Tone Pips Research Articles

Hearing threshold measurements of five stranded short-finned pilot whales (Globicephala macrorhynchus)

On May 5, 2011, 23 short-finned pilot whales, Globicephala macrorhynchus, were stranded along the coastline near Cudjoe Key, FL. Five animals (two adult females, two juvenile females, and an adult male) were transported to a rehabilitation facility in Key Largo, FL. Auditory evoked potentials (AEPs) were recorded in response to amplitude modulated tone pips modulated at 1000 Hz. AEP thresholds were determined at 10, 20, 40, 80, and 120 kHz for the four females. However, the adult male was euthanized prior to testing. Short-finned pilot whales had peak sensitivity at lower frequencies than other odontocetes such as bottlenose dolphins. Greatest sensitivity was around 40 kHz for all whales, while thresholds for the two adult females were 25-61 dB higher at 80 kHz than the juveniles. Click evoked potentials were similar between the four whales and comparable to other echolocating odontocetes. Click evoked potential data from a fifth short-finned pilot whale that had stranded in Curacao showed no response. These findings add to the limited database of pilot whale (short- and long-finned) hearing studies, of which there are only two others [Schlundt et al. (2011). J. Acoust. Soc. Am. 129, 1111-1116 and Pacini et al. (2010). J. Exp. Biol. 213, 3138-3143].

Provisional stimulus level corrections for low frequency bone-conduction ABR in babies under three months corrected age

Objective: To estimate bone-conduction stimulus level corrections by testing the auditory brainstem response (ABR) of normally-hearing newborns. The stimuli used were low frequency tone pips calibrated to reference levels derived from ISO 389 values. Design: Tone pips were presented via supra-aural earphones and a B71 Radioear bone vibrator at 0.5 or 1 kHz. ABR thresholds from both transducers were compared at each frequency. Study sample: twenty-seven newborn hearing screening referrals (33 ears) who passed an ABR discharge criterion at 4 kHz. Results: Median air- and bone-conduction ABR threshold differences were 30 dB at 0.5 kHz and 20 dB at 1 kHz. Conclusion: The 0.5 kHz data from this study and previous studies were compared. Previous studies suggested lower figures for the bone-conduction stimulus level correction. Likely sources of this discrepancy are discussed. The average 0.5 kHz bone-conduction correction value for infants < 3 months old is about 28 dB. The correction for 1 kHz is 20 dB. We recommend that calibration reference levels used in this study be adopted and that appropriate corrections be applied to bone conduction ABR thresholds in infants < 3 months old before calculation of any air-bone gap and subsequent clinical interpretation.

Vestibular receptors contribute to cortical auditory evoked potentials

Acoustic sensitivity of the vestibular apparatus is well-established, but the contribution of vestibular receptors to the late auditory evoked potentials of cortical origin is unknown. Evoked potentials from 500 Hz tone pips were recorded using 70 channel EEG at several intensities below and above the vestibular acoustic threshold, as determined by vestibular evoked myogenic potentials (VEMPs). In healthy subjects both auditory mid- and long-latency auditory evoked potentials (AEPs), consisting of Na, Pa, N1 and P2 waves, were observed in the sub-threshold conditions. However, in passing through the vestibular threshold, systematic changes were observed in the morphology of the potentials and in the intensity dependence of their amplitude and latency. These changes were absent in a patient without functioning vestibular receptors. In particular, for the healthy subjects there was a fronto-central negativity, which appeared at about 42 ms, referred to as an N42, prior to the AEP N1. Source analysis of both the N42 and N1 indicated involvement of cingulate cortex, as well as bilateral superior temporal cortex. Our findings are best explained by vestibular receptors contributing to what were hitherto considered as purely auditory evoked potentials and in addition tentatively identify a new component that appears to be primarily of vestibular origin.

