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Phase relationship between bone and air conducted impulse signals in the human head.

Using the cancellation method we investigated the phase relationship as well as the amplitude ratio between impulse signals transmitted simultaneously via both bone and air conduction channels. The psychophysical findings indicated a phase difference corresponding to a time delay of about 0.9 ms by which the air input led the bone input in the case of frontal bone stimulation. Changing the polarity of the bone conduction signal a maximum in the loudness sensation was found just in the same phase difference. The psychophysical findings, predicted mathematically, were verified with brainstem evoked potentials elicited to similar signals. Preliminary results showed that the time lag observed relied on the location of the bone vibrator on the head and furthermore on the frequency used. This suggests different transmissions of vibratory energy reaching the inner ear along the skull.

Bone Conduction↗

Intensity difference limens at high frequencies.

Thresholds for amplitude modulation detection were obtained from four subjects at frequencies of 2, 4, 6, 8, and 10 kHz for sensation levels of 15, 30, 45, and 60 dB and modulation rates of 2, 4, and 8 Hz. High-frequency difference limens calculated from amplitude modulation thresholds were found to change nonmonotonically as a function of sensation level, independent of modulation rate. This nonmonotonic relation stemmed mainly from a gradual reduction of the difference limen at the lowest sensation level with increasing frequency. Difference limens for pulsed tone discrimination were also measured in two of the subjects at 2, 6, and 10 kHz and sensation levels of 15, 30, 45, and 60 dB. The relation between intensity discrimination and sensation level was similar to that found for amplitude modulation detection. These findings are interpreted as indicating that the nonmonotonic relation between sensation level and intensity resolution is a general characteristic of stimulus processing at higher frequencies.

Adult↗

Lateralization of sinusoidally amplitude-modulated tones: effects of spectral locus and temporal variation.

It has long been recognized that listeners are sensitive to interaural temporal disparities (ITDs) of low-frequency (i.e., below 1600 Hz) stimuli. Within the last three decades, it has often been demonstrated that listeners are also sensitive to ITDs within the envelope of high-frequency, complex stimuli. Because these studies, for the most part, employed discrimination tasks, few data exist concerning the extent of laterality produced by ITDs as a function of the spectral locus of the stimulus. To this end, we employed an acoustic "pointing" task in which listeners varied the interaural intensity difference of a 500-Hz narrow-band noise (the pointer) so that it matched the intracranial position of a second, experimenter-controlled stimulus (the target). Targets were sinusoidally amplitude-modulated tones centered on 500 Hz, 1, 2, 3, or 4 kHz and modulated at rates ranging from 50 to 800 Hz. Targets were presented with either the entire waveform delayed or with only the envelope delayed. Our results suggest that: (1) for low-frequency targets, lateralization is influenced by ITDs in the envelope but is dominated by ITDs in the fine structure; (2) for high-frequency targets, envelope-based delays produce displacements of the acoustic images which are affected greatly by the rate of modulation; rather large extents of laterality could be produced with high rates of modulation; these data are consistent with those obtained previously in discrimination experiments; (3) for low rates of modulation (e.g., 100 Hz), delays of the entire waveform (both envelope and fine structure) produce much greater displacements of the acoustic image for low-frequency than for high-frequency targets (where fine-structure-based cues are not utilizable); (4) there appear to be no consistent relations among extent of laterality, rate of modulation, and the frequency of the carrier within and across listeners.

Adult↗

Localization of sound in rooms, II: The effects of a single reflecting surface.

