Catálogo de publicaciones - libros

Axons from the nucleus magnocellularis (NM) and their targets in nucleus laminaris (NL) form the circuit responsible for encoding interaural time difference (ITD). In barn owls, NL receives bilateral inputs from NM, such that axons from the ipsilateral NM enter NL dorsally, while contralateral axons enter from the ventral side. These afferents act as delay lines to create maps of ITD in NL. Since delay-line inputs are characterized by a precise latency to auditory stimulation, but the postsynaptic coincidence detectors respond to ongoing phase difference, we asked whether the latencies of a local group of axons were identical, or varied by multiples of the inverse of the frequency they respond to, i.e., to multiples of 2π phase. Intracellular recordings from NM axons were used to measure delay-line latencies in NL. Systematic shifts in conduction delay within NL accounted for the maps of ITD, but recorded latencies of individual inputs at nearby locations could vary by 2π or 4π. Therefore microsecond precision is achieved through sensitivity to phase delays, rather than absolute latencies. We propose that the auditory system “coarsely” matches ipsilateral and contralateral latencies using physical delay lines, so that inputs arrive at NL at about the same time, and then “finely” matches latency modulo 2π to achieve microsecond ITD precision.

We assessed neural sensitivity to interaural time differences (ITDs) conveyed in the temporal fine structure (TFS) of low-frequency sounds and ITDs conveyed in the temporal envelope of amplitude-modulated (AM’ed) high-frequency sounds. Using electroencephalography (EEG), we recorded brain activity to sounds in which the interaural phase difference (IPD) of the TFS (or the modulated temporal envelope) was repeatedly switched between leading in one ear or the other. When the amplitude of the tones is modulated equally in the two ears at 41 Hz, the interaural phase modulation (IPM) evokes an IPM following-response (IPM-FR) in the EEG signal. For low-frequency signals, IPM-FRs were reliably obtained, and largest for an IPM rate of 6.8 Hz and when IPD switches (around 0°) were in the range 45–90°. IPDs conveyed in envelope of high-frequency tones also generated IPM-FRs; response maxima occurred for IPDs switched between 0° and 180° IPD. This is consistent with the interpretation that distinct binaural mechanisms generate the IPM-FR at low and high frequencies, and with the reported physiological responses of medial superior olive (MSO) and lateral superior olive (LSO) neurons in other mammals. Low-frequency binaural neurons in the MSO are considered maximally activated by IPDs in the range 45–90°, consistent with their reception of excitatory inputs from both ears. High-frequency neurons in the LSO receive excitatory and inhibitory input from the two ears receptively—as such maximum activity occurs when the sounds at the two ears are presented out of phase.

The human pinna introduces spatial acoustic cues in terms of direction-dependent spectral patterns that shape the incoming sound. These cues are specifically useful for localization in the vertical dimension. Pinna cues exist at frequencies above approximately 5 kHz, a frequency range where people with hearing loss typically have their highest hearing thresholds. Since increased thresholds often are accompanied by reduced frequency resolution, there are good reasons to believe that many people with hearing loss are unable to discriminate these subtle spectral pinna-cue details, even if the relevant frequency region is amplified by hearing aids.

One potential solution to this problem is to provide hearing-aid users with artificially enhanced pinna cues—as if they were listening through oversized pinnas. In the present study, it was tested whether test subjects were better at discriminating spectral patterns similar to enlarged-pinna cues. The enlarged-pinna patterns were created by transposing (T) generic normal-sized pinna cues (N) one octave down, or by using the approach (W) suggested by Naylor and Weinrich (System and method for generating auditory spatial cues, United States Patent, 2011). The experiment was cast as a determination of simulated minimum audible angle (MAA) in the median saggital plane. 13 test subjects with sloping hearing loss and 11 normal-hearing test subjects participated. The normal-hearing test subjects showed similar discrimination performance with the T, W, and N-type simulated pinna cues, as expected. However, the results for the hearing-impaired test subjects showed only marginally lower MAAs with the W and T-cues compared to the N-cues, while the overall discrimination thresholds were much higher for the hearing-impaired compared to the normal-hearing test subjects.

