Effect of temporal frequency on habituation in migraine.

Individuals with migraine tend to experience discomfort when viewing flickering stimuli. It has been suggested that one of the characteristics of migraine is a lack of habituation to repetitive visual stimuli, although findings can be mixed. Previous work has typically used similar visual stimuli (chequerboard) and only one temporal frequency. This study systematically varied the spatial and temporal characteristics of the visual stimulus, using steady-state visual evoked potentials to assess the differences in amplitude between migraine and control group over consecutive blocks of stimulation. Twenty individuals with migraine and 18 control observers were asked to rate their visual discomfort after viewing sequences of flickering Gabor patches with a frequency of either 3 or 9 Hz across three different spatial frequencies (low 0.5 cpd; mid-range 3 cpd; high 12 cpd). Compared to the control group, the migraine group showed a reduction in SSVEP responses with increased exposure, suggesting habituation processes are intact at 3-Hz stimulation. However, at 9-Hz stimulation, there was evidence of increased responses with increasing exposure in the migraine group in particular, which might suggest a build-up of the response over repetitive presentations. Visual discomfort varied with spatial frequency, for both 3- and 9-Hz stimuli, the highest spatial frequencies were the least uncomfortable compared to the low- and mid-range spatial frequencies in both groups. This difference in SSVEP response behaviour, dependent on temporal frequency, is important to consider when researching the effects of repetitive visual stimulation in migraine and could give some indication of build-up of effects leading to aversion to visual stimuli.

Neural correlates of visual spatial selective attention are altered at early and late processing stages in human amblyopia.

Amblyopia is a neurodevelopmental visual disorder which results in reduced visual acuity in one eye and impaired binocular interactions. Previous studies suggest attentional deficits in amblyopic individuals. However, spatial cues which orient attention to a visual field improved performance. Here, we investigate the neural correlates of auditory-visual spatial selective attention in amblyopia during EEG recording. An auditory cue, that was followed by the presentation of two Gabor patches presented in the lower left and right visual fields, indicated the most likely location of an upcoming target Gabor. The target Gabor differed in orientation from the more frequently presented non-target Gabor patches. Adults with amblyopia and neurotypical observers were asked to detect the target Gabor monocularly at the cued location, while withholding their response to targets presented at the uncued location and to all non-target Gabor patches. Higher response rates were observed for cued compared to uncued targets in both groups. However, amblyopic individuals detected targets less efficiently with their amblyopic eye as compared to their fellow eye. Correspondingly, event-related potentials (ERPs) recorded to the onset of the non-target Gabor patches were delayed at early processing stages (150-300 ms: posterior N100) and reduced in amplitude at later time windows (150-350 ms: P200, 300-500 ms: sustained activity) in the amblyopic eye compared to the fellow eye. Such interocular differences were not observed in neurotypical observers. These findings suggest that neural resources allocated to the early formation of visual discrimination as well as later stimulus recognition processes are altered in the amblyopic eye.

Children can optimally integrate multisensory information after a short action-like mini game training.

Combining information across different sensory modalities is of critical importance for the animal's survival and a core feature of human's everyday life. In adulthood, sensory information is often integrated in a statistically optimal fashion, so that the combined estimates of two or more senses are more reliable than the best single one. Several studies have shown that young children use one sense to calibrate the others, which results in unisensory dominance and undermines their optimal multisensory integration abilities. In this study we trained children aged 4-5 years with action-like mini games, to determine whether it could improve their multisensory as well as their visuo-spatial skills. Multisensory integration abilities were assessed using a visuo-haptic size discrimination task, while visuo-spatial attention skills were investigated using a multiple object tracking task (MOT). We found that 2-weeks training were sufficient to observe both optimal multisensory integration and visuo-spatial enhancements selectively in the group trained with action-like mini games. This plastic change persisted up to 3 months, as assessed in a follow-up. Our novel findings reveal that abilities that are commonly known to emerge in late childhood can be promoted in younger children through action-like mini games and have long-lasting effects. Our data have clinical implications, in that they suggest that specific trainings could potentially help children with multisensory integration deficits.

Neural Correlates of Enhanced Visual Attentional Control in Action Video Game Players: An Event-Related Potential Study.

