Auditory Research publications

Objective: The objective of this study was to describe the auditory evoked response to silent gaps for a group of older adults using stimulus conditions identical to those used in psychophysical studies of gap detection. Design: The P1-N1-P2 response to the onsets of stimuli (markers) defining a silent gap for within-channel (spectrally identical markers) and across-channel (spectrally different markers) conditions was examined using four perceptually-equated gap durations. Study sample: A group of 24 older adults (mean age = 63 years) with normal hearing or minimal hearing loss participated. Results: Older adults exhibited neural activation patterns that were qualitatively different and more frontally oriented than those observed in a previous study (Lister et al., 2007) of younger listeners. Older adults showed longer P2 latencies and larger P1 amplitudes than younger adults, suggesting relatively slower neural travel time and altered auditory inhibition/arousal by irrelevant stimuli. Conclusion: Older adults appeared to recruit later-occurring T-complex-like generators for gap processing, compared to earlier-occurring T-complex-like generators by the younger group. Early and continued processing of channel cues with later processing of gap cues may represent the inefficiency of the aging auditory system and may contribute to poor speech understanding in noisy, real-world listening environments.

Elderly individuals often complain of difficulties in understanding speech, especially when heard against a background noise or when there are multiple speakers[...]

During speaking, auditory feedback is used to adjust vocalizations. The brain systems mediating this integrative ability have been investigated using a wide range of experimental strategies. In this report we examined how vocalization alters speech-sound processing within auditory cortex by directly recording evoked responses to vocalizations and playback stimuli using intracranial electrodes implanted in neurosurgery patients. Several new findings resulted from these high-resolution invasive recordings in human subjects. Suppressive effects of vocalization were found to occur only within circumscribed areas of auditory cortex. In addition, at a smaller number of sites, the opposite pattern was seen; cortical responses were enhanced during vocalization. This increase in activity was reflected in high gamma power changes, but was not evident in the averaged evoked potential waveforms. These new findings support forward models for vocal control in which efference copies of premotor cortex activity modulate sub-regions of auditory cortex.

The mammalian cochlea has two types of sensory cells; inner hair cells, which receive auditory-nerve afferent innervation, and outer hair cells, innervated by efferent axons of the medial olivocochlear (MOC) system. The role of the MOC system in hearing is still controversial. Recently, by recording cochlear potentials in behaving chinchillas, we suggested that one of the possible functions of the efferent system is to reduce cochlear sensitivity during attention to other sensory modalities (Delano et al. in J Neurosci 27:4146-4153, 2007). However, in spite of these compelling results, the physiological effects of electrical MOC activation on cochlear potentials have not been described in detail in chinchillas. The main objective of the present work was to describe these efferent effects in the chinchilla, comparing them with those in other species and in behavioral experiments. We activated the MOC efferent axons in chinchillas with sectioned middle-ear muscles by applying current pulses at the fourth-ventricle floor. Auditory-nerve compound action potentials (CAP) and cochlear microphonics (CM) were acquired in response to clicks and tones of several frequencies, using a round-window electrode. Electrical efferent stimulation produced CAP amplitude suppressions reaching up to 11 dB. They were higher for low to moderate sound levels. Additionally, CM amplitude increments were found, the largest (≤ 2.5 dB) for low intensity tones. CAP suppression was present at all stimulus frequencies, but was greatest for 2 kHz. CM increments were highest for low-frequency tones, and almost absent at high frequencies. We conclude that the effect obtained in chinchilla is similar to but smaller than that observed in cats, and that the effects seen in awake chinchillas, albeit different in magnitude, are consistent with the activation of efferent fibers.

