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The neural circuitry underlying the executive control of auditory spatial attention.

Although a fronto-parietal network has consistently been implicated in the control of visual spatial attention, the network that guides spatial attention in the auditory domain is not yet clearly understood. To investigate this issue, we measured brain activity using functional magnetic resonance imaging while participants performed a cued auditory spatial attention task. We found that cued orienting of auditory spatial attention activated a medial-superior distributed fronto-parietal network. In addition, we found cue-triggered increases of activity in the auditory sensory cortex prior to the occurrence of an auditory target, suggesting that auditory attentional control operates in part by biasing processing in sensory cortex in favor of expected target stimuli. Finally, an exploratory cross-study comparison further indicated several common frontal and parietal regions as being involved in the control of both visual and auditory spatial attention. Thus, the present findings not only reveal the network of brain areas underlying endogenous spatial orienting in the auditory modality, but also suggest that the control of spatial attention in different sensory modalities is enabled in part by some common, supramodal neural mechanisms.

Acoustic Stimulation↗

Binaural interaction in the frog dorsal medullary nucleus.

We have studied binaural and directional processing in cells in the frog dorsal medullary nucleus (DMN) stimulated with dichotic sound (couplers) and free-field sound. We present evidence that already at this stage of central processing the neural directionality is sharpened, probably by binaural interaction. Binaural interaction in the DMN was usually interpreted as inhibition, mostly driven from the contralateral side and dependent on a certain combination of interaural time differences (ITD) and interaural level differences (ILD). In free-field measurements, the strength and timing of the binaural inputs will depend on sound direction as processed by the auditory fibers. Thus, the directionality of DMN cells is caused by both monaural directional cues generated by acoustical coupling of the eardrums and non-tympanic pathways as well as binaural interaction. Most DMN cells show ovoidal directional characteristics and the directionality is sharpened compared to that of auditory nerve fibers. We suggest that the sharpening is due to the inhibitory interactions.

Acoustic Stimulation↗

The mismatch negativity (MMN): towards the optimal paradigm.

OBJECTIVE: Recent studies have shown that the mismatch negativity (MMN), a change-specific component of the event-related potential (ERP), for particular auditory features is degraded in different clinical populations. This suggests that the MMN could, in principle, reflect the whole profile and extent of the central auditory deficit. In the present article, we tested a new MMN paradigm allowing one to obtain MMNs for several auditory attributes in a short time. METHODS: MMN responses to changes in frequency, intensity, duration, location, and to a silent gap occasionally inserted in the middle of a tone were compared between the traditional 'oddball' paradigm (a single type of auditory change in each sequence) and the new paradigm (two versions) in which all the 5 types of changes appeared within the same sequence. RESULTS: The MMNs obtained in the new paradigm were equal in amplitude to those in the traditional MMN paradigm. CONCLUSIONS: We propose a new paradigm that can provide 5 different MMNs in the same time in which usually only one MMN is obtained. The new paradigm enables one to objectively determine the profile of different auditory discrimination abilities within a very short recording time.

Acoustic Stimulation↗

Cortical generators of slow evoked responses elicited by spatial and nonspatial auditory working memory tasks.

OBJECTIVE: Slow evoked responses have been extensively studied using electrophysiological and neuroimaging methods, but there is no consensus regarding their generators. We investigated the generators of the P3 and positive slow wave (PSW) in the evoked responses to probes recorded during auditory working memory tasks to find out whether there is dissociation between functional networks involved in the generation of the P3 and PSW and between spatial and nonspatial auditory processing within this time window. METHODS: Whole-head magneto-(MEG) and electroencephalography (EEG); analysis of MEG data using minimum-norm current estimates. RESULTS: The associative temporal, occipito-temporal and parietal areas contributed to the generation of the slow evoked responses. The temporal source increased while the occipito-temporal source diminished activity during transition from the P3 to PSW. The occipito-temporal generator of the P3 was activated more during the spatial than nonspatial task, and the left temporal generator of the PSW tended to be more strongly activated during the nonspatial task. CONCLUSIONS: These findings indicate that partially distinct functional networks generate the P3 and PSW and provide evidence for segregation of spatial and nonspatial auditory information processing in associative areas beyond the supratemporal auditory cortex. SIGNIFICANCE: The present results support the dual-stream model for auditory information processing.

Acoustic Stimulation↗

Evaluation of multi-attribute auditory discrimination in dyslexia with the mismatch negativity.