Auditory localization was studied in a room bounded by a single acoustically reflective surface. The position of that surface was varied so as to stimulate a floor, a ceiling, and left and right side walls. The surface was eliminated in one condition so that we could examine localization in free field for purposes of comparison. Using a source identification method we assessed the influences of these various room configurations on the localization of both slow-onset and impulsive sine tones of low frequency (500 Hz). We also measured the steady-state interaural-time-difference (ITD) and interaural-intensity-difference (IID) cues available to subjects in the different room configurations and compared these data with the perceptual judgments. Our results indicate the following: (1) A sound must include transients if the precedence effect is to operate as an aid to its localization in rooms. (2) Even if transients are present the precedence effect does not eliminate all influences of room reflections. (3) Due to the interference of reflections large interaural intensity differences may occur in a room and these have a considerable influence on localization; this is true even at low frequencies for which IID cues do not exist in a free field. (4) Listeners appear to have certain expectations about the reliability and plausibility of various directional cues and perceptually weight the cues accordingly; we suggest that this may explain, in part, the large variation in time-intensity trading ratios reported in the literature and also the differing reports regarding the importance of onsets for localization. (5) In this study we find that onset cues are of some importance to localization even in free field.

Acoustic Stimulation↗

The owl's cochlear nuclei process different sound localization cues.

This paper discusses how the barn owl's brain stem auditory pathway is divided into two physiologically and anatomically segregated channels for separate processing of interaural phase and intensity cues for sound localization. The paper also points out the power of the "downstream" approach by which the emergence of a higher-order neuron's stimulus selectivity can be traced through lower-order stations.

Animals↗

Development of auditory-evoked potentials in the cat. I. Onset of response and development of sensitivity.

Auditory-evoked potentials, originating from the brain stem and the forebrain, were studied in 30 unanesthetized kittens during the first 3 months of postnatal life, and from a smaller set of animals before and after surgical exposure of their tympanic membranes. In intact animals, responses to 135-dB peak SPL clicks were first reliably discernible on the seventh postnatal day; when stimuli were presented directly to the exposed tympanic membrane, responses were observed several days earlier. Responses progressed through three stages during maturation: an early period of gross insensitivity during which responses are evoked only by high-intensity stimuli and whose response thresholds remain essentially constant (week 1); a middle period characterized by rapid acquisition of sensitivity to near-adult values (week 2); and a late period during which adult thresholds and latencies are acquired. A sequential stage model of threshold maturation is proposed, in which thresholds decline linearly during stage two and exponentially during stage three. It is hypothesized that mechanical reorganization of the cochlea during the first 2 to 3 postnatal weeks and development of the stria vascularis are primarily responsible for the linear stage, and that neural factors primarily underlie the exponential stage and account for the gradual acquisition of adult thresholds. Rates of maturation for brain stem responses are frequency dependent, with responses to high frequencies achieving adult thresholds earlier than those to low frequencies.

Aging↗

Development of auditory-evoked potentials in the cat. II. Wave latencies.

Brain stem and forebrain auditory-evoked potentials were studied parametrically during the first 90 postnatal days in unanesthetized kittens using tonal and click stimuli. This paper describes changes that occur in transmission time through the auditory pathway during development by analyses of the maturational time courses of latencies associated with waves of both auditory brain stem responses (ABR's) and late-occurring auditory-evoked potentials (AER's), recorded subdermally from the vertex. In response to click stimuli, ABR latencies were found to decay rapidly early in postnatal life and more slowly after the third postnatal week. Those trends were modeled as a two-stage sequential process, with a linear stage occurring between 7 and 18 postnatal days followed by an exponential stage during which adult latencies were achieved. AER latencies changes during development were less complicated, and followed a single-stage exponential time course. When threshold influences were taken into account--that is, when data were adjusted so that sensation level (SL) was constant across age--the latency-maturation curves associated with all ABR waves were adequately described by a single exponential, and latencies recorded from young animals were substantially shorter than latencies associated with the same aged animals when analyses were carried out with constant sound-pressure level (SPL) stimuli across age. In addition, the difference function, generated when isoasymptotic SPL and SL latency versus age functions were subtracted from one another, was also represented by an exponential curve, suggesting that at least two processes underlie the latency decay that occurs during postnatal development. Evoked responses to tonal stimuli throughout development were consistent with the basoapical developmental gradient that is observed anatomically.

Aging↗

Effects of peak pressure and energy of impulses.