To clarify speech-elicited response patterns within auditory-responsive cortex of the posterolateral superior temporal (PLST) gyrus, time-frequency analyses of event-related band power in the high gamma frequency range (75-175 Hz) were performed on the electrocorticograms recorded from high-density subdural grid electrodes in 8 patients undergoing evaluation for medically intractable epilepsy. Stimuli were 6 stop consonant-vowel (CV) syllables that varied in their consonant place of articulation (POA) and voice onset time (VOT). Initial augmentation was maximal over several centimeters of PLST, lasted about 400 ms, and was often followed by suppression and a local outward expansion of activation. Maximal gamma power overlapped either the Nα or Pβ deflections of the average evoked potential (AEP). Correlations were observed between the relative magnitudes of gamma band responses elicited by unvoiced stop CV syllables (/pa/, /ka/, /ta/) and their corresponding voiced stop CV syllables (/ba/, /ga/, /da/), as well as by the VOT of the stimuli. VOT was also represented in the temporal patterns of the AEP. These findings, obtained in the passive awake state, indicate that PLST discriminates acoustic features associated with POA and VOT and serve as a benchmark upon which task-related speech activity can be compared.

The brainstem auditory pathway is obligatory for all aural information. Brainstem auditory neurons must encode the level and timing of sounds, as well as their time-dependent spectral properties, the fine structure, and envelope, which are essential for sound discrimination. This study focused on envelope coding in the two cochlear nuclei of the barn owl, nucleus angularis (NA) and nucleus magnocellularis (NM). NA and NM receive input from bifurcating auditory nerve fibers and initiate processing pathways specialized in encoding interaural time (ITD) and level (ILD) differences, respectively. We found that NA neurons, although unable to accurately encode stimulus phase, lock more strongly to the stimulus envelope than NM units. The spectrotemporal receptive fields (STRFs) of NA neurons exhibit a pre-excitatory suppressive field. Using multilinear regression analysis and computational modeling, we show that this feature of STRFs can account for enhanced across-trial response reliability, by locking spikes to the stimulus envelope. Our findings indicate a dichotomy in envelope coding between the time and intensity processing pathways as early as at the level of the cochlear nuclei. This allows the ILD processing pathway to encode envelope information with greater fidelity than the ITD processing pathway. Furthermore, we demonstrate that the properties of the STRFs of the neurons can be quantitatively related to spike timing reliability.

Neural prosthetic interfaces based upon penetrating microelectrode devices have broadened our understanding of the brain and have shown promise for restoring neurological functions lost to disease, stroke, or injury. However, the eventual viability of such devices for use in the treatment of neurological dysfunction may be ultimately constrained by the intrinsic brittleness of silicon, the material most commonly used for manufacture of penetrating microelectrodes. This brittleness creates predisposition for catastrophic fracture, which may adversely affect the reliability and safety of such devices, due to potential for fragmentation within the brain. Herein, we report the development of titanium-based penetrating microelectrodes that seek to address this potential future limitation. Titanium provides advantage relative to silicon due to its superior fracture toughness, which affords potential for creation of robust devices that are resistant to catastrophic failure. Realization of these devices is enabled by recently developed techniques which provide opportunity for fabrication of high-aspect-ratio micromechanical structures in bulk titanium substrates. Details are presented regarding the design, fabrication, mechanical testing, in vitro functional characterization, and preliminary in vivo testing of devices intended for acute recording in rat auditory cortex and thalamus, both independently and simultaneously.

The perception of breathiness in vowels is cued by multiple acoustic cues, including changes in aspiration noise (AH) and the open quotient (OQ) [Klatt and Klatt, J. Acoust. Soc. Am. 87(2), 820–857 (1990)]. A loudness model can be used to determine the extent to which AH masks the harmonic components in voice. The resulting “partial loudness” (PL) and loudness of AH [“noise loudness” (NL)] have been shown to be good predictors of perceived breathiness [Shrivastav and Sapienza, J. Acoust. Soc. Am. 114(1), 2217–2224 (2003)]. The levels of AH and OQ were systematically manipulated for ten synthetic vowels. Perceptual judgments of breathiness were obtained and regression functions to predict breathiness from the ratio of NL to PL (η) were derived. Results show that breathiness can be modeled as a power function of η. The power parameter of this function appears to be affected by the fundamental frequency of the vowel. A second experiment was conducted to determine if the resulting power function could estimate breathiness in a different set of voices. The breathiness of these stimuli, both natural and synthetic, was determined in a listening test. The model estimates of breathiness were highly correlated with perceptual data but the absolute predicted values showed some discrepancies.