Comparison of chirp versus click and tone pip stimulation for cervical vestibular evoked myogenic potentials

The current study explored differences among cervical vestibular evoked myogenic potentials (cVEMP) that were evoked by CE-chirp and click and tone pip in healthy controls, and tried to explain the differences of cVEMP between the three of them. Thirty normal volunteers were used as subjects for CE-chirp and click and tone-pip (Blackman pip) stimuli. The latency of P1, N1, peak-to-peak P1-N1 amplitude, and cVEMP interaural difference were obtained and analyzed. The response rates of cVEMP were 93 % for click and 100 % for both Blackman pip and CE-chirp, respectively. The P1 and N1 latencies of cVEMP evoked by CE-chirp were the shortest, followed by click, with Blackman pip the longest (F = 6,686.852, P < 0.001). All indices of cVEMP evoked by the three stimuli showed no significant difference between the left and right ears or between genders. cVEMP responses were significantly different between the three stimuli. Compared with the currently used stimulus, CE-chirp can evoke cVEMP with shorter latencies and demonstrates increased speed and reliability.

The effect of parvalbumin deficiency on the acoustic startle response and prepulse inhibition in mice

The strength of the acoustic startle response (ASR) to short bursts of broadband noise or tone pips (4, 8 and 16kHz) and the prepulse inhibition (PPI) of the ASR elicited by prepulse tones (4, 8 and 16kHz) were measured in parvalbumin-deficient (PV−/−) mice and in age-matched PV+/+ mice as controls. Hearing thresholds as determined from recordings of auditory brainstem responses were found to be similar in both genotypes. The ASRs to broadband noise and tones of low and middle frequencies were stronger than the ASRs in response to high-frequency tones in both groups. In PV−/− mice, we observed smaller ASR amplitudes in response to relatively weak startling stimuli (80–90dB sound pressure level (SPL)) of either broadband noise or 8-kHz tones compared to those recorded in PV+/+ mice. For these startling stimuli, PV−/− mice had higher ASR thresholds and longer ASR latencies. PPI of the ASR in PV−/− mice was less effective than in PV+/+ mice, for all tested prepulse frequencies (4, 8 or 16kHz) at 70dB SPL. Our findings demonstrate no effect of PV deficiency on hearing thresholds in PV−/− mice. However, the frequency-specific differences in the ASR and the significant reduction of PPI of ASR likely reflect specific changes of neuronal circuits, mainly inhibitory, in the auditory centers in PV-deficient mice.

Level dominance for the detection of changes in level distribution in sound streams

Sound streams were generated by randomly choosing the levels of tone pips from two different distributions, A and B. Of the 18 tone pips, the first nine were drawn from distribution A and the second nine from distribution B, or the opposite. The listeners' task was to indicate order, A-B or B-A. In two conditions the A and B distributions differed in mean (condition 1) or variance (condition 2). In contrast to an ideal observer, listeners' strategies were consistent across the two conditions. Analyses suggest that listeners relied primarily on the more intense tone pips in making their decisions.

Erratum to: Conductive Hearing Loss Induced by Experimental Middle-Ear Effusion in a Chinchilla Model Reveals Impaired Tympanic Membrane-Coupled Ossicular Chain Movement

Otitis media with effusion (OME) occurs when fluid collects in the middle-ear space behind the tympanic membrane (TM). As a result of this effusion, sounds can become attenuated by as much as 30-40 dB, causing a conductive hearing loss (CHL). However, the exact mechanical cause of the hearing loss remains unclear. Possible causes can include altered compliance of the TM, inefficient movement of the ossicular chain, decreased compliance of the oval window-stapes footplate complex, or altered input to the oval and round window due to conduction of sound energy through middle-ear fluid. Here, we studied the contribution of TM motion and umbo velocity to a CHL caused by middle-ear effusion. Using the chinchilla as an animal model, umbo velocity (V U) and cochlear microphonic (CM) responses were measured simultaneously using sinusoidal tone pip stimuli (125 Hz-12 kHz) before and after filling the middle ear with different volumes (0.5-2.0 mL) of silicone oil (viscosity, 3.5 Poise). Concurrent increases in CM thresholds and decreases in umbo velocity were noted after the middle ear was filled with 1.0 mL or more of fluid. Across animals, completely filling the middle ear with fluid caused 20-40-dB increases in CM thresholds and 15-35-dB attenuations in umbo velocity. Clinic-standard 226-Hz tympanometry was insensitive to fluid-associated changes in CM thresholds until virtually the entire middle-ear cavity had been filled (approximately >1.5 mL). The changes in umbo velocity, CM thresholds, and tympanometry due to experimentally induced OME suggest CHL arises primarily as a result of impaired TM mobility and TM-coupled umbo motion plus additional mechanisms within the middle ear.

