Psychoacoustics publications

Noise-exposed rat pups provide a model of early deprivation of sensory input to the central auditory system, allowing the study of developmental neuroplasticity. Our previous results have demonstrated that a brief exposure of rats to broadband noise (125 dB SPL 8 min, 14th postnatal day) at the onset of hearing resulted in an altered intensity perception and frequency discrimination in adulthood despite normal hearing thresholds. In this study, we assessed the gap-detection ability and possible presence of tinnitus- and hyperacusis-like behavior in adult rats after the same neonatal acoustic trauma, using measurements of the acoustic startle response (ASR) in quiet and noisy environments and its prepulse inhibition by gaps in noise (gap-PPI). A significant deficit in the ability to detect gap was observed in the exposed rats when 55 dB SPL broadband noise was used as background. An increase of noise intensity to 65-75 dB SPL led to strengthening of the gap-PPI in exposed animals, which approached the gap-PPI values of control animals at these levels. Behavioral signs of tinnitus (gap detection deficits in 10 kHz narrow band noise) were found in 25% of exposed rats. An increased sensitivity to continuous noise was manifested in all exposed rats by suppression of the ASR at significantly lower background noise levels than in the controls. This effect was particularly pronounced in rats with tinnitus-like behavior. Our results indicate that neonatal acoustic trauma, producing only a transient threshold shift, may produce permanent abnormalities in suprathreshold auditory functions and the development of tinnitus and hyperacusis-like behavior.

In order to survive and function in the world, we must understand the content of our environment. This requires us to gather and parse complex, sometimes conflicting, information. Yet, the brain is capable of translating sensory stimuli from disparate modalities into a cohesive and accurate percept with little conscious effort. Previous studies of multisensory integration have suggested that the brain's integration of cues is well-approximated by an ideal observer implementing Bayesian causal inference. However, behavioral data from tasks that include only one stimulus in each modality fail to capture what is in nature a complex process. Here we employed an auditory spatial discrimination task in which listeners were asked to determine on which side they heard one of two concurrently presented sounds. We compared two visual conditions in which task-uninformative shapes were presented in the center of the screen, or spatially aligned with the auditory stimuli. We found that performance on the auditory task improved when the visual stimuli were spatially aligned with the auditory stimuli-even though the shapes provided no information about which side the auditory target was on. We also demonstrate that a model of a Bayesian ideal observer performing causal inference cannot explain this improvement, demonstrating that humans deviate systematically from the ideal observer model.

It is currently unclear whether mice use their ultrasonic vocalizations (USVs) for communication purposes. It is also unknown whether mice require previous experience with USVs to understand conspecifics. There is some evidence that experience changes the perception of juvenile USVs; however, it is unclear whether similar plasticity also occurs for adult USVs. To examine whether social exposure or deprivation throughout development leads to changes in USV perception, eleven female CBA/CaJ mice were trained to discriminate between 18 USVs of three different categories using operant conditioning procedures. Mice were group housed with four females or housed individually from weaning for the duration of the experiment. Socially housed and isolated mice differed in initial training times on pure tones, suggesting isolated mice had a more difficult time learning the task. Both groups completed USV discrimination conditions quicker at the end of the testing phases relative to the beginning. The overall discrimination of USVs did not differ between the two housing conditions, but a multidimensional scaling analysis revealed that socially experienced and isolated mice perceive some USVs differently, illustrated by differences in locations of USVs on the scaling maps from the two groups. Finally, a negative correlation was found between spectrotemporal similarity and percent discrimination, and analyses support the idea that mice may show categorical perception of at least two of the three USV categories. Thus, experience with USVs changes USV perception.

