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PubMed2006

A matter of time: internal delays in binaural processing.

As an animal navigates its surroundings, the sounds reaching its two ears change in waveform similarity (interaural correlation) and in time of arrival (interaural time difference, ITD). Humans are exquisitely sensitive to these binaural cues, and it is generally agreed that this sensitivity involves coincidence detectors and internal delays that compensate for external acoustic delays (ITDs). Recent data show an unexpected relationship between the tuning of a neuron to frequency and to ITD, leading to several proposals for sources of internal delay and the neural coding of interaural temporal cues. We review the alternatives, and argue that an understanding of binaural mechanisms requires consideration of sensitivity not only to ITDs, but also to interaural correlation.

Animals↗
PubMed2001

Directing spatial attention towards the illusory location of a ventriloquized sound.

In this study, we examined whether ventriloquism can rearrange external space on which spatial reflexive attention operates. The task was to judge the elevation (up vs down) of auditory targets delivered in the left or the right periphery, taking no account of side of presentation. Targets were preceded by either auditory, visual, or audiovisual cues to that side. Auditory, but not visual cues had an effect on the speed of auditory target discrimination. On the other hand, a ventriloquized cue, consisting of a tone in central location synchronized with a light flash in the periphery, facilitated responses to targets appearing on the same side as the flash. That effect presumably resulted from the attraction of the apparent location of the tone towards the flash, a well-known manifestation of ventriloquism. Ventriloquism thus can reorganize space in which reflexive attention operates.

Adult↗
PubMed2000

The role of GABAergic inhibition on direction-dependent sharpening of frequency tuning in bat inferior collicular neurons.

This study examined the role of GABAergic inhibition on direction-dependent sharpening of frequency tuning curves (FTCs) in bat inferior collicular (IC) neurons under free field stimulation conditions. The minimum threshold (MT) at the neurons best frequency (BF) and the sharpness (Q(10), Q(20), Q(30)) of FTCs of most IC neurons increased as the sound direction changed from contralateral azimuths to ipsilateral azimuths. The application of GABA(A) antagonist, bicuculline, lowered all MTs but the application did not abolish direction-dependent variation in MT. MTs determined during bicuculline application at 40 ipsilateral were still significantly higher than those determined at 40 degrees contralateral (two-tailed paired t-test, P<0.0001). In contrast, although application of bicuculline essentially had no effect on the BFs of IC neurons, it differentially broadened neurons FTCs at different azimuths abolishing the direction-dependent sharpening of frequency tuning (i. e. Q(n) values, two-tailed paired t-test, P<0.01). These data indicate that GABAergic inhibition makes an important contribution to the direction-dependent frequency tuning of most IC neurons.

Animals↗
PubMed2003

The effect of bicuculline application on azimuth-dependent recovery cycle of inferior collicular neurons of the big brown bat, Eptesicus fuscus.

The recovery cycle of auditory neurons is an important neuronal property, which determines a neuron's ability to respond to pairs of sounds presented at short inter-sound intervals. This property is particularly important for bats, which rely upon analysis of returning echoes to extract the information about targets after emission of intense orientation sounds. Because target direction often changes throughout the course of hunting, the changing echo direction may affect the recovery cycle and thus temporal processing of auditory neurons. In this study, we examined the effect of sound azimuth on the recovery cycle of inferior collicular (IC) neurons in the big brown bat, Eptesicus fuscus, under free-field stimulation conditions. Our study showed that the recovery cycle of most IC neurons (42/49, 86%) was longer when determined with sounds delivered at 40 degrees ipsilateral (i40 degrees ) than at 40 degrees contralateral (c40 degrees ) to the recording site. To study the contribution of GABAergic inhibition to sound azimuth-dependent recovery cycle, we compared the recovery cycle of IC neurons determined at two sound azimuths before and during iontophoretic application of bicuculline, an antagonist for GABA(A) receptors. Bicuculline application produced a greater decrease of the recovery cycle of these neurons at i40 degrees than at c40 degrees. As a result, the azimuth-dependent recovery cycle of these neurons was abolished or greatly reduced. Possible mechanisms underlying these observations and biological relevance to bat echolocation are discussed.

