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1.  Reverberation impairs brainstem temporal representations of voiced vowel sounds: challenging “periodicity-tagged” segregation of competing speech in rooms 
The auditory system typically processes information from concurrently active sound sources (e.g., two voices speaking at once), in the presence of multiple delayed, attenuated and distorted sound-wave reflections (reverberation). Brainstem circuits help segregate these complex acoustic mixtures into “auditory objects.” Psychophysical studies demonstrate a strong interaction between reverberation and fundamental-frequency (F0) modulation, leading to impaired segregation of competing vowels when segregation is on the basis of F0 differences. Neurophysiological studies of complex-sound segregation have concentrated on sounds with steady F0s, in anechoic environments. However, F0 modulation and reverberation are quasi-ubiquitous. We examine the ability of 129 single units in the ventral cochlear nucleus (VCN) of the anesthetized guinea pig to segregate the concurrent synthetic vowel sounds /a/ and /i/, based on temporal discharge patterns under closed-field conditions. We address the effects of added real-room reverberation, F0 modulation, and the interaction of these two factors, on brainstem neural segregation of voiced speech sounds. A firing-rate representation of single-vowels' spectral envelopes is robust to the combination of F0 modulation and reverberation: local firing-rate maxima and minima across the tonotopic array code vowel-formant structure. However, single-vowel F0-related periodicity information in shuffled inter-spike interval distributions is significantly degraded in the combined presence of reverberation and F0 modulation. Hence, segregation of double-vowels' spectral energy into two streams (corresponding to the two vowels), on the basis of temporal discharge patterns, is impaired by reverberation; specifically when F0 is modulated. All unit types (primary-like, chopper, onset) are similarly affected. These results offer neurophysiological insights to perceptual organization of complex acoustic scenes under realistically challenging listening conditions.
PMCID: PMC4290552  PMID: 25628545
reverberation; pitch; auditory scene analysis; vowel; double vowels; cochlear nucleus
2.  Perceptual organization of sound begins in the auditory periphery 
Current biology : CB  2008;18(15):1124-1128.
Segmenting the complex acoustic mixture that makes a typical auditory scene into relevant perceptual objects is one of the main challenges of the auditory system [1], for both human and non-human species. Several recent studies indicate that perceptual auditory object formation, or “streaming”, may be based on neural activity within auditory cortex and beyond [2, 3]. Here, we find that scene analysis starts much earlier in the auditory pathways. Single units were recorded from a peripheral structure of the mammalian auditory brainstem, the cochlear nucleus. Peripheral responses were similar to cortical responses and displayed all of the functional properties required for streaming, including multi-second adaptation. Behavioral streaming was also measured in human listeners. Neurometric functions derived from the peripheral responses predicted accurately behavioral streaming. This reveals that sub-cortical structures may already contribute to the analysis of auditory scenes. This finding is consistent with the observation that species lacking a neocortex can still achieve and benefit from behavioral streaming [4]. For humans, we argue that auditory scene analysis of complex scenes is likely to be based on interactions between sub-cortical and cortical neural processes, with the relative contribution of each stage depending on nature of the acoustic cues forming the streams.
PMCID: PMC2559912  PMID: 18656355
3.  The role of auditory nerve innervation and dendritic filtering in shaping onset responses in the ventral cochlear nucleus 
Brain Research  2009;1247(C):221-234.
Neurons in the ventral cochlear nucleus (VCN) that respond primarily at the onset of a pure tone stimulus show diversity in terms of peri-stimulus-time-histograms (PSTHs), rate-level functions, frequency tuning, and also their responses to broad band noise. A number of different mechanisms have been proposed as contributing to the onset characteristic: e.g. coincidence, depolarisation block, and low-threshold potassium currents. We show that a simple point neuron receiving convergent inputs from high-spontaneous rate auditory nerve (AN) fibers, with no special currents and no peri-stimulatory shifts in firing threshold, is sufficient to produce much of the diversity seen experimentally. Three sub-classes of onset PSTHs: onset-ideal (OI), onset-chopper (OC) and onset-locker (OL) are reproduced by variations in innervation patterns and dendritic filtering. The factors shaping responses were explored by systematically varying key parameters. An OI response in this model requires a narrow range of AN input best frequencies (BF) which only produce supra-threshold depolarizations during the stimulus onset. For OC and OL responses, receptive fields were wider. Considerable low pass filtering of AN inputs away from BF results in an OL, whilst relatively unfiltered inputs produce an OC response. Rate-level functions in response to pure tones can be sloping, or plateau. These can be also reproduced in the model by the manipulation of the AN innervation. The model supports the coincidence detection hypothesis, and suggests that differences in excitatory innervation and dendritic filtering constant are important factors to consider when accounting for the variation in response characteristics seen in VCN onset units.
PMCID: PMC2653631  PMID: 18848923
Onset; Stellate; Cochlear nucleus; Point-neuron; PSTH; Rate-level functions
4.  Behavioral and physiological correlates of temporal pitch perception in electric and acoustic hearing 
In the “4-6” condition of experiment 1, normal-hearing (NH) listeners compared the pitch of a bandpass-filtered pulse train, whose inter-pulse intervals (IPIs) alternated between 4 and 6 ms, to that of isochronous pulse trains. Consistent with previous results obtained at a lower signal level, the pitch of the 4-6 stimulus corresponded to that of an isochronous pulse train having a period of 5.7 ms – longer than the mean IPI of 5 ms. In other conditions the IPI alternated between 3.5-5.5 ms and 4.5-6.5 ms. Experiment 2 was similar but presented electric pulse trains to one channel of a CI. In both cases, as overall IPI increased, the pitch of the alternating-interval stimulus approached that of an isochronous train having a period equal to the mean IPI. Experiment 3 measured compound action potentials (CAPs) to alternating-interval stimuli in guinea pigs and in NH listeners. The CAPs to pulses occurring after 4-ms intervals were smaller than responses to pulses occurring after 6-ms intervals, resulting in a modulated pattern that was independent of overall level. The results are compared to the predictions of a simple model incorporating auditory-nerve (AN) refractoriness, and where pitch is estimated from 1st-order intervals in the AN response.
PMCID: PMC2279014  PMID: 18247900

Results 1-4 (4)