Frequency-specific maturation of the eighth nerve and brain-stem auditory pathway: evidence from derived auditory brain-stem responses (ABRs).

Previous studies of human auditory development using frequency-specific auditory brain-stem responses (ABRs) have reported that maturation for both peak and interpeak latencies occurs earlier for responses generated by low-frequency stimuli. In two of these studies, low-frequency ABRs presumed to originate from apical locations in the cochlea were likely dominated by activity from higher frequency regions closer to the base. In the present study, the high-pass noise-masking technique was used to generate derived ABRs that represent activity from isolated place specific regions along the basilar membrane. Analysis of auditory brain-stem maturation based on I-V interpeak latency differences with adult means revealed a frequency-specific pattern of development. Developmental changes occurred faster and mature function was attained earlier for ABRs from the mid-center-frequency (CF) derived conditions than from either the highest or lowest CF derived conditions. The differential maturation of mid-CF derived ABRs may reflect the delayed effects of the pattern of development that occurs in the cochlea.     

The effects of sensory hearing loss on cochlear filter times estimated from auditory brainstem response latencies.

Derived-band auditory brainstem responses (ABRs) were obtained in 43 normal-hearing and 80 cochlear hearing-impaired individuals using clicks and high-pass noise masking. The response times across the cochlea [the latency difference between wave V's of the 5.7- and 1.4-kHz center frequency (CF) derived bands] were calculated for five levels of click stimulation ranging from 53 to 93 dB p.-p.e. SPL (23 to 63 dB nHL) in 10-dB steps. Cochlear response times appeared to shorten significantly with hearing loss, especially when the average pure tone (1 to 8 kHz) hearing loss exceeded 30 dB. Examination of derived-band latencies indicates that this shortening is due to a dramatic decrease of wave V latency in the lower CF derived band. Estimates of cochlear filter times in terms of the number of periods to maximum response (Nmax) were calculated from derived-band latencies corrected for gender-dependent cochlear transport and neural conduction times. Nmax decreased as a function of hearing loss, especially for the low CF derived bands. The functions were similar for both males and females. These results are consistent with broader cochlear tuning due to peripheral hearing loss. Estimating filter response times from ABR latencies enhances objective noninvasive diagnosis and allows delineation of the differential effects of pathology on the underlying cochlear mechanisms involved in cochlear transport and filter build-up times.     

Anomalous phase relations in threshold-level responses from gerbil auditory nerve fibers

Phase-locked responses at near-threshold levels obtained from single units in the auditory nerve of gerbils show that the relation between phase lag and linear frequency often contains an unexpected microstructure. In frequency regions below 1 kHz that are also an octave or more below CF, phase curves often have multiple straight-line segments, rapid slope changes, or other major inflections. These detailed features of the phase curves are not accounted for by any identified artifact. For example, the anomalous features cannot be removed simply by lowering the stimulus level; they remain down to levels where phase locking first occurs. Phase curves for single units with the same CF recorded from different animals can have similar microstructures, suggesting that the form of the phase curves may reflect some joint effect of stimulus frequency and CF or longitudinal position.     

The behavioral response of mice to gaps in noise depends on its spectral components and its bandwidth

The purpose of these experiments was to determine whether detecting brief decrements in noise level ("gaps") varies with the spectral content and bandwidth of noise in mice as it does in humans. The behavioral effect of gaps was quantified by their inhibiting a subsequent acoustic startle reflex. Gap durations from 1 to 29 ms were presented in five adjacent 1-octave noise bands and one 5-octave band, their range being 2 kHz to 64 kHz. Gaps ended 60 ms before the startle stimulus (experiment 1) or at startle onset (experiment 2). Asymptotic inhibition was greater for higher-frequency 1-octave bands and highest for the 5-octave band in both experiments, but time constants were related to frequency only in experiment 1. For the lowest band (2-4 kHz) neither noise decrements (experiment 1 and 2) nor increments (experiment 3) had any behavioral consequence, but this band was effective when presented as a pulse in quiet (experiment 4). The lowest frequencies in the most effective 1-octave band were one octave above the spectral region where mice have their best absolute thresholds. These effects are similar to those obtained in humans, and reveal a special contribution of wide band, high-frequency stimulation to temporal acuity.     