Acoustic Stimulation↗
PubMed1998

Age-related changes in auditory spatial properties of the guinea pig superior colliculus.

The map of auditory space located in the deep layers of the guinea pig superior colliculus (SC) is a complex computational representation of the auditory azimuth surrounding the animal. The map undergoes a protracted developmental profile during the first postnatal month and remains plastic until well into adulthood. However, there are no data concerning the state of the collicular auditory space map in much older animals. Multi-unit responses to broadband noise stimuli presented around the azimuthal plane under anechoic conditions were recorded from the deep SC of guinea pigs of a variety of ages, up to 44 months. The data obtained show that the map remains stable up to the age of approximately 36 months. However, after this age, the map shows rapid deterioration and at 42 months, multi-unit responses did not show features consistent with a normal map. It appears that deficits accruing within the central auditory system with increasing age, combine to overcome the ability of the mechanisms of plasticity responsible for space map maintenance to keep pace with the changes, resulting in degraded SC spatial tuning with increasing age.

Aging↗
PubMed1998

Spatial-tuning properties of auditory neurons in the optic tectum of the pigeon.

We studied the auditory neurons in the optic tectum of the unanesthetized pigeon, using single-unit recordings and acoustic free-field stimulation. Most units showed spatial tuning, with best areas located in the contralateral hemifield. All units responded also to visual stimuli, the auditory best areas being in rough alignment with visual receptive fields.

Acoustic Stimulation↗
PubMed1999

Changes in occipital cortex activity in early blind humans using a sensory substitution device.

The purpose of this study was to investigate the neural networks involved when using an ultrasonic echolocation device, which is a substitution prosthesis for blindness through audition. Using positron emission tomography with fluorodeoxyglucose, regional brain glucose metabolism was measured in the occipital cortex of early blind subjects and blindfolded controls who were trained to use this prosthesis. All subjects were studied under two different activation conditions: (i) during an auditory control task, (ii) using the ultrasonic echolocation device in a spatial distance and direction evaluation task. Results showed that the abnormally high metabolism already observed in early blind occipital cortex at rest [C. Veraart, A.G. De Volder, M.C. Wanet-Defalque, A. Bol, C. Michel, A.M. Goffinet, Glucose utilization in human visual cortex is, respectively elevated and decreased in early versus late blindness, Brain Res. 510 (1990) 115-121.] was also present during the control task and showed a trend to further increase during the use of the ultrasonic echolocation device. This specific difference in occipital cortex activity between the two tasks was not observed in control subjects. The metabolic recruitment of the occipital cortex in early blind subjects using a substitution prosthesis could reflect a concurrent stimulation of functional cross-modal sensory connections. Given the unfamiliarity of the task, it could be interpreted as a prolonged plasticity in the occipital cortex early deprived of visual afferences.

Acoustic Stimulation↗
PubMed2001

Auditory and visual objects.

Notions of objecthood have traditionally been cast in visuocentric terminology. As a result, theories of auditory and cross-modal perception have focused more on the differences between modalities than on the similarities. In this paper we re-examine the concept of an object in a way that overcomes the limitations of the traditional perspective. We propose a new, cross-modal conception of objecthood which focuses on the similarities between modalities instead of the differences. Further, we propose that the auditory system might consist of two parallel streams of processing (the 'what' and 'where' subsystems) in a manner analogous to current conceptions of the visual system. We suggest that the 'what' subsystems in each modality are concerned with objecthood. Finally, we present evidence for - and elaborate on - the hypothesis that the auditory 'where' subsystem is in the service of the visual-motor 'where' subsystem.

Auditory Perception↗
PubMed2002

Multisensory spatial representations in eye-centered coordinates for reaching.

