Contributions of Intrinsic Neural and Stimulus Variance to Binaural Sensitivity

The discrimination of a change in a stimulus is determined both by the magnitude of that change and by the variability in the neural response to the stimulus. When the stimulus is itself noisy, then the relative contributions of the neural (intrinsic) and stimulus induced variability becomes a critical question. We measured the contribution of intrinsic neural noise and interstimulus variability to the discrimination of interaural time differences (ITDs) and interaural correlation (IAC). We measured discharge rate versus characteristic frequency (CF) tone ITD functions, and CF-centered narrowband noise ITD and IAC functions in interleaved blocks in the same units in the inferior colliculus of urethane-anesthetized guinea pigs. Ten “frozen” tokens of noise were synthesized and the responses to each token were separately analyzed to allow the relative contributions of intrinsic and stimulus variability to be assessed. ITD and IAC discrimination thresholds were determined for a simulated two-interval forced-choice experiment, based on the firing rate distributions, using receiver operating characteristic analysis. On average, between stimulus variability contributed 19% (range, 1.5–30%) of the variance in noise ITD discrimination and 27% (range, 3–50%) in IAC discrimination. Noise ITD thresholds were slightly higher than tone ITD thresholds. Taking the mean of the thresholds for individual noise tokens gave a similar result to pooling across all noise tokens. This implies that although the stimulus induced variability is measurable, it is insignificant in relation to the intrinsic noise in ITD and IAC discrimination.     

Responses of Auditory Nerve and Anteroventral Cochlear Nucleus Fibers to Broadband and Narrowband Noise: Implications for the Sensitivity to Interaural Delays

The quality of temporal coding of sound waveforms in the monaural afferents that converge on binaural neurons in the brainstem limits the sensitivity to temporal differences at the two ears. The anteroventral cochlear nucleus (AVCN) houses the cells that project to the binaural nuclei, which are known to have enhanced temporal coding of low-frequency sounds relative to auditory nerve (AN) fibers. We applied a coincidence analysis within the framework of detection theory to investigate the extent to which AVCN processing affects interaural time delay (ITD) sensitivity. Using monaural spike trains to a 1-s broadband or narrowband noise token, we emulated the binaural task of ITD discrimination and calculated just noticeable differences (jnds). The ITD jnds derived from AVCN neurons were lower than those derived from AN fibers, showing that the enhanced temporal coding in the AVCN improves binaural sensitivity to ITDs. AVCN processing also increased the dynamic range of ITD sensitivity and changed the shape of the frequency dependence of ITD sensitivity. Bandwidth dependence of ITD jnds from AN as well as AVCN fibers agreed with psychophysical data. These findings demonstrate that monaural preprocessing in the AVCN improves the temporal code in a way that is beneficial for binaural processing and may be crucial in achieving the exquisite sensitivity to ITDs observed in binaural pathways.     

Channel Interaction and Current Level Affect Across-Electrode Integration of Interaural Time Differences in Bilateral Cochlear-Implant Listeners

Sensitivity to interaural time differences (ITDs) is important for sound localization. Normal-hearing listeners benefit from across-frequency processing, as seen with improved ITD thresholds when consistent ITD cues are presented over a range of frequency channels compared with when ITD information is only presented in a single frequency channel. This study aimed to clarify whether cochlear-implant (CI) listeners can make use of similar processing when being stimulated with multiple interaural electrode pairs transmitting consistent ITD information. ITD thresholds for unmodulated, 100-pulse-per-second pulse trains were measured in seven bilateral CI listeners using research interfaces. Consistent ITDs were presented at either one or two electrode pairs at different current levels, allowing for comparisons at either constant level per component electrode or equal overall loudness. Different tonotopic distances between the pairs were tested in order to clarify the potential influence of channel interaction. Comparison of ITD thresholds between double pairs and the respective single pairs revealed systematic effects of tonotopic separation and current level. At constant levels, performance with double-pair stimulation improved compared with single-pair stimulation but only for large tonotopic separation. Comparisons at equal overall loudness revealed no benefit from presenting ITD information at two electrode pairs for any tonotopic spacing. Irrespective of electrode-pair configuration, ITD sensitivity improved with increasing current level. Hence, the improved ITD sensitivity for double pairs found for a large tonotopic separation and constant current levels seems to be due to increased loudness. The overall data suggest that CI listeners can benefit from combining consistent ITD information across multiple electrodes, provided sufficient stimulus levels and that stimulating electrode pairs are widely spaced.     

