perceptual and acoustic voice parameters studied showed statistically significant differences (P < 0.001) between preoperative and postoperative voices, and these objective measurements of voice changes provided accurate and documentary evidence of the results of surgical treatment. A high degree of patient satisfaction with the MT was confirmed by a significant decrease of VHI and hoarseness on VAS. Thus, results of the present investigation confirm the functionality and effectiveness of MT in patient voice rehabilitation with UVFP. However, the means of acoustic voice parameters measured in the study did not reach normal limits, probably because of the remaining underlying condition of UVFP.

Adolescent↗

Resonant modes and musical ratios in otoacoustic emissions.

The ratios between frequency components of evoked otoacoustic emissions (OAE) were investigated for 100 ears. The signals were decomposed by means of an adaptive approximation method into basic waveforms coming from a very large and redundant dictionary of Gabor functions. The high time-frequency resolution of the method and the parametric representation of the waveforms allowed for an estimation of the frequency ratios of the basic components. A repetitive occurrence of the "fifths", "fourths" and octaves connected with the Pythagorean temperament was found. The octaves containing "fifths" were identified. This kind of sequences in OAE tend to appear in the same form for tonal stimulations of different frequencies and for broadband stimuli. The significance of the results was confirmed by comparison to Monte Carlo simulations of the null hypothesis of random distribution of frequency modes. These findings support the resonance theory of hearing, which binds musical ratios with the geometrical spacing of outer hair cells in the cochlea.

Acoustic Stimulation↗

Behaviour profiles in a population of infants later diagnosed as having autistic disorder.

In order to complement findings in the field of early autism and in the context of our studies on the quantitative neurobiological evaluation of patients with autism, we studied a large population of infants later diagnosed as having autistic disorder, using multivariate descriptive statistical methods. The population included 74 infants between six and 35 months, evaluated with the Infant Behavioural Summarised Evaluation (IBSE) scale. Thirteen of the 33 items of the IBSE scale were selected according to previous studies on older patients with autistic disorder. Correspondence analysis was applied to these 13 items, followed by a classical cluster analysis. This procedure permitted the identification of four different profiles which were distinguished on the basis of five main behaviours: activity, auditory perception, sensorimotility, eye contact and use of objects. These profiles are similar to those identified in older children with autism. This study provides an objective description of early clinical markers of the heterogeneity in autism and thus contributes to the description of the onset of autistic disorder. It identified more precise clinical subtyping which will be valuable for future bioclinical studies.

Autistic Disorder↗

Auditory processing of spectral cues for sound localization in the inferior colliculus.

The head-related transfer function (HRTF) of the cat adds directionally dependent energy minima to the amplitude spectrum of complex sounds. These spectral notches are a principal cue for the localization of sound source elevation. Physiological evidence suggests that the dorsal cochlear nucleus (DCN) plays a critical role in the brainstem processing of this directional feature. Type O units in the central nucleus of the inferior colliculus (ICC) are a primary target of ascending DCN projections and, therefore, may represent midbrain specializations for the auditory processing of spectral cues for sound localization. Behavioral studies confirm a loss of sound orientation accuracy when DCN projections to the inferior colliculus are surgically lesioned. This study used simple analogs of HRTF notches to characterize single-unit response patterns in the ICC of decerebrate cats that may contribute to the directional sensitivity of the brain's spectral processing pathways. Manipulations of notch frequency and bandwidth demonstrated frequency-specific excitatory responses that have the capacity to encode HRTF-based cues for sound source location. These response patterns were limited to type O units in the ICC and have not been observed for the projection neurons of the DCN. The unique spectral integration properties of type O units suggest that DCN influences are transformed into a more selective representation of sound source location by a local convergence of wideband excitatory and frequency-tuned inhibitory inputs.

Acoustic Stimulation↗

Forward masking additivity and auditory compression at low and high frequencies.