Action video game players (AVGPs) outperform non-action video game players (NAVGPs) on a range of perceptual and attentional tasks. Although several studies have reported neuroplastic changes within the frontoparietal networks of attention in AVGPs, little is known about possible changes in attentional modulation in low-level visual areas. To assess the contribution of these different levels of neural processing to the perceptual and attentional enhancements noted in AVGPs, visual event-related potentials (ERPs) were recorded from 14 AVGPs and 14 NAVGPs during a target discrimination task that required participants to attend to rapid sequences of Gabor patches under either focused or divided attention conditions. AVGPs responded faster to target Gabors in the focused attention condition compared with the NAVGPs. Correspondingly, ERPs to standard Gabors revealed a more pronounced negativity in the time range of the parietally generated anterior N1 component in AVGPs compared with NAVGPs during focused attention. In addition, the P2 component of the visual ERP was more pronounced in AVGPs than in NAVGPs over the hemisphere contralateral to the stimulus position in response to standard Gabors. Contrary to predictions, however, attention-modulated occipital components generated in the low-level extrastriate visual pathways, including the P1 and posterior N1, showed no significant group differences. Thus, the main neural signature of enhanced perceptual and attentional control functions in AVGPs appears linked to an attention-dependent parietal process, indexed by the anterior N1 component, and possibly to more efficient higher-order perceptual processing, indexed by the P2 component.

Activation in the angular gyrus and in the pSTS is modulated by face primes during voice recognition.

The aim of the present study was to better understand the interaction of face and voice processing when identifying people. In a S1-S2 crossmodal priming fMRI experiment, the target (S2) was a disyllabic voice stimulus, whereas the modality of the prime (S1) was manipulated blockwise and consisted of the silent video of a speaking face in the crossmodal condition or of a voice stimulus in the unimodal condition. Primes and targets were from the same speaker (person-congruent) or from two different speakers (person-incongruent). Participants had to classify the S2 as either an old or a young person. Response times were shorter after a congruent than after an incongruent face prime. The right posterior superior temporal sulcus (pSTS) and the right angular gyrus showed a significant person identity effect (person-incongruent > person-congruent) in the crossmodal condition but not in the unimodal condition. In the unimodal condition, a person identity effect was observed in the bilateral inferior frontal gyrus. Our data suggest that both the priming with a voice and with a face result in a preactivated voice representation of the respective person, which eventually facilitates (person-congruent trials) or hampers (person-incongruent trials) the processing of the identity of a subsequent voice. This process involves activation in the right pSTS and in the right angular gyrus for voices primed by faces, but not for voices primed by voices. Hum Brain Mapp 38:2553-2565, 2017. © 2017 Wiley Periodicals, Inc.

Oscillatory activity reflects differential use of spatial reference frames by sighted and blind individuals in tactile attention.

Touch can be localized either on the skin in anatomical coordinates, or, after integration with posture, in external space. Sighted individuals are thought to encode touch in both coordinate systems concurrently, whereas congenitally blind individuals exhibit a strong bias for using anatomical coordinates. We investigated the neural correlates of this differential dominance in the use of anatomical and external reference frames by assessing oscillatory brain activity during a tactile spatial attention task. The EEG was recorded while sighted and congenitally blind adults received tactile stimulation to uncrossed and crossed hands while detecting rare tactile targets at one cued hand only. In the sighted group, oscillatory alpha-band activity (8-12Hz) in the cue-target interval was reduced contralaterally and enhanced ipsilaterally with uncrossed hands. Hand crossing attenuated the degree of posterior parietal alpha-band lateralization, indicating that attention deployment was affected by external spatial coordinates. Beamforming suggested that this posture effect originated in the posterior parietal cortex. In contrast, cue-related lateralization of central alpha-band as well as of beta-band activity (16-24Hz) were unaffected by hand crossing, suggesting that these oscillations exclusively encode anatomical coordinates. In the blind group, central alpha-band activity was lateralized, but did not change across postures. The pattern of beta-band activity was indistinguishable between groups. Because the neural mechanisms for posterior alpha-band generation seem to be linked to developmental vision, we speculate that the lack of this neural mechanism in blind individuals is related to their preferred use of anatomical over external spatial codes in sensory processing.

