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Binaural noise suppresses linear click-evoked otoacoustic emissions more than ipsilateral or contralateral noise.

We studied the efferent suppression of click-evoked otoacoustic emissions with 65 dB SPL of white noise presented to left, right, or sometimes both, ears for 408 ms. Each burst of noise preceded a series of four unipolar 80 microseconds 65 dB peak Sound Pressure clicks, presented to the left ear only. The first click of the four-click group followed the end of the noise by either 1, 2, 5, 10, 20, 50, 100 or 200 ms; each subsequent click was offset by 20 additional ms via an ILO88 system with special programming modifications. Conditions were alternated so that a 'without noise' condition preceded a 'with noise' condition for three repetitions of 600 clicks per trial. Seven subjects with normal hearing participated in the study, and three of the seven participated in a test-retest reliability study. Results showed the greatest suppression followed binaural stimulation ending within one to five ms of the first click in the pulse train. Somewhat less suppression was seen following ipsilateral stimulation. The least amount of suppression was seen following contralateral stimulation, suggesting that previous research using contralateral stimulation may underestimate efferent effects. We saw no effects when the end of the noise was 100 ms or more away from the beginning of the click train.

Acoustic Stimulation↗

The relationship between sound transfer functions and hearing levels.

The effects of individuals differences in sound transfer function (STF) from free sound field to the tympanic membrane on hearing levels was studied in the right and left ears of 55 young male and 45 young female subjects. Furthermore, canal volumes and lengths have been recorded. STFs were measured in 1/3-octave bands using a miniature microphone and an attached probe. STF measurements were performed in the 0.5-16 kHz frequency range. Audiograms were registered with linear frequency sweep from 0.25 to 8 kHz. Transfer function spectra and magnitudes as well as ear canal dimensions were compared to hearing levels. There was a significant relationship between the dimensions of the ear canal and hearing levels. Large, compared to small, ear canal volumes resulted in a shift of STFs towards lower frequencies. STF spectra and magnitudes had a significant effect on hearing levels. Subject with low-frequency-dominated STFs have higher hearing thresholds than subjects with lower magnitude STFs.

Acoustic Stimulation↗

Envelope-following response and modulation transfer function in the dolphin's auditory system.

Potentials following the envelopes of sinusoidally amplitude-modulated tones (envelope response, EFR) were recorded from the head surface in bottle-nosed dolphins. EFR appeared at modulation rates from 300 to 3400 Hz. EFR amplitude was higher at rates from 500 to 1400 Hz with peaks at 600 and 1000 Hz and troughs at 700-850, 1200, and 2000 Hz; at rates above 1700 Hz it fell steeply. EFR dependence on modulation depth was linear except at the highest response amplitudes, which made it possible to obtain the modulation transfer function (MTF). EFR appears to be generated by several sources. One source had a latency of about 4 ms and followed modulation rates up to 1700 Hz, while another had a latency of 2 ms and followed modulation rates up to 3.4 kHz. The latencies of both sources coincided with those of waves of the auditory brainstem response (ABR). Comparison of MTF with the ABR spectrum had shown that several MTF peaks and troughs reflected the ABR spectrum. The latencies of the two sources were consistent with origins in the midbrain and auditory nerve, respectively.

Acoustic Stimulation↗

Cortical sources of middle latency responses of auditory evoked magnetic field.

In the recordings of middle latency responses of auditory evoked magnetic fields in 4 male subjects, we observed distinct components at 11, 19 and 33 ms after click stimulus. Equivalent current dipole sources of these components were located in the supratemporal auditory cortex, where the earliest component source was found at the most medial site.

Acoustic Stimulation↗

Evidence for spatio-topic organization of binaural processing in the human brainstem.

Three-channel Lissajous' trajectories (3-CLT) of the binaural interaction (BI) in auditory brainstem evoked potentials (ABEP) were derived from 13 normally and symmetrically hearing adults by subtracting the response to binaural clicks from the algebraic sum of monaural responses. ABEPs were recorded from four channels, three of them orthonormal to each other, in response to alternating polarity clicks, presented at a rate of 11/s with interaural time differences (ITD) of 0.2, 0.4 and 1.0 ms and an intensity of 65 dB nHL, or isochronic to both ears with interaural intensity differences (IIDs) of 5, 10 and 15 dB (65 dB nHL +/- 2.5, 5.0 and 7.5 dB, respectively). All 3-CLTs included 6 planar segments (labeled BdI, BdII, BdIII, BeI, BeII and Bf). Amplitudes of 3-CLT BI components were not significantly affected by increasing ITDs and IIDs, but latencies of all components increased significantly. The most remarkable finding was a significant change in apex orientations of BeI and BeII of the BI 3-CLT across stimulus conditions. The changes in BeI and BeII apex orientations, across stimulus conditions, may reflect differences in the anatomical representation of activity evoked by differently lateralized sounds. We suggest that this may indicate spatio-topic organization in the human brainstem.