OBJECTIVE: Dyslexia is associated with impairments in the phonological system or with more general auditory dysfunctions. We determined the discrimination of 5 sound contrasts (pitch, duration, intensity, location, and the presence of a gap) in dyslexia with the mismatch negativity (MMN). METHODS: We compared MMNs of 9 adult dyslexic and 11 control subjects with a new 5-deviant paradigm which enables one to assess the discrimination of each of these features in 15 min. Also, a control oddball condition with pitch and duration deviants was included. In the new paradigm, all deviant stimuli are presented in the same stimulus block so that the standard stimuli, of which there are 50%, alternate with the deviant stimuli. RESULTS: In the 5-deviant paradigm, a diminished pitch-MMN and an enhanced location-MMN were found in dyslexic individuals. Furthermore, pitch and duration MMNs in this and in the oddball paradigms suggested that smaller MMNs are elicited in the new than oddball paradigm in dyslexic subjects. CONCLUSIONS: Pitch discrimination is impaired in dyslexia. However, location discrimination, not addressed previously with MMN, is enhanced. Furthermore, dyslexic subjects are more impaired in detecting changes in sound streams with than without variation. SIGNIFICANCE: In dyslexia research, the new 5-deviant MMN paradigm is feasible and even more sensitive than the traditional oddball paradigm.

Acoustic Stimulation↗

No direction-specific bimodal facilitation for audiovisual motion detection.

After several decades of unimodal perceptual research, interest is turning increasingly to cross-modal interactions. At a physiological level, the existence of bimodal cells is well documented and it is known that correlated audiovisual input enhances localisation and orienting behaviours. Audiovisual perceptual interactions have also been demonstrated (e.g., the well-known McGurk effect). The present study explores motion perception and asks whether correlated audiovisual motion signals would be better detected than unimodal motions or bimodal motions in opposing directions. Using a dynamic random-dot field with variable motion coherence as a visual stimulus, together with an auditory motion defined by a stereo noise source smoothly translating along a horizontal trajectory, we find that correlated bimodal motion yields only a slight improvement (approximately a square root of two advantage) in detection threshold relative to unimodal detection. The size of this benefit is consistent with a statistical advantage rather than a bimodal facilitation account. Moreover, anticorrelated bimodal motion showed the same modest improvement, again speaking against linear summation but consistent with statistical combination of visual and auditory signals. These findings were replicated in peripheral as well as in central vision, and with translating visual objects as well as with spatially distributed visual motion. The superadditivity observed neurally (especially in deep-layer superior collicular cells), when weak unimodal signals are combined in bimodal cells does not apply to the detection of linear translational motion.

Acoustic Stimulation↗

Sound lateralization in Parkinson's disease.

The symptoms primarily associated with Parkinson's disease (PD) are of a motor and cognitive nature, but sensory deficits may also be involved. Previous studies have reported disturbed spatial perception in visual and tactile tasks. We have investigated whether PD patients show deficits in auditory spatial perception. For this purpose, we employed a simple task involving left/right judgments about dichotic stimuli presented with various interaural time differences (ITD). The acuity of sound lateralization was significantly reduced in PD: the just noticeable difference (JND) in interaural time seen in PD patients was about twice that seen for age-matched healthy controls. We propose that this deficit may be related to a potential role of the basal ganglia in spatial hearing functions, as has been suggested by neurophysiological and neuroanatomical studies on animals.

Acoustic Stimulation↗

Orderly cortical representation of vowel categories presented by multiple exemplars.

This study aimed at determining how the human brain automatically processes phoneme categories irrespective of the large acoustic inter-speaker variability. Subjects were presented with 450 different speech stimuli, equally distributed across the [a], [i], and [u] vowel categories, and each uttered by a different male speaker. A 306-channel magnetoencephalogram (MEG) was used to record N1m, the magnetic counterpart of the N1 component of the auditory event-related potential (ERP). The N1m amplitude and source locations differed between vowel categories. We also found that the spectrum dissimilarities were reproduced in the cortical representations of the large set of the phonemes used in this study: vowels with similar spectral envelopes had closer cortical representations than those whose spectral differences were the largest. Our data further extend the notion of differential cortical representations in response to vowel categories, previously demonstrated by using only one or a few tokens representing each category.

Acoustic Stimulation↗

Sound lateralization in subjects with callosotomy, callosal agenesis, or hemispherectomy.