Crossmodal plasticity in the fusiform gyrus of late blind individuals during voice recognition.

Blind individuals are trained in identifying other people through voices. In congenitally blind adults the anterior fusiform gyrus has been shown to be active during voice recognition. Such crossmodal changes have been associated with a superiority of blind adults in voice perception. The key question of the present functional magnetic resonance imaging (fMRI) study was whether visual deprivation that occurs in adulthood is followed by similar adaptive changes of the voice identification system. Late blind individuals and matched sighted participants were tested in a priming paradigm, in which two voice stimuli were subsequently presented. The prime (S1) and the target (S2) were either from the same speaker (person-congruent voices) or from two different speakers (person-incongruent voices). Participants had to classify the S2 as either coming from an old or a young person. Only in late blind but not in matched sighted controls, the activation in the anterior fusiform gyrus was modulated by voice identity: late blind volunteers showed an increase of the BOLD signal in response to person-incongruent compared with person-congruent trials. These results suggest that the fusiform gyrus adapts to input of a new modality even in the mature brain and thus demonstrate an adult type of crossmodal plasticity.

Brain systems mediating voice identity processing in blind humans.

Blind people rely more on vocal cues when they recognize a person's identity than sighted people. Indeed, a number of studies have reported better voice recognition skills in blind than in sighted adults. The present functional magnetic resonance imaging study investigated changes in the functional organization of neural systems involved in voice identity processing following congenital blindness. A group of congenitally blind individuals and matched sighted control participants were tested in a priming paradigm, in which two voice stimuli (S1, S2) were subsequently presented. The prime (S1) and the target (S2) were either from the same speaker (person-congruent voices) or from two different speakers (person-incongruent voices). Participants had to classify the S2 as either a old or a young person. Person-incongruent voices (S2) compared with person-congruent voices elicited an increased activation in the right anterior fusiform gyrus in congenitally blind individuals but not in matched sighted control participants. In contrast, only matched sighted controls showed a higher activation in response to person-incongruent compared with person-congruent voices (S2) in the right posterior superior temporal sulcus. These results provide evidence for crossmodal plastic changes of the person identification system in the brain after visual deprivation.

Is the processing of affective prosody influenced by spatial attention? An ERP study.

BACKGROUND: The present study asked whether the processing of affective prosody is modulated by spatial attention. Pseudo-words with a neutral, happy, threatening, and fearful prosody were presented at two spatial positions. Participants attended to one position in order to detect infrequent targets. Emotional prosody was task irrelevant. The electro-encephalogram (EEG) was recorded to assess processing differences as a function of spatial attention and emotional valence. RESULTS: Event-related potentials (ERPs) differed as a function of emotional prosody both when attended and when unattended. While emotional prosody effects interacted with effects of spatial attention at early processing levels (< 200 ms), these effects were additive at later processing stages (> 200 ms). CONCLUSIONS: Emotional prosody, therefore, seems to be partially processed outside the focus of spatial attention. Whereas at early sensory processing stages spatial attention modulates the degree of emotional voice processing as a function of emotional valence, emotional prosody is processed outside of the focus of spatial attention at later processing stages.

Evidence of target enhancement and distractor suppression in early visual areas.

Although the mechanisms of target enhancement and distractor suppression have been investigated along the visual processing hierarchy, there remains some unknown as to the role of perceptual load on the competition between different task-related information as attention deployment is manipulated. We present an fMRI spatial cueing paradigm, in which 32 participants had to attend to either a left or a right hemifield location and to indicate the orientation of the target Gabor that was presented simultaneously to a noise patch distractor. Critically, the target could appear at either the cued, valid location or at the uncued, invalid location; in the latter, the noise patch distractor appeared at the cued location. Perceptual load was manipulated by the presence or absence of high-contrast Gabor patches close to the fixation cross, which acted as lateral masks. Behavioural results indicated that participants performed more efficiently in validly cued trials compared to invalidly cued trials and under low compared to high load. Enhancement effects for targets and suppression effects for noise patches were greater in early visual areas at high load, that is in the presence of lateral masks. These results are in line with the hypothesis that attention results in both target enhancement and distractor suppression, and that these effects are most marked under high perceptual load. Theoretical implications of these results for different models of attention are discussed.