Acoustic Stimulation↗

Processing of modulation frequency in the dorsal cochlear nucleus of the guinea pig: sinusoidal frequency-modulated tones.

Frequency- and amplitude-modulated (FM and AM, respectively) tones are important information-bearing elements in voice sounds and can also be produced by the spatial movement of sound sources. Zhao and Liang (1995) recently reported the response features of dorsal cochlear nucleus (DCN) neurons to AM tones. In the present study, the responses of the guinea pig DCN neurons to sinusoidal FM (SFM) tones were examined. Discharges of the DCN units to the SFM tones phase-locked to the stimulus modulation frequencies (fm). The phase-locked discharge patterns existed over broad ranges of modulation parameters and at stimulus levels as high as 95 dB SPL or modulation depths (dm) as low as 2%. Robust phase-locking to the fm was observed in samples of all DCN unit types studied. The means of best fm (Bfm) and upper limit fm (ULfm) of all recorded units were 510 Hz and 940 Hz, respectively. Pauser/Buildup (P/B) units had mean maximum synchronization index (SImax) of 0.57. ON units had the highest Bfm with the mean of 646 Hz and subtype ON-S showed the highest mean of SImax at 0.63. Phase-locking to the fm was independent of discharge rates and existed even when the discharge rates were reduced to the background spontaneous rate (SR). A few units showed stronger synchronous responses to the square and triangular FM stimuli instead of the SFM tones. The relationship between the modulated responses and the unit's response area were further examined. The fm phase-locking occurred to modulation bands (or frequency ranges) within the response area, with the modulation bands as narrow as +/- 160 Hz in the central inhibitory areas of the type IV units. As the width of the modulation band changed within a unit's response area, the phases of the fm phase-locked responses of P/B units linearly changed while for Onset units, the change was lesser. The P/B and Onset units had a pi phase shift and a pi/2 phase change, respectively, as carrier frequencies (fc5) passed through characteristic frequencies (CF) and the excitatory/inhibitory response boundaries. The phase-locked responses to the fms were dependent on the SR but were independent of the CF. Low-SR (< or = 2 spikes/s) units had higher synchronization of responses to the fm than the high-SR (> 2 spikes/s) units (SImax = 0.64 and 0.42 respectively). These results suggest that the temporal characteristics of the fm is effectively represented in the responses of DCN units to the SFM tones. Such temporal encoding behavior can play an important role in the processing of the complex sounds in the auditory system. These results also have implications for a possible role for the DCN is in identifying the spatial movement of a sound source.

Acoustic Stimulation↗

Interaural time and level differences: integrated or separated processing?

The processing of internal differences in time (IDT) and sound pressure level (IDL) was studied by using the mismatch negativity auditory evoked potential (MMN), which is a probe of pre-attentive auditory sensory memory. In a passive oddball experiment, subjects were reading in a book while they were presented with a standard stimulus (P = 0.88) having no IDTs or IDLs and three different deviant stimuli revealing an IDT, IDL, or both IDT and IDL. The different deviants elicited MMNs of comparable latencies indicating that memory representations of the IDTs and IDLs have been established. The MMN amplitudes to the IDT-IDL deviant were larger than those to changes in either IDT or IDL only. Moreover, the time-courses, amplitudes, and topographies of the MMNs to the IDT-IDL deviants were very similar to the sum of the MMNs elicited by the IDT and IDL deviants. These findings suggest that the representations of the binaural location cues were (at least partly) processed in parallel. It is argued that separate azimuth representations exist for IDT and IDL at a cortical level.

Acoustic Stimulation↗

Representation of acoustic signals in the eighth nerve of the Tokay gecko. II. Masking of pure tones with noise.