Barn owls are keen hunters of moving prey. They have evolved an auditory system with impressive anatomical and physiological specializations for localizing their prey. Here we present behavioural data on the owl’s sensitivity for discriminating acoustic motion direction in azimuth that, for the first time, allow a direct comparison of neuronal and perceptual sensitivity for acoustic motion in the same model species. We trained two birds to report a change in motion direction within a series of repeating wideband noise stimuli. For any trial the starting point, motion direction, velocity (53–2400°/s), duration (30–225 ms) and angular range (12–72°) of the noise sweeps were randomized. Each test stimulus had a motion direction being opposite to that of the reference stimuli. Stimuli were presented in the frontal or the lateral auditory space. The angular extent of the motion had a large effect on the owl’s discrimination sensitivity allowing a better discrimination for a larger angular range of the motion. In contrast, stimulus velocity or stimulus duration had a smaller, although significant effect. Overall there was no difference in the owls’ behavioural performance between “inward” noise sweeps (moving from lateral to frontal) compared to “outward” noise sweeps (moving from frontal to lateral). The owls did, however, respond more often to stimuli with changing motion direction in the frontal compared to the lateral space. The results of the behavioural experiments are discussed in relation to the neuronal representation of motion cues in the barn owl auditory midbrain.

Many natural sounds have spectrotemporal signatures only on a statistical level, e.g. wind, fire or rain. While their local structure is highly variable, the spectrotemporal statistics of these auditory textures can be used for recognition. This suggests the existence of a neural representation of these statistics. To explore their encoding, we investigated the detectability of changes in the spectral statistics in relation to the properties of the change.

To achieve precise parameter control, we designed a minimal sound texture—a modified cloud of tones—which retains the central property of auditory textures: solely statistical predictability. Listeners had to rapidly detect a change in the frequency marginal probability of the tone cloud occurring at a random time.

The size of change as well as the time available to sample the original statistics were found to correlate positively with performance and negatively with reaction time, suggesting the accumulation of noisy evidence. In summary we quantified dynamic aspects of change detection in statistically defined contexts, and found evidence of integration of statistical information.

Previous studies have demonstrated that temporal envelope (E) is sufficient for speech perception, while fine structure (FS) is important for pitch perception for normal-hearing (NH) listeners. Listeners with sensorineural hearing loss (SNHL) have an impaired ability to use FS in lexical tone perception due to the reduced frequency resolution. Listeners with auditory neuropathy spectrum disorder (ANSD) may have deficits in temporal resolution. Little is known about how such deficits may impact their ability to use E and FS to perceive lexical tone, and whether it is the deficit in temporal resolution or frequency resolution that may lead to more detrimental effects on FS processing in pitch perception. Three experiments were conducted in the present study. Experiment I used the “auditory chimera” technique to investigate how SNHL and ANSD listeners would achieve lexical tone recognition using either the E or the FS cues. Experiment II tested their frequency resolution as measured with their psychophysical tuning curves (PTCs). Experiment III tested their temporal resolution as measured with the temporal gap detection (TGD) threshold. The results showed that the SNHL listeners had reduced frequency selectivity, but intact temporal resolution ability, while the ANSD listeners had degraded temporal resolution ability, but intact frequency selectivity. In comparison with the SNHL listeners, the ANSD listeners had severely degraded ability to process the FS cues and thus their ability to perceive lexical tone mainly depended on the ability to use the E cues. These results suggested that, in comparison with the detrimental impact of the reduced frequency selectivity, the impaired temporal resolution may lead to more degraded FS processing in pitch perception.

Object binding cues such as binaural and across-frequency modulation cues are likely to be used by the auditory system to separate sounds from different sources in complex auditory scenes. The present study investigates the interaction of these cues in a binaural masking pattern paradigm where a sinusoidal target is masked by a narrowband noise. It was hypothesised that beating between signal and masker may contribute to signal detection when signal and masker do not spectrally overlap but that this cue could not be used in combination with interaural cues. To test this hypothesis an additional sinusoidal interferer was added to the noise masker with a lower frequency than the noise whereas the target had a higher frequency than the noise. Thresholds increase when the interferer is added. This effect is largest when the spectral interferer-masker and masker-target distances are equal. The result supports the hypothesis that modulation cues contribute to signal detection in the classical masking paradigm and that these are analysed with modulation bandpass filters. A monaural model including an across-frequency modulation process is presented that account for this effect. Interestingly, the interferer also affects dichotic thresholds indicating that modulation cues also play a role in binaural processing.

