Sound frequency affects the auditory motion-onset response in humans.

The current study examines the modulation of the motion-onset response based on the frequency-range of sound stimuli. Delayed motion-onset and stationary stimuli were presented in a free-field by sequentially activating loudspeakers on an azimuthal plane keeping the natural percept of externalized sound presentation. The sounds were presented in low- or high-frequency ranges and had different motion direction within each hemifield. Difference waves were calculated by contrasting the moving and stationary sounds to isolate the motion-onset responses. Analyses carried out at the peak amplitudes and latencies on the difference waves showed that the early part of the motion response (cN1) was modulated by the frequency range of the sounds with stronger amplitudes elicited by stimuli with high frequency range. Subsequent post hoc analysis of the normalized amplitude of the motion response confirmed the previous finding by excluding the possibility that the frequency range had an overall effect on the waveform, and showing that this effect was instead limited to the motion response. These results support the idea of a modular organization of the motion-onset response with the processing of primary sound motion characteristics being reflected in the early part of the response. Also, the article highlights the importance of specificity in auditory stimulus design.

Auditory processing disorders with and without central auditory discrimination deficits.

Auditory processing disorder (APD) is defined as a processing deficit in the auditory modality and spans multiple processes. To date, APD diagnosis is mostly based on the utilization of speech material. Adequate nonspeech tests that allow differentiation between an actual central hearing disorder and related disorders such as specific language impairments are still not adequately available. In the present study, 84 children between 6 and 17 years of age (clinical group), referred to three audiological centers for APD diagnosis, were evaluated with standard audiological tests and additional auditory discrimination tests. Latter tests assessed the processing of basic acoustic features at two different stages of the ascending central auditory system: (1) auditory brainstem processing was evaluated by quantifying interaural frequency, level, and signal duration discrimination (interaural tests). (2) Diencephalic/telencephalic processing was assessed by varying the same acoustic parameters (plus signals with sinusoidal amplitude modulation), but presenting the test signals in conjunction with noise pulses to the contralateral ear (dichotic(signal/noise) tests). Data of children in the clinical group were referenced to normative data obtained from more than 300 normally developing healthy school children. The results in the audiological and the discrimination tests diverged widely. Of the 39 children that were diagnosed with APD in the audiological clinic, 30 had deficits in auditory performance. Even more alarming was the fact that of the 45 children with a negative APD diagnosis, 32 showed clear signs of a central hearing deficit. Based on these results, we suggest revising current diagnostic procedure to evaluate APD in order to more clearly differentiate between central auditory processing deficits and higher-order (cognitive and/or language) processing deficits.

Free-field study on auditory localization and discrimination performance in older adults.

Localization accuracy and acuity for low- (0.375-0.75 kHz; LN) and high-frequency (2.25-4.5 kHz; HN) noise bands were examined in young (20-29 years) and older adults (65-83 years) in the acoustic free-field. A pointing task was applied to quantify accuracy, while acuity was inferred from minimum audible angle (MAA) thresholds measured with an adaptive 3-alternative forced-choice procedure. Accuracy decreased with laterality and age. From young to older adults, the accuracy declined by up to 23 % for the low-frequency noise band across all lateralities. The mean age effect was even more pronounced on MAA thresholds. Thus, age was a strong predictor for MAA thresholds for both LN and HN bands. There was no significant correlation between hearing status and localization performance. These results suggest that central auditory processing of space declines with age and is mainly driven by age-related changes in the processing of binaural cues (interaural time difference and interaural intensity difference) and not directly induced by peripheral hearing loss. We conclude that the representation of the location of sound sources becomes blurred with age as a consequence of declined temporal processing, the effect of which becomes particularly evident for MAA thresholds, where two closely adjoining sound sources have to be separated. While localization accuracy and MAA were not correlated in older adults, only a weak correlation was found in young adults. These results point to an employment of different processing strategies for localization accuracy and acuity.

Differences in evoked potentials during the active processing of sound location and motion.

