Effect of prism adaptation on left dichotic listening deficit in neglect patients: glasses to hear better?

Unilateral neglect is a disabling syndrome frequently observed following right hemisphere brain damage. Symptoms range from visuo-motor impairments through to deficient visuo-spatial imagery, but impairment can also affect the auditory modality. A short period of adaptation to a rightward prismatic shift of the visual field is known to improve a wide range of hemispatial neglect symptoms, including visuo-manual tasks, mental imagery, postural imbalance, visuo-verbal measures and number bisection. The aim of the present study was to assess whether the beneficial effects of prism adaptation may generalize to auditory manifestations of neglect. Auditory extinction, whose clinical manifestations are independent of the sensory modalities engaged in visuo-manual adaptation, was examined in neglect patients before and after prism adaptation. Two separate groups of neglect patients (all of whom exhibited left auditory extinction) underwent prism adaptation: one group (n = 6) received a classical prism treatment ('Prism' group), the other group (n = 6) was submitted to the same procedure, but wore neutral glasses creating no optical shift (placebo 'Control' group). Auditory extinction was assessed by means of a dichotic listening task performed three times: prior to prism exposure (pre-test), upon prism removal (0 h post-test) and 2 h later (2 h post-test). The total number of correct responses, the lateralization index (detection asymmetry between the two ears) and the number of left-right fusion errors were analysed. Our results demonstrate that prism adaptation can improve left auditory extinction, thus revealing transfer of benefit to a sensory modality that is orthogonal to the visual, proprioceptive and motor modalities directly implicated in the visuo-motor adaptive process. The observed benefit was specific to the detection asymmetry between the two ears and did not affect the total number of responses. This indicates a specific effect of prism adaptation on lateralized processes rather than on general arousal. Our results suggest that the effects of prism adaptation can extend to unexposed sensory systems. The bottom-up approach of visuo-motor adaptation appears to interact with higher order brain functions related to multisensory integration and can have beneficial effects on sensory processing in different modalities. These findings should stimulate the development of therapeutic approaches aimed at bypassing the affected sensory processing modality by adapting other sensory modalities.

Cross-modal processing of auditory-visual stimuli in a no-task paradigm: a topographic event-related potential study.

OBJECTIVE: The aim of this study was to investigate in healthy adults the electrophysiological correlates of auditory-visual interactions involved in perception of bimodal events in a no-task paradigm. METHODS: Event-related potentials were recorded in response to unimodal auditory (A), unimodal visual (V) and bimodal (AV) stimuli. Cross-modal interactions were estimated using the additive [AV-(A+V)] model. RESULTS: The spatio-temporal analysis of ERPs and scalp current densities revealed several interaction patterns occurring at both early and late stages of sensory cortical processing: (1) amplitude decrease of the unimodal auditory N1 wave as early as 55ms, (2) amplitude increase of the unimodal auditory P2 wave from 150 to 195ms concomitant with new neural activity over the right fronto-temporal region, and (3) amplitude increase of the late unimodal visual response within 245-350ms post-stimulus. CONCLUSIONS: Our study provides evidence that several patterns of cross-modal interactions can be generated even if no task is required from subjects. SIGNIFICANCE: The paradigm used here can thus be utilized for studying the maturation of the cross-modal processes in young children and in children with pathological development.

Electrophysiological correlates of age and gender perception on human faces.

In a previous experiment using scalp event-related potentials (ERPs), we have described the neuroelectric activities associated with the processing of gender information on human faces (Mouchetant-Rostaing, Giard, Bentin, Aguera, & Pernier, 2000). Here we extend this study by examining the processing of age on faces using a similar experimental paradigm, and we compare age and gender processing. In one session, faces were of the same gender (women) and of one age range (young or old), to reduce gender and age processing. In a second session, faces of young and old women were randomly intermixed but age was irrelevant for the task, hence, age discrimination, if any, was assumed to be incidental. In the third and fourth sessions, faces had to be explicitly categorized according to their age or gender, respectively (intentional discrimination). Neither age nor gender processing affected the occipito-temporal N170 component often associated with the detection of physiognomic features and global structural encoding of faces. Rather, the three age and gender discrimination conditions induced similar fronto-central activities around 145-185 msec. In our previous experiment, this ERP pattern was also found for implicit and explicit categorization of gender from faces but not in a control condition manipulating hand stimuli (Mouchetant-Rostaing, Giard, Bentin, et al., 2000). Whatever their exact nature, these 145-185 msec effects therefore suggest, first, that similar mechanisms could be engaged in age and gender perception, and second, that age and gender may be implicitly processed irrespective of their relevance to the task, through somewhat specialized mechanisms. Additional ERP effects were found at early latencies (45-90 msec) in all three discrimination conditions, and around 200-400 msec during explicit age and gender discrimination. These effects have been previously found in control conditions manipulating nonfacial stimuli and may therefore be related to more general categorization processes.