Peak pressure has been one of the key parameters of impulse noise used to assess the hazard to hearing. It is used in most international noise exposure limits. France uses an A-weighted energy limit. There is a rough correspondence between peak pressure and the hazard to hearing for a given type of impulse noise. However, when the effects of different types of impulses are compared, this correspondence breaks down. One of the alternate measures of impulse intensity is weighted energy. Weighted energy is appealing for a number of reasons. It does not depend on details of the pressure-time history such as the peak pressure and the more common duration measures. It should be easier to integrate with continuous or intermittent noise standards. It would make it easier to use standard hearing protector attenuation to estimate the hazard when a specific hearing protector is worn. Results of previously published articles and reports will be discussed. These reports lead to the conclusion that weighted energy is a more potent determiner of hearing hazard than peak pressure if spectral effects are controlled.

Animals↗

Human temporary threshold shift (TTS) and damage risk.

Information regarding the relation of human temporary threshold shift (TTS) to properties of steady-state and intermittent noise published since the 1966 appearance of the CHABA damage risk contours is reviewed. The review focuses on results from four investigative areas relevant to potential revision of the CHABA contours including effects of long-duration exposure and asymptotic threshold shifts (ATS); equivalent quiet and/or safe noise levels; effects of intermittency; and use of noise-induced temporary threshold shift (NITTS) to predict susceptibility to noise-induced permanent threshold shift (NIPTS). These data indicate that two of three major postulates on which the original contours were based are not valid. First, recovery from TTS is not independent of the conditions that produced the TTS as was assumed. Second, the assumption that all exposures that produce equal TTS2 are equally hazardous is not substantiated. The third postulate was that NIPTS produced by 10 years of daily exposure is approximately equal to the TTS2 produced by the same noise after an 8-h exposure. Based upon several TTS experiments showing that TTS reaches an asymptote after about 8 h of exposure, the third CHABA postulate can be reworded to state the hypothesis that ATS produced by sound of fixed level and spectrum represents an upper bound on PTS produced by that sound regardless of the exposure duration or the number of times exposed. This hypothesis has a strong, logical foundation if ATS represents a true asymptote for TTS, not a temporary plateau, and if threshold shifts do not increase after the noise exposure ceases.

Auditory Fatigue↗

Hearing loss in the chinchilla from impact and continuous noise exposure.

The relative hazard posed to the peripheral auditory system by impact/impulse and continuous noise of the same power spectrum was determined. Impact noise was generated by striking a nail with a hammer and was digitally recorded. The acoustical power spectrum of the impact was determined and pink noise was filtered to produce a continuous noise stimulus with the same acoustic power spectrum. Pre-exposure auditory evoked response (AER) thresholds were obtained at 1, 2, 4, and 8 kHz on 16 adult chinchillas. The pool of animals was divided into two equal groups based upon pre-exposure AER thresholds. One group was exposed to impact noise and the other group to the filtered pink noise. Exposures were 4 h/day for 5 days. Thirty days following the exposure, auditory evoked response thresholds were remeasured. Changes in auditory sensitivity were determined by subtracting the pre-exposure thresholds from the post-exposure thresholds. Hearing threshold shifts of the impact noise group were significantly greater (p less than 0.0001) than the hearing threshold shifts of the continuous noise group. These data indicate a need to more closely examine the parameters and effects of impact noise. There may be a need to develop expanded damage-risk criteria for occupational exposure to impulse/impact noise.

Animals↗

The importance of "temporal pattern" in traumatic impulse noise exposures.

The equal energy hypothesis (EEH) was evaluated for impulse noise. Specifically, the experiments evaluated the importance of the temporal distribution of impulses; the trading relation between the number of impulses and peak level and the difference between continuous and impulse noise. Monaural chinchillas were exposed to one of seven conditions. Their hearing was evaluated before, immediately after, and 30 days after the exposure. Hair cell damage was reported in the form of a cochleogram. The experiments show that the EEH is more appropriate for low-level impulse (135-dB peak); for equal amounts of energy, 150-dB impulses produce more hearing loss and hair cell damage than 135-dB impulses; for equal amounts of energy, impulses presented in rapid bursts cause less damage than impulses presented at "1/s" and 50 microseconds. Pairs of impulses presented at "1/s" produce the largest amount of damage. The results are discussed in terms of implications for the EEH.