Difference in the processing of motion and static sounds in the human cortex was studied by electroencephalography with subjects performing an active discrimination task. Sound bursts were presented in the acoustic free-field between 47° to the left and 47° to the right under three different stimulus conditions: (i) static, (ii) leftward motion, and (iii) rightward motion. In an active oddball design, subject was asked to detect target stimuli which were randomly embedded within a stream of frequently occurring non-target events (i.e. 'standards') and rare non-target stimuli (i.e. 'deviants'). The respective acoustic stimuli were presented in blocks with each stimulus type presented in either of three stimulus conditions: as target, as non-target, or as standard. The analysis focussed on the event related potentials evoked by the different stimulus types under the respective standard condition. Same as in previous studies, all three different acoustic stimuli elicited the obligatory P1/N1/P2 complex in the range of 50-200 ms. However, comparisons of ERPs elicited by static stimuli and both kinds of motion stimuli yielded differences as early as ~100 ms after stimulus-onset, i.e. at the level of the exogenous N1 and P2 components. Differences in signal amplitudes were also found in a time window 300-400 ms ('d300-400 ms' component in 'motion-minus-static' difference wave). For motion stimuli, the N1 amplitudes were larger over the hemisphere contralateral to the origin of motion, while for static stimuli N1 amplitudes over both hemispheres were in the same range. Contrary to the N1 component, the ERP in the 'd300-400 ms' time period showed stronger responses over the hemisphere contralateral to motion termination, with the static stimuli again yielding equal bilateral amplitudes. For the P2 component a motion-specific effect with larger signal amplitudes over the left hemisphere was found compared to static stimuli. The presently documented N1 components comply with the results of previous studies on auditory space processing and suggest a contralateral dominance during the process of cortical integration of spatial acoustic information. Additionally, the cortical activity in the 'd300-400 ms' time period indicates, that in addition to the motion origin (as reflected by the N1) also the direction of motion (leftward/ rightward motion) or rather motion termination is cortically encoded. These electrophysiological results are in accordance with the 'snap shot' hypothesis, assuming that auditory motion processing is not based on a genuine motion-sensitive system, but rather on a comparison process of spatial positions of motion origin (onset) and motion termination (offset). Still, specificities of the present P2 component provides evidence for additional motion-specific processes possibly associated with the evaluation of motion-specific attributes, i.e. motion direction and/or velocity which is preponderant in the left hemisphere.

Purinergic modulation of neuronal activity in developing auditory brainstem.

In the developing nervous system, spontaneous neuronal activity arises independently of experience or any environmental input. This activity may play a major role in axonal pathfinding, refinement of topographic maps, dendritic morphogenesis, and the segregation of axonal terminal arbors. In the auditory system, endogenously released ATP in the cochlea activates inner hair cells to trigger bursts of action potentials (APs), which are transferred to the central auditory system. Here we show the modulatory role of purinergic signaling beyond the cochlea, i.e., the developmentally regulated and cell-type-specific depolarizing effects on auditory brainstem neurons of Mongolian gerbil. We assessed the effects of P2X receptors (P2XRs) on neuronal excitability from prehearing to early stages of auditory signal processing. Our results demonstrate that in neurons expressing P2XRs, extracellular ATP can evoke APs in sync with Ca(2+) signals. In cochlear nucleus (CN) bushy cells, ATP increases spontaneous and also acoustically evoked activity in vivo, but these effects diminish with maturity. Moreover, ATP not only augmented glutamate-driven firing, but it also evoked APs in the absence of glutamatergic transmission. In vivo recordings also revealed that endogenously released ATP in the CN contributes to neuronal firing activity by facilitating AP generation and prolonging AP duration. Given the enhancing effect of ATP on AP firing and confinement of P2XRs to certain auditory brainstem nuclei, and to distinct neurons within these nuclei, it is conceivable that purinergic signaling plays a specific role in the development of neuronal brainstem circuits.

Low-voltage activated Kv1.1 subunits are crucial for the processing of sound source location in the lateral superior olive in mice.