Sensitivity to Interaural Correlation of Single Neurons in the Inferior Colliculusof Guinea Pigs

Sensitivity to changes in the interaural correlation of 50-ms bursts of narrowband or broadband noise was measured in single neurons in the inferior colliculus of urethane-anaesthetized guinea pigs. Rate vs. interaural correlation functions (rICFs) were measured using two methods. These methods compensated in different ways for the inherent variance in interaural correlation between tokens with the same expected correlation. The shape of all rICFs could be best described by power functions allowing them to be summarized by two parameters. Most rICFs were best fit by a power below 2, indicating that they were only slightly nonlinear. However, there were a few fitted functions that had a power of 3–6, indicating marked curvature. Modeling results indicate that the nonlinearity of the majority of rICFs was explicable in terms of the monaural transduction stages; however, some of the rICFs with power greater than 2 require either multiple inputs to the coincidence detector or additional nonlinearities to be included in the model. Discrimination thresholds were estimated at reference correlations of −1, 0, and +1 using receiver operating characteristic (ROC) analysis of the spike-count distribution at each correlation. Thresholds spanned the full possible range, from a minimum of 0.1 to the maximum possible of 2. Thresholds were generally highest with a reference correlation of −1, intermediate with a reference of 0, and lowest with a reference correlation of +1. Thresholds were lowest for the most steeply sloped rICFs, but thresholds were not strongly correlated to the spike rate variance. The lowest thresholds occurred using narrowband noise that was compensated for internal delays, but they were still about three times larger than human psychophysical thresholds measured using similar stimuli. The data suggest that, unlike pure tone interaural time difference, discrimination of a population measure is required to account for behavioral interaural correlation discrimination performance.     

The Interaural Time Difference Pathway: a Comparison of Spectral Bandwidth and Correlation Sensitivity at Three Anatomical Levels

Temporal differences between the two ears are critical for spatial hearing. They can be described along axes of interaural time difference (ITD) and interaural correlation, and their processing starts in the brainstem with the convergence of monaural pathways which are tuned in frequency and which carry temporal information. In previous studies, we examined the bandwidth (BW) of frequency tuning at two stages: the auditory nerve (AN) and inferior colliculus (IC), and showed that BW depends on characteristic frequency (CF) but that there is no difference in the mean BW of these two structures when measured in a binaural, temporal framework. This suggested that there is little frequency convergence in the ITD pathway between AN and IC and that frequency selectivity determined by the cochlear filter is preserved up to the IC. Unexpectedly, we found that AN and IC neurons can be similar in CF and BW, yet responses to changes in interaural correlation in the IC were different than expected from coincidence patterns (“pseudo-binaural” responses) in the AN. To better understand this, we here examine the responses of bushy cells, which provide monaural inputs to binaural neurons. Using broadband noise, we measured BW and correlation sensitivity in the cat trapezoid body (TB), which contains the axons of bushy cells. This allowed us to compare these two metrics at three stages in the ITD pathway. We found that BWs in the TB are similar to those in the AN and IC. However, TB neurons were found to be more sensitive to changes in stimulus correlation than AN or IC neurons. This is consistent with findings that show that TB fibers are more temporally precise than AN fibers, but is surprising because it suggests that the temporal information available monaurally is not fully exploited binaurally.     