Animals↗

Some features of binaural input to single neurons in physiologically defined area AI of cat cerebral cortex.

1. In the ectosylvian cortex of 24 barbiturate-anesthetized cats, area AI was identified by its frequency organization and the responses to tonal stimuli of single neurons in that field were examined using sealed stimulating systems incorporating calibrated probe microphone assemblies. 2. The responsiveness to monaural and binaural best-frequency stimuli was examined quantitatively for 282 single units in AI. One hundred thirty-nine cells (49%) were excited by independent stimulation of only one ear and were classified as EO cells. In general, the effective monaural excitatory input was derived from the contralateral ear. One hundred ten (39%) neurons were excited by independent stimulation of each ear and were classified as EE units. For these neurons, the contralateral responses were generally stronger, shorter in latency, and lower in threshold than were their ipsilateral responses. Thirty-three cells (12%) gave weak or no responses to monaural stimuli but responded securely to binaural stimuli. These cells were classified as predominantly binaural (PB). 3. Binaural interactions were examined by comparison of the response to binaural, equally intense stimuli to the stronger monaural response. Among EO cells suppression was the most common form of interaction, while for EE cells summation was the more common. Less than 8% of cells were found to be monaural. 4. In electrode penetrations radial to the cortex surface, cells received their stronger or sole monaural excitatory input from a common ear, generally the contralateral. Within such penetrations, however, cells commonly differed with regard to the nature of their input from the other ear and/or in their binaural interactions. 5. Comparison of these data with data previously reported for subcortical auditory nuclei revealed that AI preserves many of the stimulus specificity characteristics of the lower nuclei. The reasons for the preservation of these characteristics at the cortex and the implications of the present data for the binaural column hypothesis are discussed.

Animals↗

Responses of single neurons in posterior field of cat auditory cortex to tonal stimulation.

In the auditory cortex of barbiturate-anesthetized cats, the posterior auditory field (field P) was identified by its tonotopic organization, and single neurons in that field were studied quantitatively for their sensitivity to the frequency and intensity of tonal stimuli presented via calibrated, sealed stimulating systems. Field P neurons had narrow, V-shaped, threshold frequency tuning curves. At suprathreshold levels, spike counts were generally greatest at frequencies at or close to the neuron's threshold best frequency (BF). Eighty-six percent of posterior-field neurons displayed spike counts that were a nonmonotonic function of the intensity of a BF tone. Of these, over 90% showed at least a 50% reduction in spike count at high stimulus levels, and almost 20% of nonmonotonic cells ceased responding entirely at high stimulus intensities. The nonmonotonic shape of spike count-versus-intensity profiles was typically preserved across the range of frequencies to which any given neuron was responsive. For some neurons, this had the consequence of generating a completely circumscribed frequency-intensity response area. That is, these neurons responded to a tonal stimulus only if the stimulus was within a restricted range of both frequency and intensity. These response areas showed internal organizations that appeared to reflect one or both of two processes. For some neurons, the optimal sound pressure level for spike counts varied with tone frequency, roughly paralleling the threshold tuning curve. For other neurons, the optimal sound pressure level tended to be constant across frequency despite threshold variations of up to 20 dB. The minimum response latencies of posterior-field neurons were generally in the range of 20-50 ms, while cells in the primary auditory cortex (AI) in the same animals generally had minimum latent periods of less than 20 ms. Comparison of these data with those previously presented for neurons in two other cortical auditory fields suggests that the cat's auditory cortex might show an interfield segregation of neurons according to their coding properties.

Animals↗

Interaural phase-sensitive units in the inferior colliculus of the unanesthetized rabbit: effects of changing frequency.