Voltage-gated potassium (Kv) channels containing Kv1.1 subunits are strongly expressed in neurons that fire temporally precise action potentials (APs). In the auditory system, AP timing is used to localize sound sources by integrating interaural differences in time (ITD) and intensity (IID) using sound arriving at both cochleae. In mammals, the first nucleus to encode IIDs is the lateral superior olive (LSO), which integrates excitation from the ipsilateral ventral cochlear nucleus and contralateral inhibition mediated via the medial nucleus of the trapezoid body. Previously we reported that neurons in this pathway show reduced firing rates, longer latencies and increased jitter in Kv1.1 knockout (Kcna1−/−) mice. Here, we investigate whether these differences have direct impact on IID processing by LSO neurons. Single-unit recordings were made from LSO neurons of wild-type (Kcna1+/+) and from Kcna1−/− mice. IID functions were measured to evaluate genotype-specific changes in integrating excitatory and inhibitory inputs. In Kcna1+/+ mice, IID sensitivity ranged from +27 dB (excitatory ear more intense) to −20 dB (inhibitory ear more intense), thus covering the physiologically relevant range of IIDs. However, the distribution of IID functions in Kcna1−/− mice was skewed towards positive IIDs, favouring ipsilateral sound positions. Our computational model revealed that the reduced performance of IID encoding in the LSO of Kcna1−/− mice is mainly caused by a decrease in temporal fidelity along the inhibitory pathway. These results imply a fundamental role for Kv1.1 in temporal integration of excitation and inhibition during sound source localization.

Spatial selective attention in a complex auditory environment such as polyphonic music.

To investigate the influence of spatial information in auditory scene analysis, polyphonic music (three parts in different timbres) was composed and presented in free field. Each part contained large falling interval jumps in the melody and the task of subjects was to detect these events in one part ("target part") while ignoring the other parts. All parts were either presented from the same location (0 degrees; overlap condition) or from different locations (-28 degrees, 0 degrees, and 28 degrees or -56 degrees, 0 degrees, and 56 degrees in the azimuthal plane), with the target part being presented either at 0 degrees or at one of the right-sided locations. Results showed that spatial separation of 28 degrees was sufficient for a significant improvement in target detection (i.e., in the detection of large interval jumps) compared to the overlap condition, irrespective of the position (frontal or right) of the target part. A larger spatial separation of the parts resulted in further improvements only if the target part was lateralized. These data support the notion of improvement in the suppression of interfering signals with spatial sound source separation. Additionally, the data show that the position of the relevant sound source influences auditory performance.

Hemispheric specialization during discrimination of sound sources reflected by MMN.

The present study reports the hemispheric specificity of spatial auditory processing in 15 healthy subjects by measuring location mismatch negativity (MMN) under free field stimulation. The aim was to decide between the partly inconsistent hypotheses of contralateral and/or right-hemispheric dominance in spatial processing in the auditory cortex. The laterality of deviant-standard positions were systematically varied covering the whole of the frontal hemifield from 90 degrees left to 90 degrees right, while the spatial separation of deviant and standard were fixed at 17 degrees. This enabled the evaluation of the specific location-MMNs relating to distinct cortical processing of acoustic space. The inter-hemispheric comparison of the amplitudes of MMNs showed that spatial deviation towards the periphery at -/+17 degrees (relative to 0 degrees -standard) and at -/+90 degrees (relative to -/+73 degrees -standard) elicited a salient contralateral activation. In contrast, positional changes towards front at -/+56 degrees -positions (relative to -/+73 degrees -standard) resulted equal bilateral MMNs. Further, MMN latencies became longer with increasing laterality of respective deviant-standard pairs. Thus, the present study suggests a contralateral pre-attentive cortical processing of acoustic space information in the free field. The direction of positional changes ('towards periphery' vs. 'towards front') seems to augment or reduce this contralateral effect. The sound source discrimination performance across space is mirrored by the location-MMN latency.

Orienting asymmetries and lateralized processing of sounds in humans.