Detection and Identification of Monaural and Binaural Pitch Contours in Dyslexic Listeners

The use of binaural pitch stimuli to test for the presence of binaural auditory impairment in reading-disabled subjects has so far led to contradictory outcomes. While some studies found that a majority of dyslexic subjects was unable to perceive binaural pitch, others obtained a clear response of dyslexic listeners to Huggins’ pitch (HP). The present study clarified whether impaired binaural pitch perception is found in dyslexia. Results from a pitch contour identification test, performed in 31 dyslexic listeners and 31 matched controls, clearly showed that dyslexics perceived HP as well as the controls. Both groups also showed comparable results with a similar-sounding, but monaurally detectable, pitch-evoking stimulus. However, nine of the dyslexic subjects were found to have difficulty identifying pitch contours both in the binaural and the monaural conditions. The ability of subjects to correctly identify pitch contours was found to be significantly correlated to measures of frequency discrimination. This correlation may be attributed to the similarity of the experimental tasks and probably reflects impaired cognitive mechanisms related to auditory memory or auditory attention rather than impaired low-level auditory processing per se.     

Models of Brainstem Responses to Bilateral Electrical Stimulation

A simple, biophysically specified cell model is used to predict responses of binaurally sensitive neurons to patterns of input spikes that represent stimulation by acoustic and electric waveforms. Specifically, the effects of changes in parameters of input spike trains on model responses to interaural time difference (ITD) were studied for low-frequency periodic stimuli, with or without amplitude modulation. Simulations were limited to purely excitatory, bilaterally driven cell models with basic ionic currents and multiple input fibers. Parameters explored include average firing rate, synchrony index, modulation frequency, and latency dispersion of the input trains as well as the excitatory conductance and time constant of individual synapses in the cell model. Results are compared to physiological recordings from the inferior colliculus (IC) and discussed in terms of ITD-discrimination abilities of listeners with cochlear implants. Several empirically observed aspects of ITD sensitivity were simulated without evoking complex neural processing. Specifically, our results show saturation effects in rate–ITD curves, the absence of sustained responses to high-rate unmodulated pulse trains, the renewal of sensitivity to ITD in high-rate trains when inputs are amplitude-modulated, and interactions between envelope and fine-structure delays for some modulation frequencies.     

Interaural Time-Delay Sensitivity in Bilateral Cochlear Implant Users: Effects of Pulse Rate, Modulation Rate, and Place of Stimulation

Electrical interaural time delay (ITD) discrimination was measured using 300-ms bursts applied to binaural pitch matched electrodes at basal, mid, and apical locations in each ear. Six bilateral implant users, who had previously shown good ITD sensitivity at a pulse rate of 100 pulses per second (pps), were assessed. Thresholds were measured as a function of pulse rate between 100 and 1,000 Hz, as well as modulation rate over that same range for high-rate pulse trains at 6,000 pps. Results were similar for all three places of stimulation and showed decreasing ITD sensitivity as either pulse rate or modulation rate increased, although the extent of that effect varied across subjects. The results support a model comprising a common ITD mechanism for high- and low-frequency places of stimulation, which, for electrical stimulation, is rate-limited in the same way across electrodes because peripheral temporal responses are largely place invariant. Overall, ITD sensitivity was somewhat better with unmodulated pulse trains than with high-rate pulse trains modulated at matched rates, although comparisons at individual rates showed that difference to be significant only at 300 Hz. Electrodes presenting with the lowest thresholds at 600 Hz were further assessed using bursts with a ramped onset of 10 ms. The slower rise time resulted in decreased performance in four of the listeners, but not in the two best performers, indicating that those two could use ongoing cues at 600 Hz. Performance at each place was also measured using single-pulse stimuli. Comparison of those data with the unmodulated 300-ms burst thresholds showed that on average, the addition of ongoing cues beyond the onset enhanced overall ITD sensitivity at 100 and 300 Hz, but not at 600 Hz. At 1,000 Hz, the added ongoing cues actually decreased performance. That result is attributed to the introduction of ambiguous cues within the physiologically relevant range and increased dichotic firing.     