We studied the interaural phase sensitivity of 85 units in the inferior colliculus (IC) of the unanesthetized rabbit. We assessed this sensitivity at several frequencies within each unit's responsive range. The interaural phase disparity was varied by delivering tones that differed by 1 Hz to the two ears, resulting in a 1-Hz binaural beat. We analyzed each unit's response to different frequencies by calculating four measures: characteristic delay (CD), characteristic phase (CP), composite peak delay, and mean peak delay. We estimated the CD and CP from the slope and phase intercept, respectively, of the regression line fitted to a plot of the mean interaural phase against stimulating frequency. The composite peak delay was estimated from the peak of a composite delay curve. This was generated by replotting the response to changes in interaural phase, as a function of the equivalent interaural delay and averaging the resultant interaural delay curves. The composite delay curve reflects the unit's average response to interaural delays across frequencies. Last, we calculated a mean peak delay, derived by converting the mean interaural phase of the response at each frequency to an equivalent delay and then averaging these delays. Interaural phase sensitivity was observed to frequencies as high as 2,150 Hz. However, the majority of units showed such sensitivity below 1,500 Hz. For most units, the interaural delay curves measured at several frequencies coincided near the peak discharge. This result is consistent with a neural model, where excitatory inputs from each ear converge upon a binaural cell, evoking maximum discharge only when the two inputs arrive simultaneously. As a first approximation, our data fit this model, indicating that IC neurons can act like coincidence detectors or cross-correlators. The distributions of CD, composite peak delay, and mean peak delay showed that most units preferred ipsilateral stimulus delays, which in the natural situation corresponds to sounds emanating from the contralateral field. Moreover, most units preferred delays that were within the estimated physiological range of the rabbit. These results support the viewpoint that neurons in the IC participate in sound localization. The distributions of CP and CD differ substantially from those found in the IC of the anesthetized cat. These differences may reflect species differences, the effects of anesthesia, or a difference in the population of units sampled. For each unit, we assessed the linearity of the plot of mean interaural phase against frequency of stimulation using a chi 2 method. For most units the plots were significantly nonlinear.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Neural encoding of single-formant stimuli in the cat. I. Responses of auditory nerve fibers.

1. We have studied auditory responses to a set of speech-related narrowband sounds, single-formant stimuli (SFSs), in populations of auditory nerve fibers (ANFs). An analytic method was developed to extract the envelope of temporal discharge patterns of the ANF responses to nonsinusoidally modulated stimuli, whose spectra have multiple clusters of components. Such responses are often encountered in the auditory system when complex stimuli are used and have traditionally been studied by analyzing the fundamental component of the responses. 2. The envelope modulation in the SFSs is shown to be represented by the response patterns of ANFs. When the whole ANF population is considered, the information on modulation in stimulus envelope does not disappear at the highest sound level tested at all best frequencies (BFs) we studied (1-10 kHz). The representation is the best at medium sound levels and degrades at high sound levels. Low/medium-spontaneous rate (SR) ANFs showed greater envelope modulation in their responses at high sound levels than do high-SR ANFs. The quality of the representation at high sound levels is, on average, proportional to BF threshold of an ANF. On the basis of populations of ANFs with all SRs, the envelope modulation in the SFSs is represented over a wide range of sound levels. 3. We found that low-BF ANFs differ from high-BF ANFs in representing envelope modulation in the SFSs. For ANFs with BFs less than approximately 6 kHz, information on stimulus envelope is not only contained in spectral components near direct current but also in components at the vicinities of frequencies equal to BF and its multiples. In fact, for ANFs with BFs < 3 kHz, the contribution from spectral components centered at BF to overall response modulation is greater than that from spectral components near direct current. These findings indicate that, by using measures solely based on the fundamental component, the amount of modulation in the responses to narrowband stimuli is underestimated for low-BF ANFs. 4. Off-BF stimulation of ANFs with SFSs was found to result in increased envelope modulation in responses at high sound levels. The further away the stimulus is centered relative to unit BF, the greater the modulation it induces, provided that the stimulus is capable of exciting the unit. An SFS centered as close as 15% off unit BF can produce a significant increase in the modulation of responses at very high sound levels.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