The Impact of Audio-Visual, Visual and Auditory Cues on Multiple Object Tracking Performance in Children with Autism.

Previous studies have documented differences in processing multisensory information by children with autism compared to typically developing children. Furthermore, children with autism have been found to track fewer multiple objects on a screen than those without autism, suggesting reduced attentional control. In the present study, we investigated whether children with autism (n = 33) and children without autism (n = 33) were able to track four target objects moving amongst four indistinguishable distractor objects while sensory cues were presented. During tracking, we presented various types of cues - auditory, visual, or audio-visual or no cues while target objects bounced off the inner boundary of a centralized circle. We found that children with autism tracked fewer targets than children without autism. Furthermore, children without autism showed improved tracking performance in the presence of visual cues, whereas children with autism did not benefit from sensory cues. Whereas multiple object tracking performance improved with increasing age in children without autism, especially when using audio-visual cues, children with autism did not show age-related improvement in tracking. These results are in line with the hypothesis that attention and the ability to integrate sensory cues during tracking are reduced in children with autism. Our findings could contribute valuable insights for designing interventions that incorporate multisensory information.

Exploring the effectiveness of auditory, visual, and audio-visual sensory cues in a multiple object tracking environment.

Maintaining object correspondence among multiple moving objects is an essential task of the perceptual system in many everyday life activities. A substantial body of research has confirmed that observers are able to track multiple target objects amongst identical distractors based only on their spatiotemporal information. However, naturalistic tasks typically involve the integration of information from more than one modality, and there is limited research investigating whether auditory and audio-visual cues improve tracking. In two experiments, we asked participants to track either five target objects or three versus five target objects amongst similarly indistinguishable distractor objects for 14 s. During the tracking interval, the target objects bounced occasionally against the boundary of a centralised orange circle. A visual cue, an auditory cue, neither or both coincided with these collisions. Following the motion interval, the participants were asked to indicate all target objects. Across both experiments and both set sizes, our results indicated that visual and auditory cues increased tracking accuracy although visual cues were more effective than auditory cues. Audio-visual cues, however, did not increase tracking performance beyond the level of purely visual cues for both high and low load conditions. We discuss the theoretical implications of our findings for multiple object tracking as well as for the principles of multisensory integration.

Intramodal cortical plastic changes after moderate visual impairment in human amblyopia.

Early blindness results in alterations in the neural responses to auditory stimuli. Here we show that even moderately reduced vision in one eye early in life is sufficient to induce neural plastic changes in voice processing. We asked individuals with reduced visual acuity in one eye due to amblyopia to attend to vocal cues during electroencephalogram recording. We found enhanced frontal auditory responses at 125 ms-225 ms, which were correlated with reduced vision in the amblyopic eye, but not the fellow eye. Our results indicate intramodal reorganization, typically observed after congenital profound visual deprivation.

The Effect of Motion Direction and Eccentricity on Vection, VR Sickness and Head Movements in Virtual Reality.

Virtual Reality (VR) experienced through head-mounted displays often leads to vection, discomfort and sway in the user. This study investigated the effect of motion direction and eccentricity on these three phenomena using optic flow patterns displayed using the Valve Index. Visual motion stimuli were presented in the centre, periphery or far periphery and moved either in depth (back and forth) or laterally (left and right). Overall vection was stronger for motion in depth compared to lateral motion. Additionally, eccentricity primarily affected stimuli moving in depth with stronger vection for more peripherally presented motion patterns compared to more central ones. Motion direction affected the various aspects of VR sickness differently and modulated the effect of eccentricity on VR sickness. For stimuli moving in depth far peripheral presentation caused more discomfort, whereas for lateral motion the central stimuli caused more discomfort. Stimuli moving in depth led to more head movements in the anterior-posterior direction when the entire visual field was stimulated. Observers demonstrated more head movements in the anterior-posterior direction compared to the medio-lateral direction throughout the entire experiment independent of motion direction or eccentricity of the presented moving stimulus. Head movements were elicited on the same plane as the moving stimulus only for stimuli moving in depth covering the entire visual field. A correlation showed a positive relationship between dizziness and vection duration and between general discomfort and sway. Identifying where in the visual field motion presented to an individual causes the least amount of VR sickness without losing vection and presence can guide development for Virtual Reality games, training and treatment programmes.