Acoustic signals are generally encoded in the peripheral auditory system of vertebrates by a duality scheme. For frequency components that fall within the excitatory tuning curve, individual eighth nerve fibers can encode the effective spectral energy by a spike-rate code, while simultaneously preserving the signal waveform periodicity of lower frequency components by phase-locked spike-train discharges. To explore how robust this duality of representation may be in the presence of noise, we recorded the responses of auditory fibers in the eighth nerve of the Tokay gecko to tonal stimuli when masking noise was added simultaneously. We found that their spike-rate functions reached plateau levels fairly rapidly in the presence of noise, so the ability to signal the presence of a tone by a concomitant change in firing rate was quickly lost. On the other hand, their synchronization functions maintained a high degree of phase-locked firings to the tone even in the presence of high-intensity masking noise, thus enabling a robust detection of the tonal signal. Critical ratios (CR) and critical bandwidths showed that in the frequency range where units are able to phaselock to the tonal periodicity, the CR bands were relatively narrow and the bandwidths were independent of noise level. However, to higher frequency tones where phaselocking fails and only spike-rate codes apply, the CR bands were much wider and depended upon noise level, so that their ability to filter tones out of a noisy background degraded with increasing noise levels. The greater robustness of phase-locked temporal encoding contrasted with spike-rate coding verifies a important advantage in using lower frequency signals for communication in noisy environments.

Acoustic Stimulation↗

Azimuthal sensitivity of rat pinna reflex: EMG recordings from cervicoauricular muscles.

Electromyographic (EMG) responses of the cervicoauricular muscles (CAM) to free-field sounds were recorded in two groups of rats whose brainstems were dissected transversely either at a pretectal or transtectal level. After the rat recovered from anesthesia, wide-band noise pulses were presented and speaker positions were varied systematically in azimuth. Sound levels were set at 10-15 dB above empirically determined threshold for an EMG response to a sound from 0 degree azimuth. In both animal groups, transient CAM EMGs with short latency were produced and three main types of azimuthal sensitivity of CAM EMG response were observed. (1) For the majority of the cases, an inverted "U' type of azimuthal sensitivity was identified: the maximum activity occurred around 0 degree azimuth, but as the speaker was moved toward either the ipsilateral or contralateral fields, the sound-evoked activity declined systematically. This directional tuning is quite different from the passive pinna directionality which is very lateral in the resting positions used in this study. (2) In a small number of cases, the spatial sensitivity curves were not symmetrical about the midline (0 degree azimuth): the EMG response was vigorous in one hemifield and dropped off systematically as the speaker was moved toward extreme positions of the other hemifield. Regardless of shapes of EMG spatial tuning curves, obstruction of either the ipsilateral or contralateral meatus reduced the sound-elicited response dramatically and eliminated the spatial sensitivity. (3) Some cases exhibited an omnidirectional function: the EMG spike rate had no or minor systematical variation as the speaker position was changed in azimuth. The results of this study indicate that with either pretectal or transtectal decerebrate preparations, the acoustically evoked CAM EMG can exhibit an azimuthal sensitivity which is based on binaural processing.

Acoustic Stimulation↗

Comparative study of fixed time versus intensity trade and fixed intensity versus time trade tests in sound lateralization.

OBJECTIVE: In the perception of sound lateralization a sound source exists in the side that the sound reaches an ear earlier in time or louder in intensity than the other ear. It is an imaginary phenomenon where the direction of sound is lateralized by an interaural time difference (ITD) and an interaural intensity difference (IID) of sound by both ears. It is speculated from pathophysiological findings that ITD and IID are processed by different pathways, but it has been not yet proved which of them is predominant in sound lateralization. The time difference can be traded by the intensity difference (fixed time versus intensity trade), and vice versa (fixed intensity versus time trade). In order to investigate predominance in this trade, we measured possible differences in effects by ITD and IID using two opposite trade phenomena of time versus intensity trade and intensity versus time trade. METHODS: The fixed time versus intensity trade test for selected narrow-band noise was investigated in 30 subjects with normal hearing. Using headphones, the subjects were instructed to push the button when the sound bias generated by fixed ITD (4 dB, 6 dB, 8 dB, 10 dB, 12 dB, 14 dB, 16 dB, 18 dB, 20 dB) was traded by gradually increased IID. The plot figure with fixed ITD and required IID to trade was made. It was compared with similar fixed intensity versus time trade test in 12 subjects with normal hearing. The fixed ITD (200-600 micros) was traded by gradually increased IID and the plot figure was made. RESULTS: In fixed time versus intensity trade test, each fixed ITD could be traded with an average of 6 dB of IID. On the other hand, in fixed intensity versus time trade test, ITD required to trade fixed IID increased, in proportion to fixed IID increased. CONCLUSION: It is concluded that predominance exists. In fixed time versus intensity trade test, the uniform amount of IID is required to trade the different ITD. However, in the novel fixed intensity versus time trade test, the phenomenon was in completely different manner that ITD in proportion to the given IID is required to trade.