Detection of interaural time differences for clicks and tone pips: effects of interaural signal disparity

The ability to detect small interaural time differences (delta t) was determined in 4 subjects using clicks or long tone pips of various interaural signal disparities which are expressed as the extent of interaural spectral overlaps. The interaural signal disparity was varied by changing (a) the interaural pulse duration difference (delta d) for clicks, or (b) the interaural carrier frequency difference (delta f) for tone pips. In either case, the sensitivity to delta t was maximal under diotic presentations and declined with delta d or delta f. The overall sensitivity to delta t was remarkably higher for clicks than for long tone pips. The results indicate that both (a) the extent of binaural spectral overlaps and (b) the structure of acoustic stimuli are important in detecting small interaural time differences of binaural sounds.     

Detection and Lateralization of Binaural Signals

A simple theory, based on the inter-aural difference of the time of stimulation that results from the difference of phase and/or the difference of level at the two ears when an antiphasic signal is presented in noise, is shown to be inadequate to account for the facts of binaural detection and lateralization. The evidence supports the conclusion that both time information and intensity information are furnished to the binaural nervous system, and that the information is utilized differently for detection from what it is for lateralization. Some combinations of time and intensity result in chance performance in lateralization while still permitting a good level of detection. No theory at the present time can account in detail for the phenomena discussed.

Résumé

Une simple théorie qui ne se baserait que sur la différence interauriculaire par rapport au moment de stimulation, différence qui résulte de la variation de phase et/ou d'intensité perçue par les deux oreilles quand le signal antiphasique se présente accompagné de bruit, se révèle comme inadéquate pour rendre compte des faits de détection et de latéralisation binauriculaire. Les évidences confirment la conclusion que l'information-temps et l'information-intensité se présentent toutes deux au système nerveux binauriculaire et que ces informations sont utilisées de façon différente pour la détection et pour la latéralisation. Certaines combinaisons int´eressant le temps et l'intensité ont pour résultat une réaction aléatoire de latéralisation, tout en permettant un niveau satisfaisant de detection. Il n'existe à l'heure actuelle aucune théorie qui puisse expliquer dans le détail les phénomènes en question.     

Acoustic Cues for Sound Source Distance and Azimuth in Rabbits, a Racquetball and a Rigid Spherical Model

There are numerous studies measuring the transfer functions representing signal transformation between a source and each ear canal, i.e., the head-related transfer functions (HRTFs), for various species. However, only a handful of these address the effects of sound source distance on HRTFs. This is the first study of HRTFs in the rabbit where the emphasis is on the effects of sound source distance and azimuth on HRTFs. With the rabbit placed in an anechoic chamber, we made acoustic measurements with miniature microphones placed deep in each ear canal to a sound source at different positions (10–160 cm distance, ±150° azimuth). The sound was a logarithmically swept broadband chirp. For comparisons, we also obtained the HRTFs from a racquetball and a computational model for a rigid sphere. We found that (1) the spectral shape of the HRTF in each ear changed with sound source location; (2) interaural level difference (ILD) increased with decreasing distance and with increasing frequency. Furthermore, ILDs can be substantial even at low frequencies when distance is close; and (3) interaural time difference (ITD) decreased with decreasing distance and generally increased with decreasing frequency. The observations in the rabbit were reproduced, in general, by those in the racquetball, albeit greater in magnitude in the rabbit. In the sphere model, the results were partly similar and partly different than those in the racquetball and the rabbit. These findings refute the common notions that ILD is negligible at low frequencies and that ITD is constant across frequency. These misconceptions became evident when distance-dependent changes were examined.     