Humans can reach for objects with their hands whether the objects are seen, heard or touched. Thus, the position of objects is recoded in a joint-centered frame of reference regardless of the sensory modality involved. Our study indicates that this frame of reference is not the only one shared across sensory modalities. The location of reaching targets is also encoded in eye-centered coordinates, whether the targets are visual, auditory, proprioceptive or imaginary. Furthermore, the remembered eye-centered location is updated after each eye and head movement. This is quite surprising since, in principle, a reaching motor command can be computed from any non-visual modality without ever recovering the eye-centered location of the stimulus. This finding may reflect the predominant role of vision in human spatial perception.

Adult↗
PubMed2004

Auditory neglect: what and where in auditory space.

A sound that we hear in a natural setting allows us to identify the sound source and to localise it in space. Several lines of evidence indicate that the two aspects are processed in anatomically distinct cortical networks. Auditory areas that are part of the What or Where processing streams have been identified recently in man and in non-human primates. Comparison between anatomical and activation studies suggests that processing within either stream can be modulated by specific attentional factors. Attending to auditory events can be affected in neglect. Bisiach et al. (1984) described systematic directional errors to the ipsilesional space, which was considered a manifestation of hemispatial neglect and interpreted as a disruption of the neural network providing the internal representation of egocentric space. The other manifestation of auditory neglect is contralesional extinction in dichotic listening condition (Heilman and Valenstein, 1972). Recently two types of auditory neglect have been described, one corresponding to a primarily attentional deficit associated with basal ganglia lesions and the other to distortions of auditory space representations associated with parieto-prefrontal lesions (Bellmann et al., 2001). Based on studies of sound detection and sound recognition following hemispheric lesions we argue that the two types of neglect correspond to disturbed processing in either the What or the Where stream.

Attention↗
PubMed2004

Auditory deficits in visuospatial neglect patients.

Since the pioneering experimental work of Bisiach et al. (1984) on deficits in sound localisation associated with unilateral brain lesions and visual neglect, a number of systematic investigations have examined auditory processing in visuospatial neglect patients. Evidence from a variety of experimental paradigms has revealed some auditory deficits in detection and identification tasks, during bilateral stimulation; plus localisation deficits for single sounds. These deficits emerge predominantly for contra-lesional sounds, although some auditory disturbances applying to both contra- and ipsilesional sounds have also been documented. Here we review evidence suggesting that some of these auditory deficits arise in relatively high-level stages of spatial processing. In addition, we present new analyses showing that auditory deficits in identification and localisation tasks often correlate with clinical measures of visual neglect, across a variety of different studies and tasks. This empirical relation suggests that a disturbance of multisensory spatial processing may often account for the joint auditory and visual spatial deficits in neglect patients, although rarer dissociations between the modalities should also be considered.

Auditory Perception↗
PubMed2002

Environmental sound recognition after unilateral subcortical lesions.

Nonverbal environmental sound recognition was investigated in 24 subjects with unilateral subcortical lesions and 20 age-matched normal controls. All patients incurred putaminal hemorrhage at least three months before examination, and had a cystic lesion under the insula on CT or MRI at the time of evaluation. A mild impairment was found in association with extensive damage to the lateral and ventral portions of the putamen and the adjacent white matter in either the right or left hemisphere. Degree of impairment and type of error did not differ significantly between the sides of lesions. These observations and a review of literature suggest that the impairment of environmental sound recognition may arise with a unilateral subcortical lesion disrupting the geniculo-auditory association cortex projection fibers.

Agnosia↗
PubMed1999

Neglect without extinction.

A patient, AB, is reported who showed clear signs of neglect but no extinction (N+ E-). Several hypotheses proposed to account for this dissociation were put to the test. The postulated association between motor neglect and extinction did not hold good, nor did the possibility that the N+ E- dissociation may be traced back to the difference in test requirements and therefore observed only in patients with object-centred neglect. Likewise, manipulating the physical features of the stimuli (relative size, exposure time, presentation synchrony) did not elicit extinction. However, when the task demands were modified by asking the patient to perform a further spatial analysis of the stimuli, rather than simply detect them, extinction emerged. Since AB performed well on several neglect tasks requiring parallel processing, while failing all tasks calling for serial processing, the hypothesis is put forward that AB's N+ E- dissociation could be interpreted within the parallel/serial distinction framework.