Unimodal and crossmodal gradients of spatial attention: Evidence from event-related potentials.

Behavioral and event-related potential (ERP) studies have shown that spatial attention is gradually distributed around the center of the attentional focus. The present study compared uni- and crossmodal gradients of spatial attention to investigate whether the orienting of auditory and visual spatial attention is based on modality specific or supramodal representations of space. Auditory and visual stimuli were presented from five speaker locations positioned in the right hemifield. Participants had to attend to the innermost or outmost right position in order to detect either visual or auditory deviant stimuli. Detection rates and event-related potentials (ERPs) indicated that spatial attention is distributed as a gradient. Unimodal spatial ERP gradients correlated with the spatial resolution of the modality. Crossmodal spatial gradients were always broader than the corresponding unimodal spatial gradients. These results suggest that both modality specific and supramodal spatial representations are activated during orienting attention in space.

The role of spatial selective attention in the processing of affective prosodies in congenitally blind adults: An ERP study.

The question whether spatial selective attention is necessary in order to process vocal affective prosody has been controversially discussed in sighted individuals: whereas some studies argue that attention is required in order to process emotions, other studies conclude that vocal prosody can be processed even outside the focus of spatial selective attention. Here, we asked whether spatial selective attention is necessary for the processing of affective prosodies after visual deprivation from birth. For this purpose, pseudowords spoken in happy, neutral, fearful or threatening prosodies were presented at the left or right loudspeaker. Congenitally blind individuals (N = 8) and sighted controls (N = 13) had to attend to one of the loudspeakers and detect rare pseudowords presented at the attended loudspeaker during EEG recording. Emotional prosody of the syllables was task-irrelevant. Blind individuals outperformed sighted controls by being more efficient in detecting deviant syllables at the attended loudspeaker. A higher auditory N1 amplitude was observed in blind individuals compared to sighted controls. Additionally, sighted controls showed enhanced attention-related ERP amplitudes in response to fearful and threatening voices during the time range of the N1. By contrast, blind individuals revealed enhanced ERP amplitudes in attended relative to unattended locations irrespective of the affective valence in all time windows (110-350 ms). These effects were mainly observed at posterior electrodes. The results provide evidence for "emotion-general" auditory spatial selective attention effects in congenitally blind individuals and suggest a potential reorganization of the voice processing brain system following visual deprivation from birth.

Faster Visual Information Processing in Video Gamers Is Associated With EEG Alpha Amplitude Modulation.

Video gaming, specifically action video gaming, seems to improve a range of cognitive functions. The basis for these improvements may be attentional control in conjunction with reward-related learning to amplify the execution of goal-relevant actions while suppressing goal-irrelevant actions. Given that EEG alpha power reflects inhibitory processing, a core component of attentional control, it might represent the electrophysiological substrate of cognitive improvement in video gaming. The aim of this study was to test whether non-video gamers (NVGs), non-action video gamers (NAVGs) and action video gamers (AVGs) exhibit differences in EEG alpha power, and whether this might account for differences in visual information processing as operationalized by the theory of visual attention (TVA). Forty male volunteers performed a visual short-term memory paradigm where they memorized shape stimuli depicted on circular stimulus displays at six different exposure durations while their EEGs were recorded. Accuracy data was analyzed using TVA-algorithms. There was a positive correlation between the extent of post-stimulus EEG alpha power attenuation (10-12 Hz) and speed of information processing across all participants. Moreover, both EEG alpha power attenuation and speed of information processing were modulated by an interaction between group affiliation and time on task, indicating that video gamers showed larger EEG alpha power attenuations and faster information processing over time than NVGs - with AVGs displaying the largest increase. An additional regression analysis affirmed this observation. From this we concluded that EEG alpha power might be a promising neural substrate for explaining cognitive improvement in video gaming.