BACKGROUND: Lateralized processing of speech is a well studied phenomenon in humans. Both anatomical and neurophysiological studies support the view that nonhuman primates and other animal species also reveal hemispheric differences in areas involved in sound processing. In recent years, an increasing number of studies on a range of taxa have employed an orienting paradigm to investigate lateralized acoustic processing. In this paradigm, sounds are played directly from behind and the direction of turn is recorded. This assay rests on the assumption that a hemispheric asymmetry in processing is coupled to an orienting bias towards the contralateral side. To examine this largely untested assumption, speech stimuli as well as artificial sounds were presented to 224 right-handed human subjects shopping in supermarkets in Germany and in the UK. To verify the lateralized processing of the speech stimuli, we additionally assessed the brain activation in response to presentation of the different stimuli using functional magnetic resonance imaging (fMRI). RESULTS: In the naturalistic behavioural experiments, there was no difference in orienting behaviour in relation to the stimulus material (speech, artificial sounds). Contrary to our predictions, subjects revealed a significant left bias, irrespective of the sound category. This left bias was slightly but not significantly stronger in German subjects. The fMRI experiments confirmed that the speech stimuli evoked a significant left lateralized activation in BA44 compared to the artificial sounds. CONCLUSION: These findings suggest that in adult humans, orienting biases are not necessarily coupled with lateralized processing of acoustic stimuli. Our results -- as well as the inconsistent orienting biases found in different animal species -- suggest that the orienting assay should be used with caution. Apparently, attention biases, experience, and experimental conditions may all affect head turning responses. Because of the complexity of the interaction of factors, the use of the orienting assay to determine lateralized processing of sound stimuli is discouraged.

The medial nucleus of the trapezoid body in rat: spectral and temporal properties vary with anatomical location of the units.

The medial nucleus of the trapezoid body (MNTB) is a distinct nucleus in the superior olivary complex that transforms excitatory input from the cochlear nucleus into a widespread inhibitory output to distinct auditory brainstem nuclei. Few studies have dealt with the response properties of MNTB neurons to sound stimulation using in vivo preparations. In order to have a better understanding of the functional significance of the MNTB in auditory processing we report the basic temporal and spectral response properties of its principal cells using single-unit extracellular recordings to acoustic stimulation with pure tones and amplitude-modulated stimuli in the rat. Ninety-seven per cent of units showed V-shaped frequency response areas. Rate level functions were mainly saturating (51%) or monotonic (45%) at high intensities. Post-stimulus time histograms typically were characterised as primary-like with notch (59%) or primary-like (33%). Units showed good phase-locking to sinusoidally amplitude-modulated signals with vector strength VS values up to 0.87. Modulation transfer functions had low-pass shapes at near-threshold levels, with cut-off frequencies ranging from 370 to 1270 Hz. Exploration of the relationship between the temporal and spectral properties and the location of the units in the MNTB yielded characteristic frequency (CF)-dependent response properties (latency, Q(10) and cut-off frequency) following a medio-lateral gradient, and CF-independent response features (maximum firing rate) following a dorso-ventral gradient.

Differential processing of terminal tone parts within structured and non-structured tones.

Recent studies utilizing the mismatch negativity (MMN) event-related potential (ERP) revealed that when a repetitive sequence of sinusoidal tones is presented, the occasional insertion of a short deviation into some of the tones leads to the elicitation of an MMN only if it occurs during the initial 300 ms, but not beyond. In contrast, deviations occurring in speech sounds elicit MMN even beyond 300 ms. We conducted two experiments to resolve this conflict. We hypothesised that an additional transient within an otherwise unstructured tone may overcome this limitation. First, we tested for MMN to a deviance at the terminal part of a 650 ms tone which did or did not contain a gap. Only when the tone included the gap, MMN was obtained. Second, we compared the gap condition with two noise conditions, in which the gap was replaced by modulated white noise. The noise conditions differed with respect to the saliency of the perceived interruption of the tone. In all three conditions, MMN was elicited. These results demonstrate that structuring a sinusoidal tone by a gap or a noise interval is sufficient to regain MMN. It is suggested that the introduction of an additional transient triggers a new integration window overcoming the temporal constraints of automatic tone representation. This resolves the seeming contradiction between MMN studies using tonal and speech sounds.

Hemispheric asymmetry for auditory processing in the human auditory brain stem, thalamus, and cortex.