The study measured listener sensitivity to increments in the inter-onset intervals (IOIs) of successive 20-ms 4000-Hz tone bursts in isochronous sequences. The stimulus sequences contained two-six tone bursts, separated equally by silent intervals, with tonal IOIs ranging from 25 to 100 ms. Difference limens (DLs) for increments of the tonal IOIs were measured to assess listener sensitivity to changes of sequence rate. Comparative DLs were also measured for increments of a single interval located within six-tone isochronous sequences with different tone rates. Listeners included younger normal-hearing adults and two groups of older adults with and without high-frequency sensorineural hearing loss. The results, expressed as Weber fractions (DL/IOI), revealed that discrimination improved as the sequence tone rate decreased and the number of tonal components increased. Discrimination of a single sequence interval also improved as the number of sequence components increased from two to six but only for brief intervals and fast sequence rates. Discrimination performance of the older listeners with and without hearing loss was equivalent and significantly poorer than that of the younger listeners. The discrimination results are examined and discussed within the context of multiple-look mechanisms and possible age-related differences in the sensory coding of signal onsets.

Listeners with sensorineural hearing loss are poorer than listeners with normal hearing at understanding one talker in the presence of another. This deficit is more pronounced when competing talkers are spatially separated, implying a reduced “spatial benefit” in hearing-impaired listeners. This study tested the hypothesis that this deficit is due to increased masking specifically during the simultaneous portions of competing speech signals. Monosyllabic words were compressed to a uniform duration and concatenated to create target and masker sentences with three levels of temporal overlap: 0% (non-overlapping in time), 50% (partially overlapping), or 100% (completely overlapping). Listeners with hearing loss performed particularly poorly in the 100% overlap condition, consistent with the idea that simultaneous speech sounds are most problematic for these listeners. However, spatial release from masking was reduced in all overlap conditions, suggesting that increased masking during periods of temporal overlap is only one factor limiting spatial unmasking in hearing-impaired listeners.

Hypothesized circuitry enabling information storage can be tested by attempting to implant memory directly in the brain in the absence of normal experience. Previously, we found that tone paired with activation of the cholinergic nucleus basalis (NB) does induce behavioral memory that shares cardinal features with natural memory; it is associative, highly specific, rapidly formed, consolidates and shows intermediate retention. Here we determine if implanted memory also exhibits long-term consolidation and retention. Adult male rats were first tested for behavioral responses (disruption of ongoing respiration) to tones (1-15 kHz), yielding pre-training behavioral frequency generalization gradients. They next received 3 days of training with a conditioned stimulus (CS) tone (8.0 kHz, 70 dB, 2s) either paired (n=7) or unpaired (n=6) with moderate electrical stimulation of the nucleus basalis (∼ 65 μA, 100 Hz, 0.2s, co-terminating with CS offset). Testing for long-term retention was performed by obtaining post-training behavioral frequency generalization gradients 24h and 2 weeks after training. At 24h post-training, the Paired group exhibited specific associative behavioral memory, manifested by larger responses to the CS frequency band than the Unpaired group. This memory was retained 2 weeks post-training. Moreover, 2 weeks later, the specificity and magnitude of memory had become greater, indicating that the implanted memory had undergone consolidation. Overall, the results demonstrate the validity of NB-implanted memory for understanding natural memory and that activation of the cholinergic nucleus basalis is sufficient to form natural associative memory.