A computer study of the relationship between noise rating assessment and dB(A) levels

The paper discusses the merits of using a single dB(A) reading in preference to the more complicated and time-consuming process of deriving an NR number. It is shown that, for commonly-encountered spectra, there is a small numerical difference between the dB(A) and NR values. This difference depends upon spectral shape and the overall level of the noise. Octave bands above 1 kHz tend to make the dB(A) level less than the NR number, while the octave bands below 1 kHz generally have the opposite effect.     

Toneburst-evoked auditory brainstem response in a leopard seal, Hydrurga leptonyx

Toneburst-evoked auditory brainstem responses (ABRs) were recorded in a captive subadult male leopard seal. Three frequencies from 1 to 4 kHz were tested at sound levels from 68 to 122 dB peak equivalent sound pressure level (peSPL). Results illustrate brainstem activity within the 1-4 kHz range, with better hearing sensitivity at 4 kHz. As is seen in human ABR, only wave V is reliably identified at the lower stimulus intensities. Wave V is present down to levels of 82 dB peSPL in the right ear and 92 dB peSPL in the left ear at 4 kHz. Further investigations testing a wider frequency range on seals of various sex and age classes are required to conclusively report on the hearing range and sensitivity in this species.     

Detection of high-frequency energy changes in sustained vowels produced by singers

The human voice spectrum above 5 kHz receives little attention. However, there are reasons to believe that this high-frequency energy (HFE) may play a role in perceived quality of voice in singing and speech. To fulfill this role, differences in HFE must first be detectable. To determine human ability to detect differences in HFE, the levels of the 8- and 16-kHz center-frequency octave bands were individually attenuated in sustained vowel sounds produced by singers and presented to listeners. Relatively small changes in HFE were in fact detectable, suggesting that this frequency range potentially contributes to the perception of especially the singing voice. Detection ability was greater in the 8-kHz octave than in the 16-kHz octave and varied with band energy level.     

DPOAE group delays versus electrophysiological measures of cochlear delay in normal human ears

Group delays of 2 f1-f2 distortion product otoacoustic emissions (DPOAEs) were determined using both f1- and f2-sweep paradigms in 24 normal-hearing subjects. These DPOAE group delays were studied in comparison with cochlear delays estimated from derived band VIIIth nerve compound action potentials (CAPs) and auditory brainstem responses (ABRs) in the same subjects. The center frequencies of the derived bands in the electrophysiological experiment were matched with the f2-frequencies in the DPOAE recording to ensure that DPOAEs and derived CAPs and ABRs were generated at the same places along the cochlear partition, thus allowing for a direct comparison. The degree to which DPOAE group delays are larger in the f2- than in the f1-sweep paradigm is consistent with a theoretical analysis of the so-called wave-fixed model. Both DPOAE group delays are highly correlated with CAP- and ABR-derived measures of cochlear delay. The principal result of this study is that "roundtrip" DPOAE group delay in the f1-sweep paradigm is exactly twice as large as the neural estimate of the "forward" cochlear delay. The interpretation of this notion in the context of cochlear wave propagation properties and DPOAE-generating mechanisms is discussed.     

Evidence for double acoustic windows in the dolphin, Tursiops truncatus

In a bottlenose dolphin positions of sound receiving areas on the head surface were determined by comparing the acoustic delays from different sound-source positions. For this investigation, auditory brainstem responses (ABRs) to short tone pips were recorded and their latencies were measured at different sound source positions. After correction for the latency dependence on response amplitude, the difference in ABR latencies was adopted as being the difference of the acoustic delays. These delay differences were used to calculate the position of the sound-receiving point. Measurements were conducted at sound frequencies from 16 to 128 kHz, in half-octave steps. At probe frequencies of 16 and 22.5 kHz, the receiving area was located 21.7-26 cm caudal of the melon tip, which is near the bulla and auditory meatus. At higher probe frequencies, from 32 to 128 kHz, the receiving area was located from 9.3 to 13.1 cm caudal of the melon tip, which corresponds to a proximal part of the lower jaw. Thus, at least two sound-receiving areas (acoustic windows) with different frequency sensitivity were identified.     