Intra and intersubject variability in auditory steady-state response amplitude with high modulation rates to 1000 Hz amplitude modulated and tone pip stimuli

Objective: This study set out to provide further information on how high modulation/stimulus rates affect the auditory steady-state response (ASSR) amplitude for a 1000-Hz tone, and how this effect varies between individuals. Both sinusoidal amplitude modulated tones and tone pip stimuli were investigated. Design: Modulation/stimulus rates were 70, 80, and 90 Hz and the peak to peak stimulus levels were matched for the two different types of stimuli, at 90.5 dBSPLppe. Study sample: The study was carried out on fourteen normally-hearing adults (9 males and 5 females) Results: Overall the ASSR amplitude to the two types of stimuli was similar. In general there was an increasing response amplitude between rates of 70 and 90 Hz; The relationship between the amplitude of the response and the modulation /stimulus rate varied considerably between subjects. Conclusions: Optimum stimulus rates based on group data may not give the best rate in a significant proportion of subjects. Currently tone pip ABR is the primary method used in assessing hearing in babies. Finding a way of avoiding suboptimal stimulus rates for ‘80-Hz’ ASSR in babies will improve the likelihood of ASSR being seen as an alternative.

Spontaneous firing rate changes in cat primary auditory cortex following long-term exposure to non-traumatic noise: Tinnitus without hearing loss?

Changes of neural activity in animal models have been correlated with tinnitus in humans. For instance, increased spontaneous firing rates (SFR), increased spontaneous neural synchrony, and cortical tonotopic map reorganization may underlie this phantom auditory percept. The aim of this study is to quantify the changes in SFR activity in the cat primary auditory cortex, after long-term exposure to different types of non-traumatic acoustic environments. For that purpose, four different groups of adult cats were exposed to moderate-level (~70dB SPL), behaviorally irrelevant sounds for several weeks to months, and their SFRs were compared with those in control cats. The sounds consisted of random multi-frequency tone pip ensembles with various bandwidths (2–4kHz, 4–20kHz, and a pair of third-octave bands centered at 4 and 16kHz), as well as a “factory noise”. Auditory brainstem response (ABR) thresholds, ABR wave 3 amplitudes at ~55 and 75dB SPL, and distortion product otoacoustic emission (DPOAE) amplitudes were unaffected by the exposure. However, we found that the SFR decreased within the exposure frequency range and increased outside the exposure range. This increased SFR for units with characteristic frequencies outside the exposure frequency range, which was slow to reverse after the exposure offset, suggests a mechanism for tinnitus in the absence of hearing loss.

Predictive value of hearing assessment by the auditory brainstem response following universal newborn hearing screening

Objective: This study set out to determine the accuracy with which tone pip ABR and click ABR, carried out in babies referred from universal newborn hearing screening, is able to predict the hearing outcome as determined by follow-up hearing tests. Study sample: The cohort of babies studied were all babies referred for hearing assessment from the universal newborn hearing screen in Sheffield, UK for the period January 2002 to September 2007, who were found to have a significant hearing impairment. Design: The results of hearing assessment following referral from the newborn hearing screen were collected together with those of follow-up tests carried out up to an age when behavioural testing had established ear- and frequency-specific thresholds at 0.5, 1, 2 and 4 kHz. Results: The standard deviation of the difference between the follow up and the tone pip ABR thresholds was 10.5 dB for the 4-kHz tone pip, 16.8 dB for the 1-kHz tone pip, and ranged between 21.7 and 24.7 dB for click ABR. Conclusions: The results of the study show that tone pip ABR following referral from newborn hearing screening has a similar accuracy to that reported in older subjects, and is a much better predictor compared to click ABR.