Background: Mammalian and avian auditory hair cells display tonotopic mapping of frequency along the length of the cochlea and basilar papilla. It is not known whether the auditory hair cells of fishes possess a similar tonotopic organization in the saccule, which is thought to be the primary auditory receptor in teleosts. To investigate this question, we determined the location of hair cell damage in the saccules of goldfish (Carassius auratus) following exposure to specific frequencies. Subjects were divided into six groups of six fish each (five treatment groups plus control). The treatment groups were each exposed to one of five tones: 100, 400, 800, 2000, and 4000 Hz at 176 dB re 1 μPa root mean squared (RMS) for 48 hours. The saccules of each fish were dissected and labeled with phalloidin in order to visualize hair cell bundles. The hair cell bundles were counted at 19 specific locations in each saccule to determine the extent and location of hair cell damage. In addition to quantification of anatomical injury, hearing tests (using auditory evoked potentials) were performed on each fish immediately following sound exposure. Threshold shifts were calculated by subtracting control thresholds from post-sound exposure thresholds. Results: All sound-exposed fish exhibited significant hair cell and hearing loss following sound exposure. The location of hair cell loss varied along the length of the saccule in a graded manner with the frequency of sound exposure, with lower and higher frequencies damaging the more caudal and rostral regions of the saccule, respectively. Similarly, fish exposed to lower frequency tones exhibited greater threshold shifts at lower frequencies, while high-frequency tone exposure led to hearing loss at higher frequencies. In general, both hair cell and hearing loss declined as a function of increasing frequency of exposure tone, and there was a significant linear relationship between hair cell loss and hearing loss. Conclusions: The pattern of hair cell loss as a function of exposure tone frequency and saccular rostral-caudal location is similar to the pattern of hearing loss as a function of exposure tone frequency and hearing threshold frequency. This data suggest that the frequency analysis ability of goldfish is at least partially driven by peripheral tonotopy in the saccule.

We herein investigated the role of the STAT signaling cascade in the production of pro-inflammatory cytokines and cisplatin ototoxicity. A significant hearing impairment caused by cisplatin injection was observed in Balb/c (wild type, WT) and STAT4(-/-), but not in STAT6(-/-) mice. Moreover, the expression levels of the protein and mRNA of pro-inflammatory cytokines, including TNF-α, IL-1β, and IL-6, were markedly increased in the serum and cochlea of WT and STAT4(-/-), but not STAT6(-/-) mice. Organotypic culture revealed that the shape of stereocilia bundles and arrays of sensory hair cell layers in the organ of Corti from STAT6(-/-) mice were intact after treatment with cisplatin, whereas those from WT and STAT4(-/-) mice were highly distorted and disarrayed after the treatment. Cisplatin induced the phosphorylation of STAT6 in HEI-OC1 auditory cells, and the knockdown of STAT6 by STAT6-specific siRNA significantly protected HEI-OC1 auditory cells from cisplatin-induced cell death and inhibited pro-inflammatory cytokine production. We further demonstrated that IL-4 and IL-13 induced by cisplatin modulated the phosphorylation of STAT6 by binding with IL-4 receptor alpha and IL-13Rα1. These findings suggest that STAT6 signaling plays a pivotal role in cisplatin-mediated pro-inflammatory cytokine production and ototoxicity.

Purpose In this study, the authors investigated the effects of age on the use of fundamental frequency differences (ΔF 0 ) in the perception of competing synthesized vowels in simulations of electroacoustic and cochlear-implant hearing. Method Twelve younger listeners with normal hearing and 13 older listeners with (near) normal hearing were evaluated in their use of ΔF 0 in the perception of competing synthesized vowels for 3 conditions: unprocessed synthesized vowels (UNP), envelope-vocoded synthesized vowels that simulated a cochlear implant (VOC), and synthesized vowels processed to simulate electroacoustic stimulation (EAS) hearing. Tasks included (a) multiplicity, which required listeners to identify whether a stimulus contained 1 or 2 sounds and (b) double-vowel identification, which required listeners to attach phonemic labels to the competing synthesized vowels. Results Multiplicity perception was facilitated by ΔF 0 in UNP and EAS but not in VOC, with no age-related deficits evident. Double-vowel identification was facilitated by ΔF 0 , with ΔF 0 benefit largest in UNP, reduced in EAS, and absent in VOC. Age adversely affected overall identification and ΔF 0 benefit on the double-vowel task. Conclusions Some but not all older listeners derived ΔF 0 benefit in EAS hearing. This variability may partly be due to how listeners are able to draw on higher-level processing resources in extracting and integrating cues in EAS hearing.