Early Components of the Averaged Electroencephalic Response to Monaural and Binaural Stimulation

Early components of the averaged electroencephalic response were recorded in 11 subjects from C_z-A₁, C_z-A₂, C₃-A₁, and C₄-A₂. The early components were examined in response to stimulation of either the left, right, or both ears with 40 dB SL clicks at a rate of 4/sec in awake subjects. Peak latency and peak-to-peak amplitude measures failed to discriminate between stimulus conditions. Left minus right difference scores obtained for each subject showed no consistent relationship to the ear stimulated     

The Role of Interaural Intensity Differences and Time Delay for Signal Detection in Noise

Résumé

Des expériences psycho-acoustiques ont démontré que l'audition binaurale non seulement rend possible la localisation de la source du son mais qu'elle améliore aussi considérablement la relation effective du signal au bruit. Les effets, qui sont écrits en détail, se produisent par des différences interaurales de l'intensité du signal ou par des différences interaurales du temps d'arrivée du signal. L'effet produit par l'un de ces deux paramètres de l'audition binaurale ne se superpose pas sur l'effet de l'autre. Ces effets sont liés à la condition que le signal et le bruit soient cohérents.     

美国特殊教育鉴定与评估对我国的启示

本文运用文献研究法,对美国特殊教育鉴定与评估的历史进行了回顾,对其特殊教育鉴定与评估的现状及特点进行了分析和总结,并对我国特殊教育鉴定与评估工作提出了一些建议,以供参考.     

The phase angle of addition in temporal masking for diotic and dichotic listening conditions

The phase angle, α, between a tonal signal and a tonal masker was varied from 0° to 135° in simultaneous masking, forward masking, and pulsation-threshold paradigms. In all conditions the frequency of the signal and masker was 500 Hz. For forward masking both diotic (MOSO) and dichotic (MOSπ) listening conditions were investigated. Only the dichotic case was studied using the pulsation threshold method. In simultaneous masking, thresholds varied as a function of a in both diotic and dichotic conditions. Thresholds in the diotic conditions were consistently different from those in the dichotic conditions — i.e., there were masking-level differences (MLDs) at most values of α tested. In forward masking and pulsation-threshold, however, thresholds were independent of α in the dichotic conditions; and thresholds were independent of α in the diotic, forward masking conditions. Nevertheless, for forward masking the dichotic thresholds remained below the diotic thresholds, yielding MLDs of 3–6 dB. Thus, in nonsimultaneous masking, there is a clear effect of the interaural signal phase, but not of the masker-signal phase relationship, on signal detectability. These results imply that masker-signal phase information is either not preserved or not used by subjects in nonsimultaneous tone-on-tone masking experiments.     

Tone-in-Noise Detection Using Envelope Cues: Comparison of Signal-Processing-Based and Physiological Models

Tone-in-noise detection tasks with reproducible noise maskers have been used to identify cues that listeners use to detect signals in noisy environments. Previous studies have shown that energy, envelope, and fine-structure cues are significantly correlated to listeners’ performance for detection of a 500-Hz tone in noise. In this study, envelope cues were examined for both diotic and dichotic tone-in-noise detection using both stimulus-based signal processing and physiological models. For stimulus-based envelope cues, a modified envelope slope model was used for the diotic condition and the binaural slope of the interaural envelope difference model for the dichotic condition. Stimulus-based models do not include key nonlinear transformations in the auditory periphery such as compression, rate and dynamic range adaptation, and rate saturation, all of which affect the encoding of the stimulus envelope. For physiological envelope cues, stimuli were passed through models for the auditory nerve (AN), cochlear nucleus, and inferior colliculus (IC). The AN and cochlear nucleus models included appropriate modulation gain, another transformation of the stimulus envelope that is not typically included in stimulus-based models. A model IC cell was simulated with a linear band-pass modulation filter. The average discharge rate and response fluctuations of the model IC cell were compared to human performance. Previous studies have predicted a significant amount of the variance across reproducible noise maskers in listeners’ detection using stimulus-based envelope cues. In this study, a physiological model that includes neural mechanisms that affect encoding of the stimulus envelope predicts a similar amount of the variance in listeners’ performance across noise maskers.     