Auditory brainstem responses to chirps delivered by different insert earphones

The frequency response and sensitivity of the ER-3A and ER-2 insert earphones are measured in the occluded-ear simulator using three ear canal extensions. Compared to the other two extensions, the DB 0370 (Bru?el & Kj?r), which is recommended by the international standards, introduces a significant resonance peak around 4500 Hz. The ER-3A has an amplitude response like a band-pass filter (1400 Hz, 6 dB/octave -4000 Hz, -36 dB/octave), and a group delay with "ripples" of up to ±0.5 ms, while the ER-2 has an amplitude response, and a group delay which are flat and smooth up to above 10000 Hz. Both earphones are used to record auditory brainstem responses, ABRs, from 22 normal-hearing ears in response to two chirps and a click at levels from 20 to 80 dB nHL. While the click-ABRs are slightly larger for ER-2 than for ER-3A, the chirp-ABRs are much larger for ER-2 than for ER-3A at levels below 60 dB nHL. With a simulated amplitude response of the ER-3A and the smooth group delay of the ER-2 it is shown that the increased chirp-ABR amplitude with the ER-2 is caused by its broader amplitude response and not by its smoother group delay.     

Masked auditory thresholds in three species of birds, as measured by the auditory brainstem response (L)

Auditory brainstem responses (ABRs) were recorded in adult budgerigars, canaries, and zebra finches in quiet and in three levels of white noise for tone stimuli between 1 and 4 kHz. Similar to behavioral results, masked ABR thresholds increased linearly with increasing noise levels. When the three species are considered together, ABR-derived CRs were higher than behavioral CRs by 18-23 dB between 2 and 4 kHz and by about 30 dB at 1 kHz. This study clarifies the utility of using ABRs for estimating masked auditory thresholds in natural environmental noises in species that cannot be tested behaviorally.     

Place specificity of multiple auditory steady-state responses

Auditory steady-state responses (ASSRs) were elicited by simultaneously presenting multiple AM (amplitude-modulated) tones with carrier frequencies of 500, 1000, 2000, and 4000 Hz and modulation frequencies of 77, 85, 93, and 102 Hz, respectively. Responses were also evoked by separately presenting single 500- or 2000-Hz AM tones. The objectives of this study were (i) to determine the cochlear place specificity of single and multiple ASSRs using high-pass noise masking and derived-band responses, and (ii) to determine if there were any differences between single- and multiple-stimulus conditions. For all carrier frequencies, derived-band ASSRs for 1-octave-wide derived bands ranging in center frequency from 0.25 to 8 kHz had maximum amplitudes within a 1/2 octave of the carrier frequency. For simultaneously presented AM tones of 500, 1000, 2000, and 4000 Hz, bandwidths for the function of derived-band ASSR amplitude by derived-band center frequency were 476, 737, 1177, and 3039 Hz, respectively. There were no significant differences when compared to bandwidths of 486 and 1371 for ASSRs to AM tones of 500 or 2000 Hz presented separately. Results indicate that ASSRs to moderately intense stimuli (60 dB SPL) reflect activation of reasonably narrow cochlear regions, regardless of presenting AM tones simultaneously or separately.     

Audiogram of a striped dolphin (Stenella coeruleoalba)

The underwater hearing sensitivity of a striped dolphin was measured in a pool using standard psycho-acoustic techniques. The go/no-go response paradigm and up-down staircase psychometric method were used. Auditory sensitivity was measured by using 12 narrow-band frequency-modulated signals having center frequencies between 0.5 and 160 kHz. The 50% detection threshold was determined for each frequency. The resulting audiogram for this animal was U-shaped, with hearing capabilities from 0.5 to 160 kHz (8 1/3 oct). Maximum sensitivity (42 dB re 1 microPa) occurred at 64 kHz. The range of most sensitive hearing (defined as the frequency range with sensitivities within 10 dB of maximum sensitivity) was from 29 to 123 kHz (approximately 2 oct). The animal's hearing became less sensitive below 32 kHz and above 120 kHz. Sensitivity decreased by about 8 dB per octave below 1 kHz and fell sharply at a rate of about 390 dB per octave above 140 kHz.     

The development of middle-ear admittance in the hamster.

A high-frequency admittance meter was developed and used to study the maturation of physiological function in the middle ear of neonatal golden hamsters (Mesocricetus auratus). The middle-ear input admittance in the frequency range of 0.8-1.8 kHz was measured in animals ranging in age from 6 to 69 days postpartum. Admittance magnitude was found to increase steadily with age, beginning on day 16, to asymptotic values at each test frequency. There were no obvious differences in admittance growth rates within the range of frequencies tested. However, an analysis of the slopes of the admittance ma;nitude frequency response curves revealed an increase from 4.6 dB/octave for animals 25 days old or younger, to 6.3 dB/octave for all older animals. This difference between younger and older subjects indicates that the development of the middle ear in the golden hamster is more complex than a simple increase in pure compliance.     