Many cochlear models assign zero longitudinal coupling in the cochlea. Although this is consistent with the transverse basilar membrane (BM) fibers, the cochlear partition contains cellular longitudinal coupling. In cochlear models, longitudinal coupling diminishes passive BM tuning; however, it has recently been employed in theories of active mechanics to enhance tuning. Our goal in this study was to probe passive longitudinal coupling by comparing BM responses in damaged cochleae with passive responses in normal cochleae. The cochleae of gerbils were damaged with intratympanic neomycin followed by a waiting period to ensure that all of the cells of the partition were missing or severely disrupted. We then measured BM motion and examined the cochleae histologically. In comparison with passive responses in normal cochleae, we observed a downward shift in characteristic frequency, an expected consequence of reduced stiffness from cellular damage. However, we did not observe enhanced passive tuning in the damaged cochleae, as would be expected if longitudinal coupling were substantially greater in the normal cochleae. Thus, we conclude that cell-based longitudinal coupling is not large enough to influence passive cochlear mechanics. This finding constrains theories of active mechanics.

Fluctuations in sound amplitude provide important cues to the identity of many sounds including speech. Of interest here was whether the ability to detect these fluctuations can be improved with practice, and if so whether this learning generalizes to untrained cases. To address these issues, normal-hearing adults (n = 9) were trained to detect sinusoidal amplitude modulation (SAM; 80-Hz rate, 3–4 kHz bandpass carrier) 720 trials/day for 6–7 days and were tested before and after training on related SAM-detection and SAM-rate-discrimination conditions. Controls (n = 9) only participated in the pre- and post-tests. The trained listeners improved more than the controls on the trained condition between the pre- and post-tests, but different subgroups of trained listeners required different amounts of practice to reach asymptotic performance, ranging from 1 (n = 6) to 4–6 (n = 3) sessions. This training-induced learning did not generalize to detection with two untrained carrier spectra (5 kHz low-pass and 0.5–1.5 kHz bandpass) or to rate discrimination with the trained rate and carrier spectrum, but there was some indication that it generalized to detection with two untrained rates (30 and 150 Hz). Thus, practice improved the ability to detect amplitude modulation, but the generalization of this learning to untrained cases was somewhat limited.

Some individuals complain of hearing difficulties in the presence of background noise even in the absence of clinically significant hearing loss (obscure auditory dysfunction). Previous evidence suggests that these listeners have impaired frequency resolution, but there has been no thorough characterization of auditory filter shapes in this population. Here, the filter shapes of adults (n = 14) who self-reported speech recognition problems in noise and performed poorly on a sentence-in-noise perception test despite having clinically normal audiograms were compared to those of controls (n = 10). The filter shapes were evaluated using a 2-kHz probe with a fixed level of 30, 40, or 50 dB sound pressure level (SPL) and notched-noise simultaneous maskers that were varied in level to determine the masker level necessary to just mask the probe. The filters of the impaired group were significantly wider than those of controls at all probe levels owing to an unusual broadening of the upper slope of the filter. In addition, absolute thresholds were statistically indistinguishable between the groups at the standard audiometric frequencies, but were elevated in the impaired listeners at higher frequencies. These results strengthen the idea that this population has a variety of hearing deficits that go undetected by standard audiometry.