The additivity of nonsimultaneous masking can be used as a measure of nonlinearity in the auditory system. For example, two equally effective forward maskers should produce 3 dB of additional masking when they are combined, assuming linearity with respect to intensity. A combined effect greater than this (excess masking) indicates compression. In the present experiments, the signal was a 10-ms pure tone presented 20 ms after a 200-ms narrowband noise masker and/or immediately after a 20-ms narrow-band noise masker. The signal frequency was 250, 500, or 4000 Hz. The signal threshold produced by combining two equally effective maskers was measured. At all three frequencies, little excess masking was observed for a signal 10 dB above absolute threshold, indicating linear additivity (no compression). At signal levels 30 dB above absolute threshold, excess masking was observed at all three frequencies. The estimated compression exponents were 0.29 at 250 Hz, 0.34 at 500 Hz, and 0.17 at 4000 Hz. In contrast with physiological studies on other mammals, the present results provide evidence for substantial compression at low frequencies in humans.

Acoustic Stimulation↗

Age-dependent changes of gap detection in the Mongolian gerbil (Meriones unguiculatus).

Gap detection using broadband noise was characterized in a group of young gerbils from the breeding colony of the University of Regensburg (RB gerbils), old RB gerbils, and old gerbils from the breeding colony of the University of South Carolina (SC gerbils). Data from old RB and old SC gerbils were not significantly different and were subsequently combined for a comparison with data from the group of young RB gerbils. Level dependence of gap-detection thresholds in young and old domesticated gerbils resembled the typical mammalian pattern of level dependence. Gap-detection thresholds of old gerbils were significantly elevated at 30 dB SL and 50 dB SPL as compared with young gerbils. Compared with young gerbils tested at 30 dB SL and 50 dB SPL, the distribution of gap-detection thresholds in old gerbils was broader with a spread to higher gap-detection thresholds. Some old animals retained excellent temporal resolution, while some showed impaired gap detection. The gap-detection data collected in young and old gerbils resemble previously published data from humans of different age and confirm that gerbils are a useful model to study age-dependent changes in temporal processing.

Acoustic Stimulation↗

Cortical responses to cochlear implant stimulation: channel interactions.

This study examined the interactions between electrical stimuli presented through two channels of a cochlear implant. Experiments were conducted in anesthetized guinea pigs. Multiunit spike activity recorded from the auditory cortex reflected the cumulative effects of electric field interactions in the cochlea as well as any neural interactions along the ascending auditory pathway. The cochlea was stimulated electrically through a 6-electrode intracochlear array. The stimulus on each channel was a single 80- micro s/phase biphasic pulse. Channel interactions were quantified as changes in the thresholds for elevation of cortical spike rates. Experimental parameters were interchannel temporal offset (0 to +/-2000 micro s), interelectrode cochlear spacing (1.5 or 2.25 mm), electrode configuration (monopolar, bipolar, or tripolar), and relative polarity between channels (same or inverted). In most conditions, presentation of a subthreshold pulse on one channel reduced the threshold for a pulse on a second channel. Threshold shifts were greatest for simultaneous pulses, but appreciable threshold reductions could persist for temporal offsets up to 640 micro s. Channel interactions varied strongly with electrode configuration: threshold shifts increased in magnitude in the order tripolar, bipolar, monopolar. Channel interactions were greater for closer electrode spacing. The results have implications for design of speech processors for cochlear implants.

Animals↗

Towards a unifying basis of auditory thresholds: the effects of hearing loss on temporal integration reconsidered.

For signal detection and identification, the auditory system needs to integrate sound over time. It is frequently assumed that the quantity ultimately integrated is sound intensity and that the integrator is located centrally. However, we have recently shown that absolute thresholds are much better specified as the temporal integral of the pressure envelope than of intensity, and we proposed that the integrator resides in the auditory pathway's first synapse. We also suggested a physiologically plausible mechanism for its operation, which was ultimately derived from the specific rate of temporal integration, i.e., the decrease of threshold sound pressure levels with increasing duration. In listeners with sensorineural hearing losses, that rate seems reduced, but it is not fully understood why. Here we propose that in such listeners there may be an elevation in the baseline above which sound pressure is effective in driving the system, in addition to a reduction in sensitivity. We test this simple model using thresholds of cats to stimuli of differently shaped temporal envelopes and durations obtained before and after hearing loss. We show that thresholds, specified as the temporal integral of the effective pressure envelope, i.e., the envelope of the pressure exceeding the elevated baseline, behave almost exactly as the lower thresholds, specified as the temporal integral of the total pressure envelope before hearing loss. Thus, the mechanism of temporal integration is likely unchanged after hearing loss, but the effective portion of the stimulus is. Our model constitutes a successful alternative to the model currently favored to account for altered temporal integration in listeners with sensorineural hearing losses, viz., reduced peripheral compression. Our model does not seem to be at variance with physiological observations and it also qualitatively accounts for a number of phenomena observed in such listeners with suprathreshold stimuli.