Memantine, a noncompetitive antagonist of the N-methyl-D-aspartate (NMDA) receptor, suppresses the release of excessive levels of glutamate that may induce neuronal excitation. Here we investigated the effects of memantine on salicylate-induced tinnitus model. The expressions of the activity-regulated cytoskeleton-associated protein (ARC) and tumor necrosis factor-alpha (TNFα) genes; as well as the NMDA receptor subunit 2B (NR2B) gene and protein, were examined in the SH-SY5Y cells and the animal model. We also used gap-prepulse inhibition of the acoustic startle reflex (GPIAS) and noise burst prepulse inhibition of acoustic startle, and the auditory brainstem level (electrophysiological recordings of auditory brainstem responses, ABR) and NR2B expression level in the auditory cortex to evaluate whether memantine could reduce salicylate-mediated behavioral disturbances. NR2B was significantly upregulated in salicylate-treated cells, but downregulated after memantine treatment. Similarly, expression of the inflammatory cytokine genes TNFα and immediate-early gene ARC was significantly increased in the salicylate-treated cells, and decreased when the cells were treated with memantine. These results were confirmed by NR2B immunocytochemistry. GPIAS was attenuated to a significantly lesser extent in rats treated with a combination of salicylate and memantine than in those treated with salicylate only. The mean ABR threshold in both groups was not significant different before and 1 day after the end of treatment. Additionally, NR2B protein expression in the auditory cortex was markedly increased in the salicylate-treated group, whereas it was reduced in the memantine-treated group. These results indicate that memantine is useful for the treatment of salicylate-induced tinnitus.

Interaural time differences (ITD) and interaural level differences (ILD) are physical cues that enable the auditory system to pinpoint the position of a sound source in space. This ability is crucial for animal communication and predator-prey interactions. The barn owl has evolved an exceptional sense of hearing and shows abilities of sound localisation that outperform most other species. So far, behavioural studies in the barn owl often used reflexive responses to investigate aspects of sound localisation. Furthermore, they predominately probed the higher frequencies of the owl's hearing range (> 3 kHz). In the present study we used a Go/NoGo paradigm to measure the barn owl's behavioural sound localisation acuity (expressed as the Minimum Audible Angle, MAA) as a function of stimulus type (narrow-band noise centred at 500, 1000, 2000, 4000 and 8000 Hz, and broad-band noise) and sound source position. We found significant effects of both stimulus type and sound source position on the barn owls' MAA. The MAA improved with increasing stimulus frequency, from 14° at 500 Hz to 6° at 8000 Hz. The smallest MAA of 4° was found for broadband noise stimuli. Comparing different sound source positions revealed smaller MAAs for frontal compared to lateral stimulus presentation, irrespective of stimulus type. These results are consistent with both the known variations in physical ITDs and variation in the width of neural ITD tuning curves with azimuth and frequency. Physical and neural characteristics combine to result in better spatial acuity for frontal compared to lateral sounds and reduced localisation acuity at lower frequencies.

Marmosets have attracted significant interest in the life sciences. Similarities with human brain anatomy and physiology, such as the granular frontal cortex, as well as the development of transgenic lines and potential for transferring rodent neuroscientific techniques to small primates make them a promising neurodegenerative and neuropsychiatric model system. However, whether marmosets can exhibit complex motor tasks in highly controlled experimental designs—one of the prerequisites for investigating higher-order control mechanisms underlying cognitive motor behavior—has not been demonstrated. We show that marmosets can be trained to perform vocal behavior in response to arbitrary visual cues in controlled operant conditioning tasks. Our results emphasize the marmoset as a suitable model to study complex motor behavior and the evolution of cognitive control underlying speech.

Bilateral cochlear implantation has provided access to some of the benefits of binaural hearing enjoyed by normal-hearing (NH) listeners. However, a gap in performance still exists between the two populations. Single-channel stimulation studies have shown that interaural place-of-stimulation mismatch (IPM) due to differences in implantation depth leads to decreased binaural fusion and lateralization of interaural time and level differences (ITDs and ILDs, respectively). While single-channel studies are informative, multi-channel stimulation is needed for good speech understanding with cochlear implants (CIs). Some multi-channel studies have shown that channel interaction due to current spread can affect ITD sensitivity. In this work, we studied the effect of IPM and channel spacing, along with their potential interaction, on binaural fusion and ITD/ILD lateralization. Experiments were conducted in adult NH listeners and CI listeners with a history of acoustic hearing. Results showed that IPM reduced the range of lateralization for ITDs but not ILDs. CI listeners were more likely to report a fused percept in the presence of IPM with multi-channel stimulation than NH listeners. However, no effect of channel spacing was found. These results suggest that IPM should be accounted for in clinical mapping practices in order to maximize bilateral CI benefits.