Masking Level Difference: Another Tool for the Evaluation of Peripheral and Cortical Defects

Masking level difference (MLD) due to binaural unmasking was measured for speech signals (5-word meaningful sentences) masked by broad-band noise.

Tests were carried out in a group of patients with unilateral cerebral lesions of vascular origin and apparently normal pure-tone audiograms (CNS patients), in a control group (normally hearing young adults) and in 5 other groups of patients (conductive symmetrical hearing loss, conductive asymmetrical hearing loss, bilateral presbyacusis, unilateral sudden deafness, Menière's disease). Testing pattern implied three or more S/N ratios in the listening conditions, of S_mN_m, SΔtN_o and S_mN_u (noise correlated), and the speech signal intensity was 70 dB SPL re 20 μPa for the control group, whilst for the pathological cases, speech level intensities were established by means of alternate binaural loudness balance and simultaneous balancing median-plane localization procedures to assess subjective suprathreshold sound image localization at the midline. The results obtained in the control group and in the patients are discussed. With specific reference to the CNS patients, a statistically significant tendency was evident for the binaural condition to produce more MLD when the ear leading in time was ipsilateral to the normal hemisphere. The importance of this and of sensitized speech testing methods in CNS disorders is discussed on the basis of the results obtained in the CNS patients.     

鼓室成形术后的咽鼓管功能检测及意义

目的探讨咽鼓管功能对慢性化脓性中耳炎鼓室成形术后疗效的影响。方法用咽鼓管鼓室-气流动态图(tube-tympanoaerodynamicgraphy,TTAG)法及音响法对53耳鼓室成形术后的咽鼓管功能进行检测,并分析鼓室成形术后咽鼓管功能正常耳与异常耳的鼓膜生长及听力变化情况;并用鼻窦镜观察咽鼓管咽口,分析咽鼓管咽口与咽鼓管功能的关系。结果53耳鼓室成形术后咽鼓管功能正常37耳,其中29耳鼓膜生长良好;咽鼓管功能异常16耳中,有4耳鼓膜生长良好,比较咽鼓管功能正常耳与异常耳鼓膜生长良好耳数,两者有显著性差异(P<0.05);53耳术后有9耳咽鼓管咽口充血肿胀,其中2耳咽鼓管功能正常,7耳咽鼓管功能不良,提示术后咽鼓管功能障碍与咽鼓管咽口病变有关。结论慢性化脓性中耳炎鼓室成形术后咽鼓管功能与疗效关系密切,术后咽鼓管功能障碍可能是导致手术失败的原因之一。     

A Comparative Study of Eya1 and Eya4 Protein Function and Its Implication in Branchio-oto-renal Syndrome and DFNA10

Allele variants of EYA1 and EYA4, two members of the vertebrate Eya gene family, underlie two types of inherited human deafness, branchio-oto-renal (BOR) syndrome and DFNA10, respectively. To clarify how mutations in these two genes and their encoded proteins impact the normal biology of hearing, we completed a number of functional studies using the yeast-two-hybrid system. We verified that bait constructs of the homologous region (Eya1HR and Eya4HR) interact with Six1 prey constructs, although no interaction with Dach1 prey was demonstrable. To compare interaction affinities, we evaluated -galactosidase activity after cotransformation of Eya1HR/Six1 and Eya4HR/Six1 and found that the latter interaction was weaker. By immunofluorescence staining, we showed Eya4HR localization to the cytoplasm. After coexpression of Six1, Eya4HR was translocated to the nucleus. Results with Eya1HR were similar. Translation of mutant constructs (Eya4HR _R564X and Eya1HR _R539X) could not be demonstrated. Using dual Eya-containing constructs (with two wild-type alleles or wild-type and mutant alleles), we confirmed no translation of the mutant allele, even if the mutation was nontruncating. These results are consistent with clinical data and implicate haploinsufficiency as the cause of BOR syndrome and DFNA10.