Conductive Hearing Loss Induced by Experimental Middle-Ear Effusion in a Chinchilla Model Reveals Impaired Tympanic Membrane-Coupled Ossicular Chain Movement

Otitis media with effusion (OME) occurs when fluid collects in the middle-ear space behind the tympanic membrane (TM). As a result of this effusion, sounds can become attenuated by as much as 30-40dB, causing a conductive hearing loss (CHL). However, the exact mechanical cause of the hearing loss remains unclear. Possible causes can include altered compliance of the TM, inefficient movement of the ossicular chain, decreased compliance of the oval window-stapes footplate complex, or altered input to the oval and round window due to conduction of sound energy through middle-ear fluid. Here, we studied the contribution of TM motion and umbo velocity to a CHL caused by middle-ear effusion. Using the chinchilla as an animal model, umbo velocity (V U) and cochlear microphonic (CM) responses were measured simultaneously using sinusoidal tone pip stimuli (125Hz-12kHz) before and after filling the middle ear with different volumes (0.5-2.0mL) of silicone oil (viscosity, 3.5 Poise). Concurrent increases in CM thresholds and decreases in umbo velocity were noted after the middle ear was filled with 1.0mL or more of fluid. Across animals, completely filling the middle ear with fluid caused 20-40-dB increases in CM thresholds and 15-35-dB attenuations in umbo velocity. Clinic-standard 226-Hz tympanometry was insensitive to fluid-associated changes in CM thresholds until virtually the entire middle-ear cavity had been filled (approximately >1.5mL). The changes in umbo velocity, CM thresholds, and tympanometry due to experimentally induced OME suggest CHL arises primarily as a result of impaired TM mobility and TM-coupled umbo motion plus additional mechanisms within the middle ear.

Comparison of ABR response amplitude, test time, and estimation of hearing threshold using frequency specific chirp and tone pip stimuli in newborns

Objective: To evaluate the auditory brainstem response (ABR) amplitudes evoked by tone pip and narrowband chirp (NB CE-Chirp) stimuli when testing post-screening newborns and to determine the difference in estimated hearing level correction values. Design: Tests were performed with tone pips and NB CE-Chirps at 4 kHz or 1 kHz. The response amplitude, response quality (Fmp), and residual noise were compared for both stimuli. Study sample: Thirty babies (42 ears) who passed our ABR discharge criterion at 4 kHz following referral from their newborn hearing screen. Results: Overall, NB CE-Chirp responses were 64% larger than the tone pip responses, closer to those evoked by clicks. Fmp was significantly higher for NB CE-Chirps. Conclusion: It is anticipated that there could be significant reductions in test time for the same signal to noise ratio by using NB CE-Chirps when testing newborns. This effect may vary in practice and is likely to be most beneficial for babies with low amplitude ABR responses. We propose that the ABR nHL threshold to eHL correction for NB CE-Chirps should be approximately 5 dB less than the corrections for tone pips at 4 and 1 kHz.

Psychoacoustic and Phoneme Identification Measures in Cochlear-Implant and Normal-Hearing Listeners

The purpose of this study is to identify precise and repeatable measures for assessing cochlear-implant (CI) hearing. The study presents psychoacoustic and phoneme identification measures in CI and normal-hearing (NH) listeners, with correlations between measures examined. Psychoacoustic measures included pitch discrimination tasks using pure tones, harmonic complexes, and tone pips; intensity perception tasks included intensity discrimination for tones and modulation detection; spectral-temporal masking tasks included gap detection, forward and backward masking, tone-on-tone masking, synthetic formant-on-formant masking, and tone in noise detection. Phoneme perception measures included vowel and consonant identification in quiet and stationary and temporally gated speech-shaped noise. Results on psychoacoustic measures illustrate the effects of broader filtering in CI hearing contributing to reduced pitch perception and increased spectral masking. Results on consonant and vowel identification measures illustrate a wide range in performance across CI listeners. They also provide further evidence that CI listeners obtain little to no release of masking in temporally gated noise compared to stationary noise. The forward and backward-masking measures had the highest correlation with the phoneme identification measures for CI listeners. No significant correlations between speech reception and psychoacoustic measures were observed for NH listeners. The superior NH performance on measures of phoneme identification, especially in the presence of background noise, is a key difference between groups.

Repeated exposure to a tone transiently alters spectral tuning bandwidth of neurons in the central nucleus of inferior colliculus in juvenile rats

Early acoustic experience changes tonal frequency tuning in the inferior colliculus (IC) and the primary auditory cortex. The contributions of IC plasticity to cortical frequency map reorganization are not entirely clear. While most cortical plasticity studies exposed animals to pulsed tones, studies of IC plasticity used either noise or a continuous tone. Here we compared the effects of repeated exposure to single-frequency tone pips on cortical and IC frequency representations in juvenile rats. We found that while tone exposure caused a long-lasting increase in cortical representations of the exposure frequency, changes to IC neurons were limited to a transient narrowing of tuning bandwidth. These results suggest that previously documented cortical frequency map reorganization does not depend on similar changes in the subcortical auditory nuclei.