Separating sound sources in acoustic environments relies on making ongoing, highly accurate spectro-temporal comparisons. However, listeners with hearing impairment may have varying quality of temporal encoding within or across ears, which may limit the listeners' ability to make spectro-temporal comparisons between places-of-stimulation. In this study in normal hearing listeners, depth of amplitude modulation (AM) for sinusoidally amplitude modulated (SAM) tones was manipulated in an effort to reduce the coding of periodicity in the auditory nerve. The ability to judge differences in AM rates was studied for stimuli presented to different cochlear places-of-stimulation, within- or across-ears. It was hypothesized that if temporal encoding was poorer for one tone in a pair, then sensitivity to differences in AM rate of the pair would decrease. Results indicated that when the depth of AM was reduced from 50% to 20% for one SAM tone in a pair, sensitivity to differences in AM rate decreased. Sensitivity was greatest for AM rates near 90 Hz and depended upon the places-of-stimulation being compared. These results suggest that degraded temporal representations in the auditory nerve for one place-of-stimulation could lead to deficits comparing that temporal information with other places-of-stimulation.

The CHRNA7 gene that encodes the α 7 -subunit of the nicotinic acetylcholine receptor (α 7 -nAChR) has been associated with some autism spectrum disorders and other neurodevelopmental conditions characterized, in part, by auditory and language impairment. These conditions may include auditory processing disorders that represent impaired timing of neural activity, often accompanied by problems understanding speech. Here, we measure timing properties of sound-evoked activity via the auditory brainstem response (ABR) of α 7 -nAChR knockout mice of both sexes and wild-type colony controls. We find a significant timing delay in evoked ABR signals that represents midbrain activity in knockouts. We also examine spike-timing properties of neurons in the inferior colliculus, a midbrain nucleus that exhibits high levels of α 7 -nAChR during development. We find delays of evoked responses along with degraded spiking precision in knockout animals. We find similar timing deficits in responses of neurons in the superior paraolivary nucleus and ventral nucleus of the lateral lemniscus, which are brainstem nuclei thought to shape temporal precision in the midbrain. In addition, we find that other measures of temporal acuity including forward masking and gap detection are impaired for knockout animals. We conclude that altered temporal processing at the level of the brainstem in α 7 -nAChR-deficient mice may contribute to degraded spike timing in the midbrain, which may underlie the observed timing delay in the ABR signals. Our findings are consistent with a role for the α 7 -nAChR in types of neurodevelopmental and auditory processing disorders and we identify potential neural targets for intervention. NEW & NOTEWORTHY Disrupted signaling via the α 7 -nicotinic acetylcholine receptor (α 7 -nAChR) is associated with neurodevelopmental disorders that include impaired auditory processing. The underlying causes of dysfunction are not known but a common feature is abnormal timing of neural activity. We examined temporal processing of α 7 -nAChR knockout mice and wild-type controls. We found degraded spike timing of neurons in knockout animals, which manifests at the level of the auditory brainstem and midbrain.

An experiment was performed with 10 young normal-hearing listeners that attempted to determine if envelope modulations affected binaural processing in bandlimited pulse trains. Listeners detected an interaurally out-of-phase carrier pulse train in the presence of different amplitude modulations. The peaks of the pulses were constant (called "flat" or F), followed envelope modulations from an interaurally correlated 50-Hz bandwidth noise (called CM), or followed modulations from an interaurally uncorrelated noise (called UM). The pulse rate was varied from 50 to 500 pulses per second (pps) and the center frequency (CF) was 4 or 8 kHz. It was hypothesized that CM would cause no change or an increase in performance compared to F; UM would cause a decrease because of the blurring of the binaural detection cue. There was a small but significant decrease from F to CM (inconsistent with the hypothesis) and a further decrease from CM to UM (consistent with the hypothesis). Critically, there was a significant envelope by rate interaction caused by a decrease from F to CM for the 200-300 pps rates. The data can be explained by a subject-based factor, where some listeners experienced interaural envelope decorrelation when the sound was encoded by the auditory system that reduced performance when the modulations were present. Since the decrease in performance between F and CM conditions was small, it seems that most young normal-hearing listeners have very similar encoding of modulated stimuli across the ears. This type of task, when further optimized, may be able to assess if hearing-impaired populations experience interaural decorrelation from encoding modulated stimuli and therefore could help better understand the limited spatial hearing in populations like cochlear-implant users.