Cerebral mechanisms underlying orienting of attention towards auditory frequency changes.

Brain mechanisms underlying detection of auditory frequency changes were studied with event-related potentials (ERPs) in 14 human subjects discriminating visual stimuli. Scalp-current density mapping revealed bilateral components of mismatch negativity (MMN) in frontal and auditory cortices. Deviance-related activations in frontal and temporal cortex began to be significant at 94 ms and 154 ms in the right hemisphere, and at 128 ms and 132 ms in the left hemisphere. The magnitude of MMN-neuroelectric currents from the left temporal cortex correlated significantly (r = -0.56, p < 0.05) with distraction caused by MMN-eliciting deviant tones. These results suggest a complex cerebral circuitry involved in frequency change detection and strongly support the role of this circuitry in driving attention involuntarily towards potentially relevant frequency changes in the acoustic environment.

Maturation of frontal and temporal components of mismatch negativity (MMN) in children.

The mismatch negativity (MMN) response of auditory ERPs in adults appears to result from several overlapping components involving both frontal and temporal brain areas. Our aim was to test whether a similar configuration could be observed in children, and to examine the maturation rates of the different components. MMN (standard tones: 1000 Hz, deviants: 1100 Hz) was recorded from 28 scalp electrodes in 24 healthy children aged from 5 to 10 and in eight adults for comparison. Scalp current density analysis revealed both temporal and frontal components in children of all ages as well as in adults. Moreover the amplitudes of the temporal components were significantly greater in children than in adults, whereas the frontal components were similar at all ages. The results strongly suggest that MMN is mediated by at least two separate neural systems, and that the frontal system matures earlier than the sensory-specific system.

Early signs of visual categorization for biological and non-biological stimuli in humans.

In a previous experiment aimed at studying gender processing from faces, we had found unexpected early ERP differences (45-85 ms) in task-irrelevant stimuli between a condition in which the stimuli of each gender were delivered in separate runs, and a condition in which the stimuli of both genders were mixed. Similar effects were observed with hand stimuli. These early ERP differences were tentatively related to incidental categorization processes between male and female stimuli. The present study was designed to test the robustness of these early effects for faces, and to examine whether similar effects can also be generated between two classes of non-biological stimuli. We replicated the previous findings for faces, and found similar early differential effects (50-65 ms) for non-biological stimuli (grey and hatched geometrical shapes) only, however, when the two shape categories were separated by conspicuous visual characteristics. While these results can partly be explained by phenomena related to neuronal habituation in the visual cortex, they may also suggest the existence of coarse and automatic categorization processes for rapid distinction between two wide classes of stimuli with strong psychosocial significance for humans.

Auditory distraction: event-related potential and behavioral indices.

OBJECTIVE: The aim of this study was to illuminate behavioral and event-related potential (ERP) effects of attentional orienting and reorienting obtained in a newly developed auditory distraction paradigm, to provide more precise indicators about the neural generators of the ERP effects using scalp current density (SCD) analysis, and to evaluate the stability of the distraction effects. METHODS: In two sessions separated by 25 days, 10 subjects were presented with tones being of short (200 ms) and long (400 ms) duration equiprobably; tones were of high-probability standard or of low-probability deviant frequency. In Distraction condition, subjects had to behaviorally discriminate short from long tones. In Ignore condition, subjects were reading a book. Behavioral performance and multi-channel EEG were recorded. RESULTS: Task-irrelevant frequency deviations prolonged reaction times in the duration discrimination task by more than 35 ms and elicited the MMN and P3a components of the event-related potential. The P3a was followed by a negative deflection called RON (reorienting negativity). P3a and RON were absent in Ignore condition. All effects were found to be highly stable between sessions (product-moment correlations between 0.76 and 0.90). SCD analysis suggested frontal generators for P3a and for RON. CONCLUSIONS: It is demonstrated that small frequency deviations may yield distinct distraction effects in a tone duration discrimination task on a behavioral and on an electrophysiological level. Results support the hypothesis that frontal areas are involved in the exogenous orienting of attention (P3a) and in the reorienting of attention (RON). Due to the high stability of the deviance-related behavioral and ERP effects, this distraction paradigm may be utilized for clinical research.

