Auditory Research publications

Development of electrophysiological means to assess the medial olivocochlear (MOC) system in humans is important to further our understanding of the function of that system and for the refinement and validation of psychoacoustical and otoacoustic emission methods which are thought to probe the MOC. Based on measurements in anesthetized animals it has been hypothesized that the MOC-reflex (MOCR) can enhance the response to signals in noise, and several lines of evidence support such a role in humans. A difficulty in these studies is the isolation of efferent effects. Efferent activation can be triggered by acoustic stimulation of the contralateral or ipsilateral ear, but ipsilateral stimulation is thought to be more effective. However, ipsilateral stimulation complicates interpretation of effects since these sounds can affect the perception of other ipsilateral sounds by mechanisms not involving olivocochlear efferents. We assessed the ipsilaterally evoked MOCR in human using a transtympanic procedure to record mass-potentials from the cochlear promontory or the niche of the round window. Averaged compound action potential (CAP) responses to masked probe tones of 4 kHz with and without a precursor (designed to activate the MOCR but not the stapedius reflex) were extracted with a polarity alternating paradigm. The masker was either a simultaneous narrow band noise masker or a short (20-ms) tonal ON- or OFF-frequency forward masker. The subjects were screened for normal hearing (audiogram, tympanogram, threshold stapedius reflex) and psychoacoustically tested for the presence of a precursor effect. We observed a clear reduction of CAP amplitude by the precursor, for different masking conditions. Even without an MOCR, this is expected because the precursor will affect the response to subsequent stimuli via neural adaptation. To determine whether the precursor also activated the efferent system, we measured the CAP over a range of masker levels, with or without precursor, and for different types of masker. The results show CAP reduction consistent with the type of gain reduction caused by the MOCR. These results generally support psychoacoustical paradigms designed to probe the efferent system as indeed activating the MOCR system, but not all observations are consistent with this mechanism.

Perceptual constancy requires neural representations that are selective for object identity, but also tolerant for identity-preserving transformations. How such representations arise in the brain and contribute to perception remains unclear. Here we studied tolerant representations of sound identity in the auditory system by recording multi-unit activity in tonotopic auditory cortex of ferrets discriminating the identity of vowels which co-varied across orthogonal stimulus dimensions (fundamental frequency, sound level, location and voicing). We found that neural decoding of vowel identity was most successful across the same orthogonal dimensions over which animals generalized their behavior. We also decoded orthogonal sound features and behavioral variables including choice and accuracy to show a behaviorally-relevant, multivariate and multiplexed representation of sound, with each variable represented over a distinct time-course. Finally, information content and timing of sound feature encoding was modulated by task-engagement and training, suggesting that tolerant representations during perceptual constancy are attentionally and experience-dependent.

The objective of this study was to demonstrate the efficacy of acute inactivation of brain areas by cooling in the behaving ferret and to demonstrate that cooling auditory cortex produced a localisation deficit that was specific to auditory stimuli. The effect of cooling on neural activity was measured in anesthetized ferret cortex. The behavioural effect of cooling was determined in a benchmark sound localisation task in which inactivation of primary auditory cortex (A1) is known to impair performance. Cooling strongly suppressed the spontaneous and stimulus-evoked firing rates of cortical neurons when the cooling loop was held at temperatures below 10°C, and this suppression was reversed when the cortical temperature recovered. Cooling of ferret auditory cortex during behavioural testing impaired sound localisation performance, with unilateral cooling producing selective deficits in the hemifield contralateral to cooling, and bilateral cooling producing deficits on both sides of space. The deficit in sound localisation induced by inactivation of A1 was not caused by motivational or locomotor changes since inactivation of A1 did not affect localisation of visual stimuli in the same context.

Enhanced detection and discrimination, along with faster reaction times, are the most typical behavioural manifestations of the brain's capacity to integrate multisensory signals arising from the same object. In this study, we examined whether multisensory behavioural gains are observable across different components of the localization response that are potentially under the command of distinct brain regions. We measured the ability of ferrets to localize unisensory (auditory or visual) and spatiotemporally coincident auditory–visual stimuli of different durations that were presented from one of seven locations spanning the frontal hemifield. During the localization task, we recorded the head movements made following stimulus presentation, as a metric for assessing the initial orienting response of the ferrets, as well as the subsequent choice of which target location to approach to receive a reward. Head‐orienting responses to auditory–visual stimuli were more accurate and faster than those made to visual but not auditory targets, suggesting that these movements were guided principally by sound alone. In contrast, approach‐to‐target localization responses were more accurate and faster to spatially congruent auditory–visual stimuli throughout the frontal hemifield than to either visual or auditory stimuli alone. Race model inequality analysis of head‐orienting reaction times and approach‐to‐target response times indicates that different processes, probability summation and neural integration, respectively, are likely to be responsible for the effects of multisensory stimulation on these two measures of localization behaviour.