Effects of passive, moderate-level sound exposure on the mature auditory cortex: Spectral edges, spectrotemporal density, and real-world noise

Persistent, passive exposure of adult cats to bandlimited tone pip ensembles or sharply-filtered white noise at moderate levels (∼70 dB SPL) leads to a long-term suppression of spontaneous and sound-evoked activity in the region(s) of primary auditory cortex (AI) normally tuned to the exposure spectrum, and to an enhancement of activity in one or more neighboring regions of AI, all in the apparent absence of hearing loss. Here, we first examined the effects of passive exposure to a more structured, real-world noise, consisting of a mix of power tool and construction sounds. This “factory noise” had less pronounced effects on adult cat AI than our previous random tone pip ensembles and white noise, and these effects appeared limited to the region of AI tuned to frequencies near the sharp factory noise cutoff at 16 kHz. To further investigate the role of sharp spectral edges in passive exposure-induced cortical plasticity, a second group of adult cats was exposed to a tone pip ensemble with a flat spectrum between 2 and 4 kHz and shallow cutoff slopes (12 dB/oct) on either side. Compared to our previous ensemble with the same power in the 2–4 kHz band but very steep slopes, exposure to the overall more intense, sloped stimulus had much weaker effects on AI. Finally, we explored the issue of exposure stimulus spectrotemporal density and found that low aggregate tone pip presentation rates of about one per second sufficed to induce changes in the adult AI similar to those characteristic of our previous, much denser exposures. These results are discussed in light of the putative mechanisms underlying exposure-induced auditory cortical plasticity, and the potential adverse consequences of working or living in moderately noisy environments.

Investigating the auditory enhancement phenomenon using behavioral temporal masking patterns.

A narrowband signal is subjected to less masking from a simultaneously presented notched masker if it is preceded by a precursor that occupies the same spectral region as the masker, a phenomenon referred to as enhancement. The present study investigated (i) the amount of enhancement for the detection of a narrowband noise added to a notched masker, and (ii) masking patterns associated with the detection of tone pips added to the narrowband signal. The resulting psychophysical data were compared to predictions generated using a model similar to the neural adaptation-of-inhibition model proposed by Nelson and Young [(2010b). J. Neurosci. 30, 6577-6587]. The amount of enhancement was measured as a function of the temporal separation between the precursor and masker in Experiment I, and as a function of precursor level in Experiment II. The model captured the temporal dynamics of psychophysical enhancement reasonably well for both the long-duration noise signals and the masking patterns. However, in contrast to psychophysical data which indicated reliable enhancement only when the precursor and masker shared the same levels, the model predicated enhancement at all precursor levels.

Plasticity of Peripheral Auditory Frequency Sensitivity in Emei Music Frog

In anurans reproductive behavior is strongly seasonal. During the spring, frogs emerge from hibernation and males vocalize for mating or advertising territories. Female frogs have the ability to evaluate the quality of the males' resources on the basis of these vocalizations. Although studies revealed that central single torus semicircularis neurons in frogs exhibit season plasticity, the plasticity of peripheral auditory sensitivity in frog is unknown. In this study the seasonally plasticity of peripheral auditory sensitivity was test in the Emei music frog Babina daunchina, by comparing thresholds and latencies of auditory brainstem responses (ABRs) evoked by tone pips and clicks in the reproductive and non-reproductive seasons. The results show that both ABR thresholds and latency differ significantly between the reproductive and non-reproductive seasons. The thresholds of tone pip evoked ABRs in the non-reproductive season increased significantly about 10 dB than those in the reproductive season for frequencies from 1 KHz to 6 KHz. ABR latencies to waveform valley values for tone pips for the same frequencies using appropriate threshold stimulus levels are longer than those in the reproductive season for frequencies from 1.5 to 6 KHz range, although from 0.2 to 1.5 KHz range it is shorter in the non-reproductive season. These results demonstrated that peripheral auditory frequency sensitivity exhibits seasonal plasticity changes which may be adaptive to seasonal reproductive behavior in frogs.