The effect of the coupling between the top plate and the fingerboard on the acoustic power radiated by a classical guitar (L)

An experimental investigation on the coupling between the fingerboard and the top plate of a classical guitar at low frequencies is presented. The study was carried out using a finished top plate under fixed boundary conditions and a commercial guitar. Radiated sound power was determined in one-third octave bands up to the band of 1 kHz based on measurements of sound intensity. The results provide evidence that the way in which the fingerboard and top plate are coupled is not a relevant factor in the radiated acoustic power of the classical guitar in the studied frequency range.     

Medial efferent effects on auditory-nerve responses to tail-frequency tones. I. Rate reduction.

One way medial efferents are thought to inhibit responses of auditory-nerve fibers (ANFs) is by reducing the gain of the cochlear amplifier thereby reducing motion of the basilar membrane. If this is the only mechanism of medial efferent inhibition, then medial efferents would not be expected to inhibit responses where the cochlear amplifier has little effect, i.e., at sound frequencies in the tails of tuning curves. Inhibition at tail frequencies was tested for by obtaining randomized rate-level functions from cat ANFs with high characteristic frequencies (CF > or = 5 kHz), stimulated with tones two or more octaves below CF. It was found that electrical stimulation of medial efferents can indeed inhibit ANF responses to tail-frequency tones. The amplitude of efferent inhibition depended on both sound level (largest near to threshold) and frequency (largest two to three octaves below CF). On average, inhibition of high-CF ANFs responding to 1 kHz tones was around 5 dB. Although an efferent reduction of basilar-membrane motion cannot be ruled out as the mechanism producing the inhibition of ANF responses to tail frequency tones, it seems more likely that efferents produce this effect by changing the micromechanics of the cochlear partition.     

Low frequency sound measurement in vehicles

Sound pressure level measurements in cars travelling at motorway speeds have shown that, in many cases, the overall level is very high in relation to the dB(A) and octave band levels, suggesting that much of the sound energy is in the low frequency and infrasonic regions. A technique has been developed to extend accurate octave band measurements down to the octave centred on 2 Hz. The system uses a calibrated sound level meter feeding a frequency modulation tape-recorder to record noise below 64 Hz, and an octave band analysis system to analyse the resultant tape recordings. Typical results are presented for a number of vehicles and it is found that sound pressure levels as high as 120 dB can be found in the octave bands between 2 and 16 Hz.     

Relationships between arithmetic averages of sound pressure level calculated in octave bands and Zwicker’s loudness level

Previously it has been found through a series of psychoacoustical experiments that the arithmetic average of sound pressure level calculated in octave bands is a good estimator of loudness for various kinds of environmental noise. Remarkably, the arithmetic average of sound pressure level in octave bands from 63 Hz to 4 kHz, L_m,1/1(63-4k), strongly correlates with the loudness level specified in ISO 532B, LL(Z), as well as with loudness assessment. To investigate this relationship further, a numerical study has been carried out based on Zwicker’s loudness model. As a result, practical expressions to estimate the loudness levels of general environmental noises from the sound pressure levels in octave bands from 63 Hz or 125 Hz to 4 kHz are proposed.     

Simulations of tonotopically mapped speech processors for cochlear implant electrodes varying in insertion depth

It has been claimed that speech recognition with a cochlear implant is dependent on the frequency alignment of analysis bands in the speech processor with characteristic frequencies (CFs) at electrode locations. However, the most apical electrode location can often have a CF of 1 kHz or more. The use of filters aligned in frequency to relatively basal electrode arrays leads to the loss of lower frequency speech information. This study simulates a frequency-aligned speech processor and common array insertion depths to assess this significance of this loss. Noise-excited vocoders simulated processors driving eight electrodes 2 mm apart. Analysis filters always had center frequencies matching the CFs of the simulated stimulation sites. The simulated insertion depth of the most apical electrode was varied in 2-mm steps between 25 mm (CF 502 Hz) and 17 mm (CF 1851 Hz) from the cochlear base. Identification of consonants, vowels, and words in sentences all showed a significant decline between each of the three more basal simulated electrode configurations. Thus, if implant processors used analysis filters frequency-aligned to electrode CFs, patients whose most apical electrode is 19 mm (CF 1.3 kHz) or less from the cochlear base would suffer a significant loss of speech information.