The question of whether there is a right-hemisphere dominance in the processing of auditory spatial information in human cortex as well as the role of the corpus callosum in spatial hearing functions is still a matter of debate. Here, we approached this issue by investigating two late-callosotomized subjects and one subject with agenesis of the corpus callosum, using a task of sound lateralization with variable interaural time differences. For comparison, three subjects with left or right hemispherectomy were also tested by employing identical methods. Besides a significant reduction in their acuity, subjects with total or partial section of the corpus callosum exhibited a considerable leftward bias of sound lateralization compared to normal controls. No such bias was found in the subject with callosal agenesis, but merely a marginal reduction of general acuity. Also, one subject with complete resection of the left cerebral cortex showed virtually normal performance, whereas another subject with left hemispherectomy and one subject with right hemispherectomy exhibited severe deficits, with almost total loss of sound-lateralization ability. The results obtained in subjects with callosotomy indicate that the integrity of the corpus callosum is not indispensable for preservation of sound-lateralization ability. On the other hand, transcallosal interhemispheric transfer of auditory information obviously plays a significant role in spatial hearing functions that depend on binaural cues. Moreover, these data are compatible with the general view of a dominance of the right cortical hemisphere in auditory space perception.

Acoustic Stimulation↗

Reassessing mechanisms of low-frequency sound localisation.

Interaural time differences (ITDs) are the dominant cues for human localisation of low-frequency sounds. Although a mechanism for ITD processing proposed in 1948 seems applicable to birds, and is consistent with many aspects of the responses found in mammals, recent data suggest that key tenets of the model might need to be reconsidered. The model requires, at every frequency, a distribution of cells with firing rate peaks across all ITD values within the animal's physiological range. The ITD tuning relies on internal delays in the form of a neural delay line. The evidence for such a delay line structure in mammals is not as convincing as it is in birds and, in some small animals the full range of physiological ITDs are not fully represented by peak firing of neurones at every frequency channel. Alternative means of achieving internal delays such as inhibitory inputs or the delays associated with cochlear filtering are being considered.

Animals↗

Neuroscience: what you see and hear is what you get.

The brain receives signals from a variety of sources; for example, visual and auditory signals can both indicate the direction of a stimulus, but with differing precision. A recent study has shed light on the way that the brain combines these signals to achieve the best estimate possible.

Cues↗

Visually induced plasticity of auditory spatial perception in macaques.

When experiencing spatially disparate visual and auditory stimuli, a common percept is that the sound originates from the location of the visual stimulus, an illusion known as the ventriloquism effect. This illusion can persist for tens of minutes, a phenomenon termed the ventriloquism aftereffect. The underlying neuronal mechanisms of this rapidly induced plasticity remain unclear; indeed, it remains untested whether similar multimodal interactions occur in other species. We therefore tested whether macaque monkeys experience the ventriloquism aftereffect similar to the way humans do. The ability of two monkeys to determine which side of the midline a sound was presented from was tested before and after a period of 20-60 min in which the monkeys experienced either spatially identical or spatially disparate auditory and visual stimuli. In agreement with human studies, the monkeys did experience a shift in their auditory spatial perception in the direction of the spatially disparate visual stimulus, and the aftereffect did not transfer across sounds that differed in frequency by two octaves. These results show that macaque monkeys experience the ventriloquism aftereffect similar to the way humans do in all tested respects, indicating that these multimodal interactions are a basic phenomenon of the central nervous system.

Acoustic Stimulation↗

Early- and late-onset blind individuals show supra-normal auditory abilities in far-space.

Blind individuals manifest remarkable abilities in navigating through space despite their lack of vision. They have previously been shown to perform normally or even supra-normally in tasks involving spatial hearing in near space, a region that, however, can be calibrated with sensory-motor feedback. Here we show that blind individuals not only properly map auditory space beyond their peri-personal environment but also demonstrate supra-normal performance when subtle acoustic cues for target location and distance must be used to carry out the task. Moreover, it is generally postulated that such abilities rest in part on cross-modal cortical reorganizations, particularly in the immature brain, where important synaptogenesis is still possible. Nonetheless, we show for the first time that even late-onset blind subjects develop above-normal spatial abilities, suggesting that significant compensation can occur in the adult.

Acoustic Stimulation↗

Directionality of hearing in two CF/FM bats, Pteronotus parnellii and Rhinolophus rouxi.

The head-related transfer function (HRTF) has been measured in two CF/FM bats, Pteronotus parnellii and Rhinolophus rouxi from 575 positions in the frontal hemisphere. P. parnellii showed an increase of the elevation angle of the axis of highest pinna gain with increasing frequency followed by a specific decrease at 75 kHz. Such a drop of elevation angle of the acoustic axis was not seen in R. rouxi. The HRTF further showed a spectral notch dependent on elevation and frequency in P. parnellii, but not in R. rouxi. The functional implications of this difference between both bat species are discussed. Frequencies at maximum pinna gain values did not clearly match the frequencies of the harmonics of the echolocation calls whereas spatial resolution of interaural intensity differences was best in a frequency range that included the higher harmonics of the echolocation calls in both bat species. However, specializations of HRTF patterns matching the exact frequencies of the harmonics of the echolocation calls could not be observed in both bat species.