Our sensory environment changes constantly. Accordingly, neural systems continually adapt to the concurrent stimulus statistics to remain sensitive over a wide range of conditions. Such dynamic range adaptation (DRA) is assumed to increase both the effectiveness of the neuronal code and perceptual sensitivity. However, direct demonstrations of DRA-based efficient neuronal processing that also produces perceptual benefits are lacking. Here, we investigated the impact of DRA on spatial coding in the rodent brain and the perception of human listeners. Complex spatial stimulation with dynamically changing source locations elicited prominent DRA already on the initial spatial processing stage, the Lateral Superior Olive (LSO) of gerbils. Surprisingly, on the level of individual neurons, DRA diminished spatial tuning because of large response variability across trials. However, when considering single-trial population averages of multiple neurons, DRA enhanced the coding efficiency specifically for the concurrently most probable source locations. Intrinsic LSO population imaging of energy consumption combined with pharmacology revealed that a slow-acting LSO gain-control mechanism distributes activity across a group of neurons during DRA, thereby enhancing population coding efficiency. Strikingly, such "efficient cooperative coding" also improved neuronal source separability specifically for the locations that were most likely to occur. These location-specific enhancements in neuronal coding were paralleled by human listeners exhibiting a selective improvement in spatial resolution. We conclude that, contrary to canonical models of sensory encoding, the primary motive of early spatial processing is efficiency optimization of neural populations for enhanced source separability in the concurrent environment.

Speech comprehension difficulties are ubiquitous to aging and hearing loss, particularly in noisy environments. Older adults' poorer speech-in-noise (SIN) comprehension has been related to abnormal neural representations within various nodes (regions) of the speech network, but how senescent changes in hearing alter the transmission of brain signals remains unspecified. We measured electroencephalograms in older adults with and without mild hearing loss during a SIN identification task. Using functional connectivity and graph-theoretic analyses, we show that hearing-impaired (HI) listeners have more extended (less integrated) communication pathways and less efficient information exchange among widespread brain regions (larger network eccentricity) than their normal-hearing (NH) peers. Parameter optimized support vector machine classifiers applied to EEG connectivity data showed hearing status could be decoded (> 85% accuracy) solely using network-level descriptions of brain activity, but classification was particularly robust using left hemisphere connections. Notably, we found a reversal in directed neural signaling in left hemisphere dependent on hearing status among specific connections within the dorsal-ventral speech pathways. NH listeners showed an overall net "bottom-up" signaling directed from auditory cortex (A1) to inferior frontal gyrus (IFG; Broca's area), whereas the HI group showed the reverse signal (i.e., "top-down" Broca's → A1). A similar flow reversal was noted between left IFG and motor cortex. Our full-brain connectivity results demonstrate that even mild forms of hearing loss alter how the brain routes information within the auditory-linguistic-motor loop.

Age‐related hearing loss (ARHL) is a neurodegenerative disorder characterized by a gradual decrease in hearing sensitivity. Previous electrophysiological and behavioral studies have demonstrated that the CBA/CaJ mouse strain is an appropriate model for the late‐onset hearing loss found in humans. However, few studies have characterized hearing in these mice behaviorally using longitudinal methodologies. The goal of this research was to utilize a longitudinal design and operant conditioning procedures with positive reinforcement to construct audiograms and temporal integration functions in aging CBA/CaJ mice. In the first experiment, thresholds were collected for 8, 16, 24, 42, and 64 kHz pure tones in 30 male and 35 female CBA/CaJ mice. Similar to humans, mice had higher thresholds for high frequency tones than for low frequency pure tones across the lifespan. Female mice had better hearing acuity than males after 645 days of age. In the second experiment, temporal integration functions were constructed for 18 male and 18 female mice for 16 and 64 kHz tones varying in duration. Mice showed an increase in thresholds for tones shorter than 200 ms, reaching peak performance at shorter durations than other rodent species. Overall, CBA/CaJ mice experience ARHL for pure tones of different frequencies and durations, making them a good model for studies on hearing loss. These findings highlight the importance of using a wide range of stimuli and a longitudinal design when comparing presbycusis across different species.