Neurophysiological mechanisms of auditory selective attention in humans.

This chapter reviews the main data on the physiological substrates of auditory selective attention and their contribution to theoretical models of cognitive psychology. While event-related potentials, magnetoencephalography, and more recently neuroimaging techniques have provided fundamental information on the neural correlates of attention in the central cortical system, measurements of the frequency-following responses in the brainstem and evoked otoacoustic emissions at the cochlea strongly suggest attentional phenomena at the auditory periphery. We propose an adaptive filtering mechanism for selective auditory attention that can be flexibly and dynamically tuned depending on the attentional demand.

Neurophysiological correlates of face gender processing in humans.

Event-related potentials (ERPs) were recorded while subjects were involved in three gender-processing tasks based on human faces and on human hands. In one condition all stimuli were only of one gender, preventing any gender discrimination. In a second condition, faces (or hands) of men and women were intermixed but the gender was irrelevant for the subject's task; hence gender discrimination was assumed to be incidental. In the third condition, the task required explicit gender discrimination; gender processing was therefore assumed to be intentional. Gender processing had no effect on the occipito-temporal negative potential at approximately 170 ms after stimulation (N170 component of the ERP), suggesting that the neural mechanisms involved in the structural encoding of faces are different from those involved in the extraction of gender-related facial features. In contrast, incidental and intentional processing of face (but not hand) gender affected the ERPs between 145 and 185 ms from stimulus onset at more anterior scalp locations. This effect was interpreted as evidence for the direct visual processing of faces as described in Bruce and Young's model [Bruce, V. & Young, A. (1986) Br. J. Psychol., 77, 305-327]. Additional gender discrimination effects were observed for both faces and hands at mid-parietal sites around 45-85 ms latency, in the incidental task only. This difference was tentatively assumed to reflect an early mechanism of coarse visual categorization. Finally, intentional (but not incidental) gender processing affected the ERPs during a later epoch starting from approximately 200 ms and ending at approximately 250 ms for faces, and approximately 350 ms for hands. This later effect might be related to attention-based gender categorization or to a more general categorization activity.

Auditory-visual integration during multimodal object recognition in humans: a behavioral and electrophysiological study.

The aim of this study was (1) to provide behavioral evidence for multimodal feature integration in an object recognition task in humans and (2) to characterize the processing stages and the neural structures where multisensory interactions take place. Event-related potentials (ERPs) were recorded from 30 scalp electrodes while subjects performed a forced-choice reaction-time categorization task: At each trial, the subjects had to indicate which of two objects was presented by pressing one of two keys. The two objects were defined by auditory features alone, visual features alone, or the combination of auditory and visual features. Subjects were more accurate and rapid at identifying multimodal than unimodal objects. Spatiotemporal analysis of ERPs and scalp current densities revealed several auditory-visual interaction components temporally, spatially, and functionally distinct before 200 msec poststimulus. The effects observed were (1) in visual areas, new neural activities (as early as 40 msec poststimulus) and modulation (amplitude decrease) of the N185 wave to unimodal visual stimulus, (2) in the auditory cortex, modulation (amplitude increase) of subcomponents of the unimodal auditory N1 wave around 90 to 110 msec, and (3) new neural activity over the right fronto-temporal area (140 to 165 msec). Furthermore, when the subjects were separated into two groups according to their dominant modality to perform the task in unimodal conditions (shortest reaction time criteria), the integration effects were found to be similar for the two groups over the nonspecific fronto-temporal areas, but they clearly differed in the sensory-specific cortices, affecting predominantly the sensory areas of the nondominant modality. Taken together, the results indicate that multisensory integration is mediated by flexible, highly adaptive physiological processes that can take place very early in the sensory processing chain and operate in both sensory-specific and nonspecific cortical structures in different ways.

ERP manifestations of processing printed words at different psycholinguistic levels: time course and scalp distribution.