Detection of low‐frequency tones and whale predator sounds by the American sand lance Ammodytes americanus

Auditory evoked potentials (AEP) were used to measure the hearing range and auditory sensitivity of the American sand lance Ammodytes americanus. Responses to amplitude-modulated tone pips indicated that the hearing range extended from 50 to 400 Hz. Sound pressure thresholds were lowest between 200 and 400 Hz. Particle acceleration thresholds showed an improved sensitivity notch at 200 Hz but not substantial differences between frequencies and only a slight improvement in hearing abilities at lower frequencies. The hearing range was similar to Pacific sand lance Ammodytes personatus and variations between species may be due to differences in threshold evaluation methods. AEPs were also recorded in response to pulsed sounds simulating humpback whale Megaptera novaeangliae foraging vocalizations termed megapclicks. Responses were generated with pulses containing significant energy below 400 Hz. No responses were recorded using pulses with peak energy above 400 Hz. These results show that A. americanus can detect the particle motion component of low-frequency tones and pulse sounds, including those similar to the low-frequency components of megapclicks. Ammodytes americanus hearing may be used to detect environmental cues and the pulsed signals of mysticete predators.

Low‐Cost System for Auditory Brainstem Evoked Response

Objective: 1) Be able to design-low-cost ABR equipment. 2) Learn the possibility of mainstream technology applied for laboratory equipment use and be able to standardize the results with commercially available ABR equipment. 3) Apply this technology for research and neurophysiology laboratories. Method: Animal experimentation study. This system comprises a differential amplifier, high-pass and low-pass filters, a main rejection notch filter, an analog to digital converter, and software programs for data acquisition and processing automation. Amplifier and filters were designed using low-cost and readily available commercial components. Auditory stimuli were reproduced by a portable music player and a miniature rechargeable speaker. The stimuli comprising auditory clicks, tone pips and tone bursts, were generated numerically by the software and transferred digitally to the music player. 25 female and male Sprague-Dawley rats were tested with amplitude and threshold protocols. Results: Results demonstrate a linear correlation and morphology with ABR previously recorded by other teams. Amplitude in the response was little affected by the hardware but strongly modified by electrode position. A regression analysis demonstrated a coincidence r = 0.92 with the physiological measured threshold using RMS analysis. Ketamine affects the first 20 minutes of recordings by delaying latency in all waves, the effect fades away after that. Morphological changes were observed by blocking the ear canal in accordance with previous studies. Conclusion: The ABRs obtained compare favorably with other published results in similar conditions. Clear amplification with subdermal electrodes are seen consistently and very little EEG noise is obtain with the recordings, with no significant difference between individuals and results from other studies. The system will rapid determine auditory threshold with good accuracy. We are confident that this approach will make hardware more accessible for low budget research facilities.

Reversible Long-Term Changes in Auditory Processing in Mature Auditory Cortex in the Absence of Hearing Loss Induced by Passive, Moderate-Level Sound Exposure

It has become increasingly clear that even occasional exposure to loud sounds in occupational or recreational settings can cause irreversible damage to the hair cells of the cochlea and the auditory nerve fibers, even if the resulting partial loss of hearing sensitivity, usually accompanied by tinnitus, disappears within hours or days of the exposure. Such exposure may explain at least some cases of poor speech intelligibility in noise in the face of a normal or near-normal audiogram. Recent findings from our laboratory suggest that long-term changes to auditory brain function-potentially leading to problems with speech intelligibility-can be effected by persistent, passive exposure to more moderate levels of noise (in the 70 dB SPL range) in the apparent absence of damage to the auditory periphery (as reflected in normal distortion product otoacoustic emissions and auditory brainstem responses). Specifically, passive exposure of adult cats to moderate levels of band-pass-filtered noise, or to band-limited ensembles of dense, random tone pips, can lead to a profound decrease of neural activity in the auditory cortex roughly in the exposure frequency range, and to an increase of activity outside that range. This can progress to an apparent reorganization of the cortical tonotopic map, which is reminiscent of the reorganization resulting from hearing loss restricted to a part of the hearing frequency range, although again, no hearing loss was apparent after our moderate-level sound exposure. Here, we review this work focusing specifically on the potential hearing problems that may arise despite a normally functioning auditory periphery.