Successful speech perception in real-world environments requires that the auditory system segregate competing voices that overlap in frequency and time into separate streams. Vowels are major constituents of speech and are comprised of frequencies (harmonics) that are integer multiples of a common fundamental frequency (F0). The pitch and identity of a vowel are determined by its F0 and spectral envelope (formant structure), respectively. When two spectrally overlapping vowels differing in F0 are presented concurrently, they can be readily perceived as two separate “auditory objects” with pitches at their respective F0s. A difference in pitch between two simultaneous vowels provides a powerful cue for their segregation, which in turn, facilitates their individual identification. The neural mechanisms underlying the segregation of concurrent vowels based on pitch differences are poorly understood. Here, we examine neural population responses in macaque primary auditory cortex (A1) to single and double concurrent vowels (/a/ and /i/) that differ in F0 such that they are heard as two separate auditory objects with distinct pitches. We find that neural population responses in A1 can resolve, via a rate-place code, lower harmonics of both single and double concurrent vowels. Furthermore, we show that the formant structures, and hence the identities, of single vowels can be reliably recovered from the neural representation of double concurrent vowels. We conclude that A1 contains sufficient spectral information to enable concurrent vowel segregation and identification by downstream cortical areas.

In patients with bilateral hearing loss, the use of two hearing aids (HAs) offers the potential to restore the benefits of binaural hearing, including sound source localization and segregation. However, existing evidence suggests that bilateral HA users’ access to binaural information, namely interaural time and level differences (ITDs and ILDs), can be compromised by device processing. Our objective was to characterize the nature and magnitude of binaural distortions caused by modern digital behind-the-ear HAs using a variety of stimuli and HA program settings. Of particular interest was a common frequency-lowering algorithm known as nonlinear frequency compression, which has not previously been assessed for its effects on binaural information. A binaural beamforming algorithm was also assessed. Wide dynamic range compression was enabled in all programs. HAs were placed on a binaural manikin, and stimuli were presented from an arc of loudspeakers inside an anechoic chamber. Stimuli were broadband noise bursts, 10-Hz sinusoidally amplitude-modulated noise bursts, or consonant–vowel–consonant speech tokens. Binaural information was analyzed in terms of ITDs, ILDs, and interaural coherence, both for whole stimuli and in a time-varying sense (i.e., within a running temporal window) across four different frequency bands (1, 2, 4, and 6 kHz). Key findings were: (a) Nonlinear frequency compression caused distortions of high-frequency envelope ITDs and significantly reduced interaural coherence. (b) For modulated stimuli, all programs caused time-varying distortion of ILDs. (c) HAs altered the relationship between ITDs and ILDs, introducing large ITD–ILD conflicts in some cases. Potential perceptual consequences of measured distortions are discussed.

Auditory information relayed by auditory nerve fibers and somatosensory information relayed by granule cell parallel fibers converge on the fusiform cells (FCs) of the dorsal cochlear nucleus, the first brain station of the auditory pathway. In vitro, parallel fiber synapses on FCs exhibit spike-timing-dependent plasticity with Hebbian learning rules, partially mediated by the NMDA receptor (NMDAr). Well-timed bimodal auditory-somatosensory stimulation, in vivo equivalent of spike-timing-dependent plasticity, can induce stimulus-timing-dependent plasticity (StTDP) of the FCs spontaneous and tone-evoked firing rates. In healthy guinea pigs, the resulting distribution of StTDP learning rules across a FC neural population is dominated by a Hebbian profile while anti-Hebbian, suppressive and enhancing LRs are less frequent. In this study, we investigate in vivo, the NMDAr contribution to FC baseline activity and long term plasticity. We find that blocking the NMDAr decreases the synchronization of FC- spontaneous activity and mediates differential modulation of FC rate-level functions such that low, and high threshold units are more likely to increase, and decrease, respectively, their maximum amplitudes. Three significant alterations in mean learning-rule profiles were identified: transitions from an initial Hebbian profile towards (1) an anti-Hebbian; (2) a suppressive profile; and (3) transitions from an anti-Hebbian to a Hebbian profile. FC units preserving their learning rules showed instead, NMDAr-dependent plasticity to unimodal acoustic stimulation, with persistent depression of tone-evoked responses changing to persistent enhancement following the NMDAr antagonist. These results reveal a crucial role of the NMDAr in mediating FC baseline activity and long-term plasticity which have important implications for signal processing and auditory pathologies related to maladaptive plasticity of dorsal cochlear nucleus circuitry.

From behavioral studies it is known that auditory spatial resolution of azimuthal space declines over age. To date, it is not clear how age affects the respective sensory auditory processing at the pre-attentive level. Here we tested the hypothesis that pre-attentive processing of behaviorally perceptible spatial changes is preserved in older adults. An EEG-study was performed in older adults (65-82 years of age) and a mismatch negativity (MMN) paradigm employed. Sequences of frequent standard stimuli of defined azimuthal positions were presented together with rarely occurring deviants shifted by 10° or 20° to the left or to the right of the standard. Standard positions were at +5° (central condition) from the midsagittal plane and at 65° in both lateral hemifields (±65°; lateral condition). The results suggest an effect of laterality on the pre-attentive change processing of spatial deviations in older adults: While for the central conditions deviants close to MAA threshold (i.e., 10°) yielded discernable MMNs, for lateral positions the respective MMN responses were only elicited by spatial deviations of 20° toward the midline (i.e., ±45°). Furthermore, MMN amplitudes were found to be insensitive to the magnitude of deviation (10°, 20°), which is contrary to recent studies with young adults (Bennemann et al., 2013) and hints to a deteriorated pre-attentive encoding of sound sources in older adults. The discrepancy between behavioral MAA data and present results are discussed with respect to the possibility that under the condition of active stimulus processing older adults might benefit from recruiting additional attentional top-down processes to detect small magnitudes of spatial deviations even within the lateral acoustic field.