Neural adaptation enables sensory information to be represented optimally in the brain despite large fluctuations over time in the statistics of the environment. Auditory contrast gain control represents an important example, which is thought to arise primarily from cortical processing. We find, however, that neurons in both the auditory thalamus and midbrain of mice show robust contrast gain control, and that this is implemented independently of cortical activity. Although neurons at each level exhibit contrast gain control to similar degrees, adaptation time constants become longer at later stages of the processing hierarchy, resulting in progressively more stable representations. We also show that auditory discrimination thresholds in human listeners compensate for changes in contrast, and that the strength of this perceptual adaptation can be predicted from physiological measurements. Contrast adaptation is therefore a robust property of both the subcortical and cortical auditory system and accounts for the short-term adaptability of perceptual judgments.

The brain has a remarkable capacity to adapt to changes in sensory inputs and to learn from experience. However, the neural circuits responsible for this flexible processing remain poorly understood. Using optogenetic silencing of ArchT-expressing neurons in adult ferrets, we show that within-trial activity in primary auditory cortex (A1) is required for training-dependent recovery in sound-localization accuracy following monaural deprivation. Because localization accuracy under normal-hearing conditions was unaffected, this highlights a specific role for cortical activity in learning. A1-dependent plasticity appears to leave a memory trace that can be retrieved, facilitating adaptation during a second period of monaural deprivation. However, in ferrets in which learning was initially disrupted by perturbing A1 activity, subsequent optogenetic suppression during training no longer affected localization accuracy when one ear was occluded. After the initial learning phase, the reweighting of spatial cues that primarily underpins this plasticity may therefore occur in A1 target neurons.

Auditory cortex is required for sound localisation, but how neural firing in auditory cortex underlies our perception of sound sources in space remains unclear. Specifically, whether neurons in auditory cortex represent spatial cues or an integrated representation of auditory space across cues is not known. Here, we measured the spatial receptive fields of neurons in primary auditory cortex (A1) while ferrets performed a relative localisation task. Manipulating the availability of binaural and spectral localisation cues had little impact on ferrets’ performance, or on neural spatial tuning. A subpopulation of neurons encoded spatial position consistently across localisation cue type. Furthermore, neural firing pattern decoders outperformed two-channel model decoders using population activity. Together, these observations suggest that A1 encodes the location of sound sources, as opposed to spatial cue values.

When two voices compete, listeners can segregate and identify concurrent speech sounds using pitch (fundamental frequency, F0) and timbre (harmonic) cues. Speech perception is also hindered by the signal-to-noise ratio (SNR). How clear and degraded concurrent speech sounds are represented at early, pre-attentive stages of the auditory system is not well understood. To this end, we measured scalp-recorded frequency-following responses (FFR) from the EEG while human listeners heard two concurrently presented, steady-state (time-invariant) vowels whose F0 differed by zero or four semitones (ST) presented diotically in either clean (no noise) or noise-degraded (+5dB SNR) conditions. Listeners also performed a speeded double vowel identification task in which they were required to identify both vowels correctly. Behavioral results showed that speech identification accuracy increased with F0 differences between vowels, and this perceptual F0 benefit was larger for clean compared to noise degraded (+5dB SNR) stimuli. Neurophysiological data demonstrated more robust FFR F0 amplitudes for single compared to double vowels and considerably weaker responses in noise. F0 amplitudes showed speech-on-speech masking effects, along with a non-linear constructive interference at 0ST, and suppression effects at 4ST. Correlations showed that FFR F0 amplitudes failed to predict listeners' identification accuracy. In contrast, FFR F1 amplitudes were associated with faster reaction times, although this correlation was limited to noise conditions. The limited number of brain-behavior associations suggests subcortical activity mainly reflects exogenous processing rather than perceptual correlates of concurrent speech perception. Collectively, our results demonstrate that FFRs reflect pre-attentive coding of concurrent auditory stimuli that only weakly predict the success of identifying concurrent speech.