The speech intelligibility index (SII) calculation is based on the assumption that the effective range of signal-to-noise ratio (SNR) regarding speech intelligibility is [‑15 dB; +15 dB]. In a specific frequency band, speech intelligibility would remain constant by varying the SNRs above + 15 dB or below ‑15 dB. These assumptions were tested in four experiments measuring speech reception thresholds (SRTs) with a speech target and speech-spectrum noise, while attenuating target or noise above or below 1400 Hz, with different levels of attenuation in order to test different SNRs in the two bands. SRT varied linearly with attenuation at low-attenuation levels and an asymptote was reached for high-attenuation levels. However, this asymptote was reached (intelligibility was not influenced by further attenuation) for different attenuation levels across experiments. The ‑15-dB SII limit was confirmed for high-pass filtered targets, whereas for low-pass filtered targets, intelligibility was further impaired by decreasing the SNR below ‑15 dB (until ‑37 dB) in the high-frequency band. For high-pass and low-pass filtered noises, speech intelligibility kept improving when increasing the SNR in the rejected band beyond + 15 dB (up to 43 dB). Before reaching the asymptote, a 10-dB increase of SNR obtained by filtering the noise resulted in a larger decrease of SRT than a corresponding 10-dB decrease of SNR obtained by filtering the target (the slopes SRT/attenuation were different depending on which source was filtered). These results question the use of the SNR range and the importance function adopted by the SII when considering sharply filtered signals.

Dynamic aspects of cochlear mechanical compression were studied by recording basilar membrane (BM) vibrations evoked by tone pairs (“beat stimuli”) in the 11–19 kHz region of the gerbil cochlea. The frequencies of the stimulus components were varied to produce a range of “beat rates” at or near the characteristic frequency (CF) of the BM site under study, and the amplitudes of the components were balanced to produce near perfect periodic cancellations, visible as sharp notches in the envelope of the BM response. We found a compressive relation between instantaneous stimulus intensity and BM response magnitude that was strongest at low beat rates (e.g., 10–100 Hz). At higher beat rates, the amount of compression reduced progressively (i.e. the responses became linearized), and the rising and falling flanks of the response envelope showed increasing amounts of hysteresis; the rising flank becoming steeper than the falling flank. This hysteresis indicates that cochlear mechanical compression is not instantaneous, and is suggestive of a gain control mechanism having finite attack and release times. In gain control terms, the linearization that occurs at higher beat rates occurs because the instantaneous gain becomes smoothened, or low-pass filtered, with respect to the magnitude fluctuations in the stimulus. In terms of peripheral processing, the linearization corresponds to an enhanced coding, or decompression, of rapid amplitude modulations. These findings are relevant both to those who wish to understand the underlying mechanisms and those who need a realistic model of nonlinear processing by the auditory periphery.

The perceptual insensitivity to low frequency (LF) sound in humans has led to an underestimation of the physiological impact of LF exposure on the inner ear. It is known, however, that intense, LF sound causes cyclic changes of indicators of inner ear function after LF stimulus offset, for which the term “Bounce” phenomenon has been coined.

Here, we show that the mechanical amplification of hair cells (OHCs) is significantly affected after the presentation of LF sound. First, we show the Bounce phenomenon in slow level changes of quadratic, but not cubic, distortion product otoacoustic emissions (DPOAEs). Second, Bouncing in response to LF sound is seen in slow, oscillating frequency and correlated level changes of spontaneous otoacoustic emissions (SOAEs). Surprisingly, LF sound can induce new SOAEs which can persist for tens of seconds. Further, we show that the Bounce persists under free-field conditions, i.e. without an in-ear probe occluding the auditory meatus. Finally, we show that the Bounce is affected by contralateral acoustic stimulation synchronised to the ipsilateral LF sound. These findings clearly demonstrate that the origin of the Bounce lies in the modulation of cochlear amplifier gain. We conclude that activity changes of OHCs are the source of the Bounce, most likely caused by a temporary disturbance of OHC calcium homeostasis. In the light of these findings, the effects of long-duration, anthropogenic LF sound on the human inner ear require further research.