We report evidence for a context- and not stimulus-dependent functional asymmetry in the left and right human auditory midbrain, thalamus, and cortex in response to monaural sounds. Neural activity elicited by left- and right-ear stimulation was measured simultaneously in the cochlear nuclei, inferior colliculi (ICs), medial geniculate bodies (MGBs), and auditory cortices (ACs) in 2 functional magnetic resonance imaging experiments. In experiment 1, pulsed noise was presented monaurally to either ear, or binaurally, simulating a moving sound source. In experiment 2, only monaural sounds were presented. The results show a modulation of the neural responses to monaural sounds by the presence of binaural sounds at a time scale of tens of seconds: In the absence of binaural stimulation, the left and right ICs, MGBs, and ACs responded stronger to stimulation of the contralateral ear. When blocks of binaural stimuli were interspersed in the sound sequence, the contralateral preference vanished in those structures in the right hemisphere. The resulting hemispheric asymmetry was similar to the asymmetry demonstrated for spatial sound processing. Taken together, the data demonstrate that functional asymmetries in auditory processing are modulated by context. The observed long time constant suggests that this effect results from a "top-down" mechanism.

Hemispheric asymmetry for spectral and temporal processing in the human antero-lateral auditory belt cortex.

The present study investigates the acoustic basis of the hemispheric asymmetry for the processing of speech and music. Experiments on this question ideally involve stimuli that are perceptually unrelated to speech and music, but contain acoustic characteristics of both. Stimuli in previous studies were derived from speech samples or tonal sequences. Here we introduce a new class of noise-like sound stimuli with no resemblance of speech or music that permit independent parametric variation of spectral and temporal acoustic complexity. Using these stimuli in a functional MRI experiment, we test the hypothesis of a hemispheric asymmetry for the processing of spectral and temporal sound structure by seeking cortical areas in which the blood oxygen level dependent (BOLD) signal covaries with the number of simultaneous spectral components (spectral complexity) or the temporal modulation rate (temporal complexity) of the stimuli. BOLD-responses from the left and right Heschl's gyrus (HG) and part of the right superior temporal gyrus covaried with the spectral parameter, whereas covariation analysis for the temporal parameter highlighted an area on the left superior temporal gyrus. The portion of superior temporal gyrus in which asymmetrical responses are apparent corresponds to the antero-lateral auditory belt cortex, which has been implicated with spectral integration in animal studies. Our results support a similar function of the anterior auditory belt in humans. The findings indicate that asymmetrical processing of complex sounds in the cerebral hemispheres does not depend on semantic, but rather on acoustic stimulus characteristics.

Hierarchical processing of sound location and motion in the human brainstem and planum temporale.

Horizontal sound localization relies on the extraction of binaural acoustic cues by integration of the signals from the two ears at the level of the brainstem. The present experiment was aimed at detecting the sites of binaural integration in the human brainstem using functional magnetic resonance imaging and a binaural difference paradigm, in which the responses to binaural sounds were compared with the sum of the responses to the corresponding monaural sounds. The experiment also included a moving sound condition, which was contrasted against a spectrally and energetically matched stationary sound condition to assess which of the structures that are involved in general binaural processing are specifically specialized in motion processing. The binaural difference contrast revealed a substantial binaural response suppression in the inferior colliculus in the midbrain, the medial geniculate body in the thalamus and the primary auditory cortex. The effect appears to reflect an actual reduction of the underlying activity, probably brought about by binaural inhibition or refractoriness at the level of the superior olivary complex. Whereas all structures up to and including the primary auditory cortex were activated as strongly by the stationary as by the moving sounds, non-primary auditory fields in the planum temporale responded selectively to the moving sounds. These results suggest a hierarchical organization of auditory spatial processing in which the general analysis of binaural information begins as early as the brainstem, while the representation of dynamic binaural cues relies on non-primary auditory fields in the planum temporale.

Spectral and temporal processing in the human auditory cortex--revisited.

We use novel noise-like sound stimuli to identify cortical areas in which the functional magnetic resonance signal covaries with spectral and temporal acoustic complexity. The results support a model of hemispheric functional asymmetry for fine-grained spectral and fast temporal processing.