This study describes sound localization and speech-recognition-in-noise abilities of a cochlear-implant user with electro-acoustic stimulation (EAS) in one ear, and a hearing aid in the contralateral ear. This listener had low-frequency, up to 250 Hz, residual hearing within the normal range in both ears. The objective was to determine how hearing devices affect spatial hearing for an individual with substantial unaided low-frequency residual hearing. Sound-localization performance was assessed for three sounds with different bandpass characteristics: low center frequency (100-400 Hz), mid center frequency (500-1,500 Hz) and high frequency broad-band (500-20,000 Hz) noise. Speech recognition was assessed with the Dutch Matrix sentence test presented in noise. Tests were performed while the listener used several on-off combinations of the devices. The listener localized low-center frequency sounds well in all hearing conditions, but mid-center frequency and high frequency broadband sounds were localized well almost exclusively in the completely unaided condition (mid-center frequency sounds were also localized well with the EAS device alone). Speech recognition was best in the fully aided condition with speech presented in the front and noise presented at either side. Furthermore, there was no significant improvement in speech recognition with all devices on, compared to when the listener used her cochlear implant only. Hearing aids and cochlear implant impair high frequency spatial hearing due to improper weighing of interaural time and level difference cues. The results reinforce the notion that hearing symmetry is important for sound localization. The symmetry is perturbed by the hearing devices for higher frequencies. Speech recognition depends mainly on hearing through the cochlear implant and is not significantly improved with the added information from hearing aids. A contralateral hearing aid provides benefit when the noise is spatially separated from the speech. However, this benefit is explained by the head shadow in that ear, rather than by an ability to spatially segregate noise from speech, as sound localization was perturbed with all devices in use.

Purpose Growing evidence supports the inclusion of perceptual tests that quantify the processing of temporal fine structure (TFS) in clinical hearing assessment. Many tasks have been used to evaluate TFS in the laboratory that vary greatly in the stimuli used and whether the judgments require monaural or binaural comparisons of TFS. The purpose of this study was to compare laboratory measures of TFS for inclusion in a battery of suprathreshold auditory tests. A subset of available TFS tasks were selected on the basis of potential clinical utility and were evaluated using metrics that focus on characteristics important for clinical use. Method TFS measures were implemented in replication of studies that demonstrated clinical utility. Monaural, diotic, and dichotic measures were evaluated in 11 young listeners with normal hearing. Measures included frequency modulation (FM) tasks, harmonic frequency shift detection, interaural phase difference (TFS–low frequency), interaural time difference (ITD), monaural gap duration discrimination, and tone detection in noise with and without a difference in interaural phase (N 0 S 0 , N 0 S π ). Data were compared with published results and evaluated with metrics of consistency and efficiency. Results Thresholds obtained were consistent with published data. There was no evidence of predictive relationships among the measures consistent with a homogenous group. The most stable tasks across repeated testing were TFS–low frequency, diotic and dichotic FM, and N 0 S π . Monaural and diotic FM had the lowest normalized variance and were the most efficient accounting for differences in total test duration, followed by ITD. Conclusions Despite a long stimulus duration, FM tasks dominated comparisons of consistency and efficiency. Small differences separated the dichotic tasks FM, ITD, and N 0 S π . Future comparisons following procedural optimization of the tasks will evaluate clinical efficiency in populations with impairment.

Hearing loss is a major risk factor for tinnitus, hyperacusis, and central auditory processing disorder. Although recent studies indicate that hearing loss causes neuroinflammation in the auditory pathway, the mechanisms underlying hearing loss-related pathologies are still poorly understood. We examined neuroinflammation in the auditory cortex following noise-induced hearing loss (NIHL) and its role in tinnitus in rodent models. Our results indicate that NIHL is associated with elevated expression of proinflammatory cytokines and microglial activation-two defining features of neuroinflammatory responses-in the primary auditory cortex (AI). Genetic knockout of tumor necrosis factor alpha (TNF-α) or pharmacologically blocking TNF-α expression prevented neuroinflammation and ameliorated the behavioral phenotype associated with tinnitus in mice with NIHL. Conversely, infusion of TNF-α into AI resulted in behavioral signs of tinnitus in both wild-type and TNF-α knockout mice with normal hearing. Pharmacological depletion of microglia also prevented tinnitus in mice with NIHL. At the synaptic level, the frequency of miniature excitatory synaptic currents (mEPSCs) increased and that of miniature inhibitory synaptic currents (mIPSCs) decreased in AI pyramidal neurons in animals with NIHL. This excitatory-to-inhibitory synaptic imbalance was completely prevented by pharmacological blockade of TNF-α expression. These results implicate neuroinflammation as a therapeutic target for treating tinnitus and other hearing loss-related disorders.