Objectives: Recent research suggests that older listeners may have difficulty processing information related to the fundamental frequency (F0) of voiced speech. In this study, the focus was on the mechanisms that may underlie this reduced ability. We examined whether increased age resulted in decreased ability to perceive F0 using fine-structure cues provided by the harmonic structure of voiced speech sounds or cues provided by high-rate envelope fluctuations (periodicity). Design: Younger listeners with normal hearing and older listeners with normal to near-normal hearing completed two tasks of F0 perception. In the first task (steady state F0), the fundamental frequency difference limen (F0DL) was measured adaptively for synthetic vowel stimuli. In the second task (time-varying F0), listeners relied on variations in F0 to judge intonation of synthetic diphthongs. For both tasks, three processing conditions were created: eight-channel vocoding that preserved periodicity cues to F0; a simulated electroacoustic stimulation condition, which consisted of high-frequency vocoder processing combined with a low-pass-filtered portion, and offered both periodicity and fine-structure cues to F0; and an unprocessed condition. Results: F0 difference limens for steady state vowel sounds and the ability to discern rising and falling intonations were significantly worse in the older subjects compared with the younger subjects. For both older and younger listeners, scores were lowest for the vocoded condition, and there was no difference in scores between the unprocessed and electroacoustic simulation conditions. Conclusions: Older listeners had difficulty using periodicity cues to obtain information related to talker fundamental frequency. However, performance was improved by combining periodicity cues with (low frequency) acoustic information, and that strategy should be considered in individuals who are appropriate candidates for such processing. For cochlear implant candidates, this effect might be achieved by partial electrode insertion providing acoustic stimulation in the low frequencies or by the combination of a traditional implant in one ear and a hearing aid in the opposite ear.

The speech signal may be divided into frequency bands, each containing temporal properties of the envelope and fine structure. For maximal speech understanding, listeners must allocate their perceptual resources to the most informative acoustic properties. Understanding this perceptual weighting is essential for the design of assistive listening devices that need to preserve these important speech cues. This study measured the perceptual weighting of young normal-hearing listeners for the envelope and fine structure in each of three frequency bands for sentence materials. Perceptual weights were obtained under two listening contexts: (1) when each acoustic property was presented individually and (2) when multiple acoustic properties were available concurrently. The processing method was designed to vary the availability of each acoustic property independently by adding noise at different levels. Perceptual weights were determined by correlating a listener’s performance with the availability of each acoustic property on a trial-by-trial basis. Results demonstrated that weights were (1) equal when acoustic properties were presented individually and (2) biased toward envelope and mid-frequency information when multiple properties were available. Results suggest a complex interaction between the available acoustic properties and the listening context in determining how best to allocate perceptual resources when listening to speech in noise.

Low-frequency masking by intense high-frequency noise bands, referred to as remote masking (RM), was the first evidence to challenge energy-detection models of signal detection. Its underlying mechanisms remain unknown. RM was measured in five normal-hearing young-adults at 250, 350, 500, and 700 Hz using equal-power, spectrally matched random-phase noise (RPN) and low-noise noise (LNN) narrowband maskers. RM was also measured using equal-power, two-tone complex (TC2) and eight-tone complex (TC8). Maskers were centered at 3000 Hz with one or two equivalent rectangular bandwidths (ERBs). Masker levels varied from 80 to 95 dB sound pressure level in 5 dB steps. LNN produced negligible masking for all conditions. An increase in bandwidth in RPN yielded greater masking over a wider frequency region. Masking for TC2 was limited to 350 and 700 Hz for one ERB but shifted to only 700 Hz for two ERBs. A spread of masking to 500 and 700 Hz was observed for TC8 when the bandwidth was increased from one to two ERBs. Results suggest that high-frequency noise bands at high levels could generate significant low-frequency masking. It is possible that listeners experience significant RM due to the amplification of various competing noises that might have significant implications for speech perception in noise.