Acoustic Stimulation↗

Effects of increasing visual load on aurally and visually guided target acquisition in a virtual environment.

The aim of the present study is to investigate interactions between vision and audition during a target acquisition task performed in a virtual environment. We measured the time taken to locate a visual target (acquisition time) signalled by auditory and/or visual cues in conditions of variable visual load. Visual load was increased by introducing a secondary visual task. The auditory cue was constructed using virtual three-dimensional (3D) sound techniques. The visual cue was constructed in the form of a 3D updating arrow. The results suggested that both auditory and visual cues reduced acquisition time as compared to an uncued condition. Whereas the visual cue elicited faster acquisition time than the auditory cue, the combination of the two cues produced the fastest acquisition time. The introduction of secondary visual task differentially affected acquisition time depending on cue modality. In conditions of high visual load, acquiring a target signalled by the auditory cue led to slower and more error-prone performance than acquiring a target signalled by either the visual cue alone or by both the visual and auditory cues.

Adolescent↗

Poor hand-pointing to sounds in right brain-damaged patients: not just a problem of spatial-hearing.

We asked 22 right brain-damaged (RBD) patients and 11 elderly healthy controls to perform hand-pointing movements to free-field unseen sounds, while modulating two non-auditory variables: the initial position of the responding hand (left, centre or right) and the presence or absence of task-irrelevant ambient vision. RBD patients suffering from visual neglect, unlike RBD patients without neglect and healthy controls, showed a systematic rightward error in sound localisation, which was modulated by the non-auditory variables. Localisation errors were exacerbated by initial hand-position to the right of the body-midline, and reduced by the leftwards initial hand-position. Moreover, for the visual neglect patients, mere presence of ambient vision worsened localisation errors. These results demonstrate that although hand-pointing to sounds has often been considered a straightforward approach to investigate sound-localisation abilities in brain-damaged patients, in some patients it may actually reveal localisation deficits that reflect a combination of impaired spatial-hearing and spatial biases from other sensory modalities (i.e., vision and proprioception).

Aged↗

A left-ear disadvantage for the presentation of irrelevant sound: manipulations of task requirements and changing state.

Three experiments attempted to clarify the effect of altering the spatial presentation of irrelevant auditory information. Previous research using serial recall tasks demonstrated a left-ear disadvantage for the presentation of irrelevant sounds (). Experiments 1 and 2 examined the effects of manipulating the location of irrelevant sound on either a mental arithmetic task () or a missing-item task (; Experiment 4). Experiment 3 altered the amount of change in the irrelevant stream to assess how this affected the level of interference elicited. Two prerequisites appear necessary to produce the left-ear disadvantage; the presence of ordered structural changes in the irrelevant sound and the requirement for serial order processing of the attended information. The existence of a left-ear disadvantage highlights the role of the right hemisphere in the obligatory processing of auditory information.

Adolescent↗

Directional hearing in a silicon cricket.

Phonotaxis is the ability to orient towards or away from sound sources. Crickets can locate conspecifics by phonotaxis to the calling (mating) song they produce, and can evade bats by negative phonotaxis from echolocation calls. The behaviour and underlying physiology have been studied in some depth, and the auditory system solves this complex problem in a unique manner. Experiments conducted on a simulation model of the system indicated that the mechanism output a directional signal to sounds ahead at calling song frequency and to sounds behind at echolocation frequencies. We suggest that this combination of responses helps simplify later processing in the cricket. To further explore this result, an analogue, very large scale integrated (aVLSI) circuit model of the mechanism was designed and built; results from testing this agreed with the simulation. The aVLSI circuit was used to test a further hypothesis about the potential advantages of the positioning of the acoustic inputs for sound localisation during walking. There was no clear advantage to the directionality of the system in their location. The aVLSI circuitry is now being extended to use on a robot along with previously modelled neural circuitry to better understand the complete sensorimotor pathway.