The aim of the present study was to examine the time course and scalp distribution of electrophysiological manifestations of the visual word recognition mechanism. Event-related potentials (ERPs) elicited by visually presented lists of words were recorded while subjects were involved in a series of oddball tasks. The distinction between the designated target and nontarget stimuli was manipulated to induce a different level of processing in each session (visual, phonological/phonetic, phonological/lexical, and semantic). The ERPs of main interest in this study were those elicited by nontarget stimuli. In the visual task the targets were twice as big as the nontargets. Words, pseudowords, strings of consonants, strings of alphanumeric symbols, and strings of forms elicited a sharp negative peak at 170 msec (N170); their distribution was limited to the occipito-temporal sites. For the left hemisphere electrode sites, the N170 was larger for orthographic than for nonorthographic stimuli and vice versa for the right hemisphere. The ERPs elicited by all orthographic stimuli formed a clearly distinct cluster that was different from the ERPs elicited by nonorthographic stimuli. In the phonological/phonetic decision task the targets were words and pseudowords rhyming with the French word vitrail, whereas the nontargets were words, pseudowords, and strings of consonants that did not rhyme with vitrail. The most conspicuous potential was a negative peak at 320 msec, which was similarly elicited by pronounceable stimuli but not by nonpronounceable stimuli. The N320 was bilaterally distributed over the middle temporal lobe and was significantly larger over the left than over the right hemisphere. In the phonological/lexical processing task we compared the ERPs elicited by strings of consonants (among which words were selected), pseudowords (among which words were selected), and by words (among which pseudowords were selected). The most conspicuous potential in these tasks was a negative potential peaking at 350 msec (N350) elicited by phonologically legal but not by phonologically illegal stimuli. The distribution of the N350 was similar to that of the N320, but it was broader and including temporo-parietal areas that were not activated in the "rhyme" task. Finally, in the semantic task the targets were abstract words, and the nontargets were concrete words, pseudowords, and strings of consonants. The negative potential in this task peaked at 450 msec. Unlike the lexical decision, the negative peak in this task significantly distinguished not only between phonologically legal and illegal words but also between meaningful (words) and meaningless (pseudowords) phonologically legal structures. The distribution of the N450 included the areas activated in the lexical decision task but also areas in the fronto-central regions. The present data corroborated the functional neuroanatomy of word recognition systems suggested by other neuroimaging methods and described their timecourse, supporting a cascade-type process that involves different but interconnected neural modules, each responsible for a different level of processing word-related information.

Mismatch negativity in dichotic listening: evidence for interhemispheric differences and multiple generators.

The characteristics of mismatch negativity (MMN) elicited by dichotic stimulation were examined using frequency-deviant stimuli presented to the right, to the left, or to both sides. The experiment was run twice, once using earphones and once using loudspeakers in free field. With both modes of stimulation, deviants presented in the left, right, or both ears, or tones that were switched between ears, elicited comparable MMNs, with a peak latency of about 180 ms. With earphones, the amplitude of the MMN was bigger at the frontal-lateral right hemisphere sites than at the homologous left-hemisphere sites for all deviance conditions. Scalp current density analysis revealed that deviance in the right side elicited bilaterally equivalent frontal current sinks and a trend towards stronger contralateral current sources at the mastoid sites. In contrast, left side deviance elicited frontal sinks and temporal current sources stronger over the right hemiscalp. These results are compatible with the multiple-generator model of MMN. The attention-related role of the MMN is discussed, suggesting comparable attention mechanisms for vision and audition.

Auditory cortex activation in deaf subjects during cochlear electrical stimulation. Evaluation by functional magnetic resonance imaging.

RATIONALE AND OBJECTIVES: The authors detect activation in the auditory cortex during cochlear electrical stimulation in deaf patients using functional magnetic resonance (MR) imaging. METHODS: Stimulating electrode was inserted gently under local anesthesia close to the round window membrane of the cochlea in seven cochlear implant candidates. These patients suffered from postlingual-acquired deafness. Four patients were stimulated above the electrical perception threshold and three below the electrical discomfort threshold. Functional scans (fast low-angle shot 91 mseconds/60 mseconds) were acquired in an oblique axial plane running parallel to the sylvian fissure. Four consecutive series of six images were obtained in 6 minutes. The acquisition time of each image was 15 seconds. RESULTS: During electrical cochlear stimulation below the discomfort threshold, the three patients described "auditory" sensations with activation of the superior temporal regions. In two patients with electrical stimulation of the left ear, the maximum signal intensity increased by 8.42% in the right auditory cortex and 5.69% in the left. In one patient with a right electrical stimulation only the left cortex was activated. Electrical cochlear stimulation above the perception threshold induces no significant activation in the auditory cortex. CONCLUSION: Functioning MR imaging can detect activation in the auditory cortex during cochlear electrical stimulation in deaf patients using a conventional 1.5-tesla system in a routine hospital environment. Further studies are needed to investigate its usefulness in clinical practice.

Separate representation of stimulus frequency, intensity, and duration in auditory sensory memory: an event-related potential and dipole-model analysis.