Event-Related Potentials Reveal Evidence for Late Integration of Emotional Prosody and Facial Expression in Dynamic Stimuli: An ERP Study.

The aim of the present study was to test whether multisensory interactions of emotional signals are modulated by intermodal attention and emotional valence. Faces, voices and bimodal emotionally congruent or incongruent face-voice pairs were randomly presented. The EEG was recorded while participants were instructed to detect sad emotional expressions in either faces or voices while ignoring all stimuli with another emotional expression and sad stimuli of the task irrelevant modality. Participants processed congruent sad face-voice pairs more efficiently than sad stimuli paired with an incongruent emotion and performance was higher in congruent bimodal compared to unimodal trials, irrespective of which modality was task-relevant. Event-related potentials (ERPs) to congruent emotional face-voice pairs started to differ from ERPs to incongruent emotional face-voice pairs at 180 ms after stimulus onset: Irrespectively of which modality was task-relevant, ERPs revealed a more pronounced positivity (180 ms post-stimulus) to emotionally congruent trials compared to emotionally incongruent trials if the angry emotion was presented in the attended modality. A larger negativity to incongruent compared to congruent trials was observed in the time range of 400-550 ms (N400) for all emotions (happy, neutral, angry), irrespectively of whether faces or voices were task relevant. These results suggest an automatic interaction of emotion related information.

Alpha-band oscillations reflect external spatial coding for tactile stimuli in sighted, but not in congenitally blind humans.

We investigated the function of oscillatory alpha-band activity in the neural coding of spatial information during tactile processing. Sighted humans concurrently encode tactile location in skin-based and, after integration with posture, external spatial reference frames, whereas congenitally blind humans preferably use skin-based coding. Accordingly, lateralization of alpha-band activity in parietal regions during attentional orienting in expectance of tactile stimulation reflected external spatial coding in sighted, but skin-based coding in blind humans. Here, we asked whether alpha-band activity plays a similar role in spatial coding for tactile processing, that is, after the stimulus has been received. Sighted and congenitally blind participants were cued to attend to one hand in order to detect rare tactile deviant stimuli at this hand while ignoring tactile deviants at the other hand and tactile standard stimuli at both hands. The reference frames encoded by oscillatory activity during tactile processing were probed by adopting either an uncrossed or crossed hand posture. In sighted participants, attended relative to unattended standard stimuli suppressed the power in the alpha-band over ipsilateral centro-parietal and occipital cortex. Hand crossing attenuated this attentional modulation predominantly over ipsilateral posterior-parietal cortex. In contrast, although contralateral alpha-activity was enhanced for attended versus unattended stimuli in blind participants, no crossing effects were evident in the oscillatory activity of this group. These findings suggest that oscillatory alpha-band activity plays a pivotal role in the neural coding of external spatial information for touch.

Spatial coordinate systems for tactile spatial attention depend on developmental vision: evidence from event-related potentials in sighted and congenitally blind adult humans.

Changes in limb posture (such as crossing the hands) can impair people's performance in tasks such as those involving temporal order judgements, when one tactile stimulus is presented to either hand. This crossed hands deficit has been attributed to a conflict between externally and anatomically anchored reference systems when people localize tactile stimuli. Interestingly, however, the performance of congenitally blind adults does not seem to be affected by crossing the hands, suggesting a default use of an anatomically rather than an externally anchored reference system for tactile localization. In the present study, 12 congenitally blind and 12 sighted adults were instructed to attend to either the left or the right hand on a trial-by-trial basis in order to detect rare deviants (consisting of a double touch) at that hand, while ignoring both deviants at the other hand and frequent standard stimuli (consisting of a single touch) presented to either hand. Only the sighted participants performed less accurately when they crossed their hands. Concurrent electroencephalogram recordings revealed an early contralateral attention positivity, followed by an attention negativity in the sighted group when they adopted the uncrossed hands posture. For the crossed hand posture, only the attention negativity was observed with reduced amplitude in the sighted group. By contrast, the congenitally blind group displayed an event-related potential attention negativity that did not vary when the posture of their hands was changed. These results demonstrate that the default use of an external frame of reference for tactile localization seems to depend on developmental vision.