Animals↗

Effects of spatial separation and stimulus probability on the event-related potentials elicited by occasional changes in sound location.

The ability to extract information about the spatial location of sounds plays an important role in auditory scene analysis. The present study examined the effects of spatial separation and stimulus probability on auditory event-related potentials (ERPs) to changes in sound location. In Experiment 1, we found that difference waves between ERPs elicited by standard and deviant stimuli showed a biphasic negative-positive response peaking around 126 and 226 ms after deviant onset. The amplitude of both responses increased with decreasing deviant stimulus probability, and increasing stimulus deviance. When the same stimuli were presented with equal probability for all locations (Experiment 2), there were no significant differences in the ERP amplitude and latency. These results suggest that the data reported in Experiment 1 are the result of contextual changes, rather than changes in simple acoustic features. Brain electrical source analyses are consistent with generators located in auditory cortices posterior to Heschel's gyrus. Although occasional changes in sound location elicit earlier peaks than the mismatch negativity (MMN) response reported for other types of deviation, their topographical distribution and behavior are consistent with MMN. The early latency of MMN for changes in sound location is interpreted in the context of an early-warning system to alert the organism to new sound sources in the environment.

Acoustic Stimulation↗

Occasional changes in sound location enhance middle latency evoked responses.

Rapid processing of sound location is critical for orienting attention. The present study investigated whether contextually sensitive early neural responses elicited by occasional changes in sound location could be measured. Using an oddball paradigm with stimuli consisting of brief noise bursts whose location was occasionally varied using head-related transfer functions, we found significant enhanced negativities in the event-related potentials elicited by deviant stimuli as early as 25 ms after stimulus onset, in addition to the differences around 125 ms which have previously been reported. Recent research suggests that occasional changes in auditory location information are processed in areas beyond primary auditory cortex. Our data suggest that any such processing is in fact preceded by activation in primary auditory cortex.

Acoustic Stimulation↗

Selective attention to sound location or pitch studied with fMRI.

We used 3-T functional magnetic resonance imaging to compare the brain mechanisms underlying selective attention to sound location and pitch. In different tasks, the subjects (N = 10) attended to a designated sound location or pitch or to pictures presented on the screen. In the Attend Location conditions, the sound location varied randomly (left or right), while the pitch was kept constant (high or low). In the Attend Pitch conditions, sounds of randomly varying pitch (high or low) were presented at a constant location (left or right). Both attention to location and attention to pitch produced enhanced activity (in comparison with activation caused by the same sounds when attention was focused on the pictures) in widespread areas of the superior temporal cortex. Attention to either sound feature also activated prefrontal and inferior parietal cortical regions. These activations were stronger during attention to location than during attention to pitch. Attention to location but not to pitch produced a significant increase of activation in the premotor/supplementary motor cortices of both hemispheres and in the right prefrontal cortex, while no area showed activity specifically related to attention to pitch. The present results suggest some differences in the attentional selection of sounds on the basis of their location and pitch consistent with the suggested auditory "what" and "where" processing streams.

Acoustic Stimulation↗

The 'Franssen' illusion for short duration tones is preattentive: a study using mismatch negativity.

When a tone burst is divided into two parts, an onset transient and a sustained tone smoothly fading on, and these parts are delivered to two stereophonically located loudspeakers in a room, a listener gains the impression that the whole sound is coming from the loudspeaker that actually emits merely the transient. Due to this auditory illusion known as the 'Franssen effect' (FE), the physical and the perceived lateralizations of the sustained sound become different. A two-block mismatch negativity (MMN) paradigm was used to investigate the stage of auditory processing at which this segregation would take place. In one block, standard stimuli were 100 ms, 1 kHz tone bursts emitted by one of the loudspeakers, and deviant stimuli were their split version, with the sustained part switched to the other loudspeaker. In the other block, the roles of the two stimuli were swapped. A room acoustics software was used for generating the signals to a headphone. The responses recorded from 10 subjects displayed no MMN, although the same stimuli but without the transients evoked prominent MMNs. This indicated that the mechanism underlying this illusion modifies the neural representation of the stimulus with FE in such a way that it becomes similar to that of the stimulus without FE before reaching the input of the preattentive mechanism indexed by the MMN. Considering the possible relationship of this illusion to the precedence effect and also the relevant electrophysiological findings in the literature, we conclude that the primary auditory cortex is the most plausible site of the mechanism leading to the FE.

Acoustic Stimulation↗