Acoustic Stimulation↗

Velocity discrimination of auditory image moving in vertical plane.

This research investigated the ability of humans to discriminate changes in velocity of apparent movement of a sound source (as defined by the time required for the sound to traverse a 180 deg arc) in the median vertical plane. Apparent auditory movement was created by successive switching of the loudspeakers situated over the arc. The broadband noise with band width of 0.25-4 kHz (low-pass noise - LP) and of 4-12.5 kHz (high-pass noise - HP) was employed as stimuli. Discrimination thresholds were calculated for reference velocities of 58 and 115 deg/s under four stimuli types: movement of stepped movement of noise for low-pass noise and for high-pass noise and continuous motion for low-pass noise and for high-pass noise. The result showed that discrimination of the velocity depended on the signal frequency spectrum. The differential thresholds for the signals with low pass noise were significantly higher than those for signals with high pass noise (F(1;78) >> 3.96, p < 0.05). Magnitude of the absolute thresholds was the highest when the velocity was 115 deg/s for each of four types. It is interesting to note that the thresholds magnitude depended on the type of signal motion, that is on whether it was continuous or stepped.

Acoustic Stimulation↗

Excitability of auditory brainstem neurons, in vivo, is increased by cyclic-AMP.

Physiological control of auditory neural responses is critical for accurate representation of acoustic information, such as sound source localization and speech perception. Central auditory neural responses are almost certainly regulated by a range of mechanisms, including second messenger systems, such as the cAMP pathway. An increase in spontaneous neural discharge is known to accompany cochlear insults. Here we report that an increase in spontaneous as well as tone-evoked discharge can also be induced by pressure application of forskolin, a pharmacological agent that elevates intracellular cAMP level by activating adenyl cyclase. The forskolin induced increase in superior olivary complex (SOC) brainstem neurons is specific, dose-dependent, and reversible, whereas application of artificial cerebrospinal fluid (aCSF, the vehicle) does not alter activity. Forskolin-application also has a relatively greater effect on spontaneous activity compared to tone evoked responses. Blockade of the hyperpolarization-activated current, Ih, by ZD7288, consistently reversed the effects of forskolin. Based on these findings, we propose that the second messenger, cAMP, can significantly modulate neural excitability and spontaneous discharge in SOC neurons, principally by shifting the activation of Ih channels.

Acoustic Stimulation↗

Attribute capture in the precedence effect for long-duration noise sounds.

Listeners perceptually fuse the direct wave from a sound source with its reflections off nearby surfaces into a single sound image, located at or near the sound source (the precedence effect). This study investigated how a brief gap presented in the middle of either a direct wave or simulated reflection is incorporated into the fused image. For short (<9.5 ms) delays between the direct (leading) and reflected (lagging) waves, no sound was perceived from the direction of the lagging wave. For delays between 10 and 15 ms, both sounds were perceived, but the gap was heard only on the leading side. When the gap was only in the correlated lagging sound at short delays, it also was perceived as occurring on the leading side. Moreover, gap detection thresholds were the same for gaps in the leading and lagging sounds, suggesting that the perception of the gap was not suppressed, but rather incorporated into the leading sound. Finally, scalp event-related potentials were not associated with the precedence effect until the gap occurred. This suggests that cortical mechanisms are engaged to maintain fusion when attributes in direct or reflected waves change.

Adult↗

Time-intensity trading in bilateral congenital aural atresia patients.

In an effort to examine the rules by which information of bilaterally applied bone-conducted signals arising from interaural time differences (ITD) and interaural intensity differences (IID) is combined, data were measured for continuous 500 Hz narrow band noise at 65-70 dB HL in 11 patients with bilateral congenital aural atresia. Time-intensity trading functions were obtained by shifting the sound image towards one side using ITD, and shifting back to a centered sound image by varying the IID in the same ear (auditory midline task). ITD values were varied from -600 to +600 micros at 200 micros steps, where negative values indicate delays to the right ear. The results indicate that time-intensity trading is present in patients with bilateral aural atresia. The gross response properties of time-intensity trading in response to bone-conducted signals were comparable in patients with bilateral aural atresia and normal-hearing subjects, though there was a larger inter-subject variability and higher discrimination thresholds across IIDs in the atresia group. These results suggest that the mature auditory brainstem has a potential to employ binaural cues later in life, although to a restricted degree. A binaural fitting of a bone-conducted hearing aid might optimize binaural hearing and improve sound lateralization, and we recommend now systematically bilateral fitting in aural atresia patients.

Acoustic Stimulation↗