Aged↗
PubMed2005

Staring in the eye of auditory neglect: comments on 'gaze direction modulates auditory spatial deficits in stroke patients with neglect'.

In the current issue of Cortex, Pavani and colleagues show that directing gaze toward the contralesional hemifield significantly diminishes the auditory deficits associated with unilateral neglect. The authors suggest that this beneficial effect of gaze direction may arise due to the recruitment of crossmodal attentional links between audition and vision. A complementary interpretation of these findings is that directing gaze toward neglected sound sources encourages the recoding of auditory spatial location in a relatively preserved frame of reference, and that it is through this coordinate transformation process that awareness of auditory stimuli may be regained. Rehabilitation strategies aimed at recoding stimuli within relatively preserved reference frames may be a useful step forward in managing unilateral neglect.

Attention↗
PubMed2005

Gaze direction modulates auditory spatial deficits in stroke patients with neglect.

We investigated the effects of eye position on auditory spatial deficits in four patients with left neglect and right-hemisphere damage, using three blocked gaze directions (35 degrees to the right, central, or 35 degrees to the left), while preventing any head-movement to ensure that initial auditory inputs remained constant regardless of eye-in-orbit position. The auditory task required speeded discrimination of sound elevation, with patients moving a central lever up or down according to the vertical position of a peripheral target sound, regardless of its side (left or right). Replicating previous auditory research, the patients' vertical discrimination performance was worse for auditory targets on the contralesional (left) versus the ipsilesional side, indicating neglect-related auditory deficits on this task. Critically, while this worse performance for left than right auditory targets was present (for both reaction times and errors) when gaze was directed centrally or rightwards, it was considerably reduced when gaze was directed leftwards. These results demonstrate that lateral gaze-direction can modulate neglect-related auditory spatial deficits, even though eye-position did not alter the initial auditory inputs. This outcome may relate to audio-visual links in spatial orienting and potentially some retinocentric influences on perceived sound location, although the latter alone could not explain all our results. Such findings might involve multisensory brain structures in which responses to sounds are modulated by eye-in-orbit position.

Attention↗
PubMed1986

Manual pointing to auditory targets: performances of right versus left handed subjects.

In order to test the existence of an hemispheric asymmetry at a basic level of spatial information processing, six right handed and six left handed normal subjects were submitted to a manual pointing task to auditory targets. Results showed a shift in perceived target position according to the hand used for pointing, a striking asymmetry between the two auditory hemispaces reflected in both groups in directional error and dispersion.

Adult↗
PubMed1984

The directionality of the frog ear described by a mechanical model.

The frog ear can be modelled as two coupled eardrums with an additional sound pathway through the mouth cavity. In a previous version of this model, with realistic parameters, we were able to account very well for empirical measurements of the eardrum vibration under free- and closed-field acoustic conditions. This earlier model does not, however, adequately predict the directional characteristics of the frog ear as determined empirically. In this paper we present a modified model which will account for the directionality together with the free- and closed-field frequency responses, and which is also consistent with anatomical considerations.

Animals↗
PubMed2000

Modulation of slow brain potentials by working memory load in spatial and nonspatial auditory tasks.

Slow event-related brain potentials were recorded from the human scalp during spatial and nonspatial auditory delayed matching-to-sample and n-back tasks to find out whether there are differences in the distribution of slow potentials during the retention of audiospatial and pitch information. The performance of both the location and pitch tasks produced slow potentials during the delay phase of the memory tasks. The delay-related slow potential was modulated by the amount of information to be processed during the tasks at the parietal-occipital sites. The distribution of mnemonic modulation was, however, not different between the tasks. The results suggest that there is integration of auditory information processing in the neuronal networks engaged in mnemonic processing of pitch and location.

Adolescent↗