Hearing Protection Devices (HPDs) can protect the ear against loud potentially damaging sounds while allowing lower-level sounds such as speech to be perceived. However, the impact of these devices on the ability to localize sound sources is not well known. To address this question, we propose two different methods: one behavioral and one dealing with acoustical measurements. For the behavioral method, sound localization performance was measured with, and without, HPDs on 20 listeners. Five HPDs, including both passive (non-linear attenuation) and three active (talk-through) systems were evaluated. The results showed a significant increase in localization errors, especially front-back and up-down confusions relative to the "naked ear" test condition for all of the systems tested, especially for the talk-through headphone system. For the acoustic measurement method, Head-Related Transfer Functions (HRTFs) were measured on an artificial head both without, and with the HPDs in place. The effects of the HPDs on the spectral cues for the localization of different sound sources in the horizontal plane were analyzed. Alterations of the Interaural Spectral Difference (ISD) cues were identified, which could explain the observed increase in front-back confusions caused by the talk-through headphone protectors.

The encoding of auditory spatial acuity (measured as the precision to distinguish between two spatially distinct stimuli) by neural circuits in both auditory cortices is a matter of ongoing research. Here, the event-related potential (ERP) mismatch negativity (MMN), a sensitive indicator of preattentive auditory change detection, was used to tap into the underlying mechanism of cortical representation of auditory spatial information. We characterized the MMN response affected by the degree of spatial deviance in lateral acoustic space using a passive oddball paradigm. Two stimulation conditions (SCs)-specifically focusing on the investigation of the mid- and far-lateral acoustic space-were considered: (1) 65° left standard position with deviant positions at 70, 75, and 80°; and (2) 95° left standard position with deviant positions at 90, 85, and 80°. Additionally, behavioral data on the minimum audible angle (MAA) were acquired for the respective standard positions (65, 95° left) to quantify spatial discrimination in separating distinct sound sources. The two measurements disclosed the linkage between the (preattentive) MMN response and the (attentive) behavioral threshold. At 65° spatial deviations as small as 5° reliably elicited MMNs. Thereby, the MMN amplitudes monotonously increased as a function of spatial deviation. At 95°, spatial deviations of 15° were necessary to elicit a valid MMN. The behavioral data, however, yielded no difference in mean MAA thresholds for position 65 and 95°. The different effects of laterality on MMN responses and MAA thresholds suggest a role of spatial selective attention mechanisms particularly relevant in active discrimination of neighboring sound sources, especially in the lateral acoustic space.

Stereotyped sequences of neural activity underlie learned vocal behavior in songbirds; principle neurons in the cortical motor nucleus HVC fire in stereotyped sequences with millisecond precision across multiple renditions of a song. The geometry of neural connections underlying these sequences is not known in detail though feed-forward chains are commonly assumed in theoretical models of sequential neural activity. In songbirds, a well-defined cortical-thalamic motor circuit exists but little is known the fine-grain structure of connections within each song nucleus. To examine whether the structure of song is critically dependent on long-range connections within HVC, we bilaterally transected the nucleus along the anterior-posterior axis in normal-hearing and deafened birds. The disruption leads to a slowing of song as well as an increase in acoustic variability. These effects are reversed on a time-scale of days even in deafened birds or in birds that are prevented from singing post-transection. The stereotyped song of zebra finches includes acoustic details that span from milliseconds to seconds--one of the most precise learned behaviors in the animal kingdom. This detailed motor pattern is resilient to disruption of connections at the cortical level, and the details of song variability and duration are maintained by offline homeostasis of the song circuit.

Determining if we, or another agent, were responsible for a sensory event can require an accurate sense of timing. Our sense of appropriate timing relationships must, however, be malleable as there is a variable delay between the physical timing of an event and when sensory signals concerning that event are encoded in the brain. One dramatic demonstration of such malleability involves having people repeatedly press a button thereby causing a beep. If a delay is inserted between button presses and beeps, when it is subsequently taken away beeps can seem to precede the button presses that caused them. For this to occur it is important that people feel they were responsible for instigating the beeps. In terms of their timing, as yet it is not clear what combination of events is important for motor-sensory temporal recalibration. Here, by introducing ballistic reaches of short or longer extent before a button press, we varied the delay between the intention to act and the sensory consequence of that action. This manipulation failed to modulate recalibration magnitude. By contrast, introducing a similarly lengthened delay between button presses and consequent beeps eliminated recalibration. Thus it would seem that the critical timing relationship for motor-sensory temporal recalibration is between tactile signals relating to the completion of an action and the subsequent auditory percept.