Animals↗

The role of suppression in the upward spread of masking.

The upward spread of masking refers to the higher growth rate of masking for maskers lower in frequency than the signal, compared to maskers at the signal frequency (Wegel RL, Lane CE. The auditory masking of one pure tone by another and its possible relation to the dynamics of the inner ear. Physics Rev. 23:266-285, 1924; Egan JP, Hake HW. On the masking pattern of a simple auditory stimulus. J. Acoust. Soc. Am. 22:622-630, 1950; Delgutte B. Physiological mechanisms of psychophysical masking: Observations from auditory-nerve fibres. J. Acoust. Soc. Am. 87:791-809, 1990a, Delgutte B. Two-tone rate suppression in auditory-nerve fibres: Dependence on suppressor frequency and level. Hear Res. 49:225-246, 1990b). The upward spread of simultaneous masking may arise from a combination of excitatory and suppressive effects. In this study, growth of masking functions were obtained for a 4-kHz signal masked by an on-frequency (4 kHz) or off-frequency (2.4 kHz), simultaneous or forward masker, in the presence of a notched noise with a center frequency of 4 kHz presented to restrict off-frequency listening. Compression was estimated from the slopes of the off-frequency growth of masking functions. Suppression was estimated by comparing the off-frequency simultaneous- and forward-masked growth of masking functions. Results showed that, for midlevel signals (35-60 dB SPL), the compression exponent estimated from simultaneous and forward masking averaged 0.31 and 0.26, respectively. The maximum amount of suppression (defined as the decrease in the basilar-membrane response to the signal) was variable, ranging from about 6 to 17 dB across subjects. Despite the substantial reduction in the response to the signal, the results suggest that suppression has a minimal effect on the slope of the masking function at mid levels. Rather, upward spread of masking seems to be mainly determined by the compressive basilar-membrane response to the signal in relation to the linear response to the lower-frequency masker.

Acoustic Stimulation↗

Temporal masking in electric hearing.

Temporal masking can be defined as the detection threshold of a brief signal as a function of the signal delay in a relatively long masker. The temporal masking pattern in normal acoustic hearing reveals temporal edge enhancement in which the signal detection threshold is greater near the masker onset than in the steady-state portion. Both peripheral and central mechanisms appear to underlie temporal edge enhancement, but their relative contributions remain elusive. Cochlear implants bypass cochlear mechanical processing and stimulate the auditory nerve directly, thereby providing a unique opportunity to separate the peripheral mechanisms from the central mechanisms. Here, we systematically measured temporal masking in electric hearing by examining whether a brief signal was harder to detect at the onset than in the steady-state portion of a long masker (the "overshoot" effect). The signal and the masker were presented (1) either to the same electrode or to different electrodes, (2) at the same stimulation or different rates, and (3) in a simultaneous or an interleaved fashion. A consistent pattern of results was observed, depending on the stimulus configuration between the signal and the masker. Simultaneous stimulation at the same rate and with the same electrode produced no difference in sensitivity between the onset and the steady-state conditions, but interleaved stimulation at different rates or with different electrodes produced a significant difference. Unlike acoustic hearing, high masker levels produced an overshoot effect, and low masker levels produced an undershoot effect. Although the present results are consistent with the "on-frequency vs. off-frequency" hypothesis for the overshoot effect, results also suggest a central "same vs. different" mechanism underlying temporal masking. These results have practical implications for improving cochlear implant design.

Acoustic Stimulation↗

Assessing tinnitus and prospective tinnitus therapeutics using a psychophysical animal model.