Abstract The present study analyzed the neural correlates of acoustic stimulus representation in echoic sensory memory. The neural traces of auditory sensory memory were indirectly studied by using the mismatch negativity (MMN), an event-related potential component elicited by a change in a repetitive sound. The MMN is assumed to reflect change detection in a comparison process between the sensory input from a deviant stimulus and the neural representation of repetitive stimuli in echoic memory. The scalp topographies of the MMNs elicited by pure tones deviating from standard tones by either frequency, intensity, or duration varied according to the type of stimulus deviance, indicating that the MMNs for different attributes originate, at least in part, from distinct neural populations in the auditory cortex. This result was supported by dipole-model analysis. If the MMN generator process occurs where the stimulus information is stored, these findings strongly suggest that the frequency, intensity, and duration of acoustic stimuli have a separate neural representation in sensory memory.

Two separate frontal components in the N1 wave of the human auditory evoked response.

Scalp current density analysis of the auditory evoked response to 1-kHz tone bursts delivered at various interstimulus intervals (ISIs) (from 1 s to 2 min in separate runs) shows that two different frontal components can be observed and functionally dissociated in the N1 time range: one is elicited for all ISIs, peaks at about 95 ms poststimulus, and has a full recovery time below 8 s; the second is elicited only by infrequent stimuli (ISIs > 4 s), peaks around 140 ms, and significantly increases in amplitude with increasing ISIs. The first component can be considered a new obligatory component in N1 elicited simultaneously with the responses in auditory cortex; the later component could correspond to the orienting Component III of Näätänen and Picton (1987).

Dissociation of temporal and frontal components in the human auditory N1 wave: a scalp current density and dipole model analysis.

This study reports a combined scalp current density (SCD) and dipole model analysis of the N1 wave of the auditory event-related potentials evoked by 1 kHz tone bursts delivered every second. The SCD distributions revealed: (i) a sink and a source of current reversing in polarity at the inferotemporal level of each hemiscalp, compatible with neural generators in and around the supratemporal plane of the auditory cortex, as previously reported; and (ii) bilateral current sinks over frontal areas. Consistently, dynamic dipole model analysis showed that generators in and outside the auditory cortex are necessary to account for the observed current fields between 65 and 140 msec post stimulus. The frontal currents could originate from the motor cortex, the supplementary motor area and/or the cingulate gyrus. The dissociation of an exogenous, obligatory frontal component from the sensory-specific response in the auditory N1 suggests that parallel processes served by distinct neural systems are activated during acoustic stimulation. Implications for recent models of auditory processing are discussed.

Auditory selective attention in the human cochlea.

According to current theories, auditory selective attention alters the sensory analysis of acoustic inputs only in the central auditory system. Despite numerous attempts, no evidence of attentional selection has been found in the auditory periphery. Measurements of evoked otoacoustic emissions (EOAEs) during a selective dichotic listening task showed that the EOAEs to tones in one ear had larger amplitude when attention was directed to this ear than when attention was directed to the opposite ear. The results indicate that genuine effects of auditory selective attention can be observed at the cochlear receptor.

Brain generators implicated in the processing of auditory stimulus deviance: a topographic event-related potential study.

The neurophysiological mechanisms underlying mismatch negativity (MMN) can be inferred from an examination of some of the brain generators involved in the process of this event-related potential (ERP) component. ERPs were recorded in two studies in which the subjects were involved in a selective dichotic listening task. Subjects were required to silently count rare stimuli deviating in pitch from a sequence of standard stimuli in one ear, while ignoring all the stimuli (standards and deviants) delivered randomly to the other ear. The results showed that, in all cases, the negative wave elicited by the deviant stimuli showed the highest amplitudes over the right hemiscalp irrespective of the ear of stimulation or the direction of attention. Scalp radial current density analysis showed that this asymmetric potential distribution could be attributed to the sum of activities of two sets of neural generators: one temporal, located in the vicinity of the primary auditory cortex, predominantly activated in the hemisphere contralateral to the ear of stimulation, and the other frontal, involving mainly the right hemisphere. The results are discussed in light of Näätänen's model: we suggest the dissociation of two functional processes on the basis of activity of distinct brain areas: a sensory memory mechanism related to the temporal generators, and an automatic attention-switching process related to the frontal generators.

Precautions in topographic mapping and in evoked potential map reading.

First, we consider the main points that must be addressed when constructing topographic maps: types of projection, methods of interpolation, number and locations of recording electrodes, and color scales. Data integrity and precautions in map interpretation are then examined for the case of evoked potential data.