Subjective tinnitus is a common and often debilitating disorder that is difficult to study because it is a perceptual state without an objective stimulus correlate. Studying tinnitus in humans is further complicated by the heterogeneity of tinnitus quality, severity, and associated hearing loss. As a consequence, the pathophysiology of tinnitus is poorly understood and treatments are often unsuccessful. In the present study, an animal psychophysical model was developed to reflect several features of tinnitus observed in humans. Chronic tinnitus was induced in rats by a single intense unilateral exposure to noise. The tinnitus was measured using a psychophysical procedure, which required the animals to discriminate between auditory test stimuli consisting of tones, noise, and 0 dB. Tinnitus was indicated by a frequency-specific shift in discrimination functions with respect to control subjects not exposed to noise. The psychophysical consequences of the noise exposure were best explained by a tinnitus hypothesis and could not be explained easily by other consequences of noise exposure such as hearing loss. The qualitative features of the tinnitus were determined and related to the duration of noise exposure and the associated cochlear trauma. The tinnitus was found to persist and intensify over 17 months of testing. Finally, the tinnitus was reversibly attenuated by treatment with gabapentin, a GABA agonist. It was concluded that this model reflected several features of human tinnitus, such as its tonality and persistence, and could be useful as a screen for potential therapeutics as well as a tool to help unravel the pathophysiology of the disorder of phantom auditory perception.

Acetates↗

Informational masking in listeners with sensorineural hearing loss.

Measures of energetic and informational masking were obtained from 46 listeners with sensorineural hearing loss. The task was to detect the presence of a sequence of eight contiguous 60-ms bursts of a pure tone embedded in masker bursts that were played synchronously with the signal. The masker was either a sequence of Gaussian noise bursts (energetic masker) or a sequence of random-frequency 2-tone bursts (informational masker). The 2-tone maskers were of two types: one type that normally tends to produce large amounts of informational masking and a second type that normally tends to produce very little informational masking. The two informational maskers are called "multiple-bursts same" (MBS), because the same frequency components are present in each burst of a sequence, and "multiple-bursts different" (MBD), because different frequency components are presented in each burst of a sequence. The difference in masking observed for these two maskers is thought to occur because the signal perceptually segregates from the masker in the MBD condition but fuses with the masker in MBS. In the present study, the effectiveness of the MBD masker, measured as the signal-to-masker ratio at masked threshold, increased with increasing hearing loss. In contrast, the signal-to-masker ratio at masked threshold for the MBS masker changed much less as a function of hearing loss. These results suggest that sensorineural hearing loss interferes with the ability of the listener to perceptually segregate individual components of complex sounds. The results from the energetic masking condition, which included critical ratio estimates for all listeners and auditory filter characteristics for a subset of the listeners, indicated that increasing hearing loss also reduced frequency selectivity at the signal frequency. Overall, these results suggest that the increased susceptibility to masking observed in listeners with sensorineural hearing loss is a consequence of both peripheral and central processes.

Acoustic Stimulation↗

Varying feedback to evaluate detection strategies: the detection of a tone added to noise.

The effectiveness of rewarding strategies associated with different models of detection was evaluated for a tone-in-noise detection task. In three experiments, a tone was added to a narrowband noise masker in a two-interval, forced-choice procedure with feedback. In the first experiment the observer's task was to detect which interval had the added tone. In the second experiment the task was to indicate which interval was more intense. In the third experiment the task was to indicate which interval had the sound with the less modulated envelope. In terms of correct signal detection, thresholds were nearly the same for all three experiments. However, for three of the observers the psychometric functions indicate that observers altered their responses in accordance with the changes in feedback contingencies. This indicates that neither changes in level nor changes in envelope pattern alone can account for the detection of a tone added to noise for these observers. The fourth observer was sufficiently insensitive that the pattern of feedback associated with the level and envelope models did not differ substantially, and for her feedback had little impact on behavior. The results indicate that changes in feedback provide a means of testing alterative psychophysical models. Moreover, the results suggest that it is difficult to model the detection of a tone added to noise in terms of a single cue or a simple combination of cues. Observers are free to choose from at least two cues, and can do so.

Acoustic Stimulation↗