Attention to central and peripheral visual space in a movement detection task: an event-related potential and behavioral study. II. Congenitally deaf adults

We compared the effects of focussed attention upon event-related brain potentials (ERPs) to peripherally and centrally located visual stimuli in congenitally deaf subjects (Ss). The results were compared with those obtained from a group of normal hearing Ss in the same paradigm. ERPs from deaf and hearing Ss displayed similar attention-related changes with attention to the centrally located stimuli. These included enhanced amplitudes of the N1 component (157 ms) over the occipital regions of both hemispheres. By contrast, with attention to peripheral visual stimuli, ERPs from deaf Ss displayed attention-related increases that were several times larger than those from hearing Ss and different in scalp distribution. Whereas for hearing Ss the principal effects of attention to peripheral stimuli occurred over the contralateral parietal region, in deaf Ss the effects were also observed over the occipital regions of both hemispheres. In addition, lateral asymmetries in behavior and the ERPs indicated a greater role for the right hemisphere in this task in hearing Ss, but predominance of the left hemisphere in deaf Ss. These results suggest that auditory deprivation since birth has major effects on the development of the peripheral visual system. The specific pattern of group differences is discussed in relation to other studies of the effects of unimodal deprivation on the development of remaining modalities.     

Attention to central and peripheral visual space in a movement detection task: an event-related potential and behavioral study. I. Normal hearing adults

The effects of focussed attention to peripherally and centrally located visual stimuli were compared via an analysis of event-related brain potentials (ERPs) while subjects detected the direction of motion of a white square in a specified location. While attention to both peripheral and foveal stimuli produced enhancements of the early ERP components, the distribution over the scalp of the attention-related changes varied according to stimulus location. The attention-related increase in the amplitude of the N1 wave (157 ms) to the peripheral stimuli was greater over the parietal region of the hemisphere contralateral to the attended visual field. By contrast, the largest effects of foveally directed attention occurred over the occipital regions where the increase was bilaterally symmetrical. Additionally, the effects of attention on the ERPs were significantly larger for moving than for stationary stimuli, and this effect was greater for peripheral than for central attention. A long-latency positive displacement component (300-600 ms) was larger over the right than the left hemisphere during attention to the lateral visual fields, but was symmetrical in amplitude when central stimuli were attended. These results suggest that different pathways are modulated when attention is deployed to different regions of the visual fields. Further, they suggest that the special role of the right hemisphere in spatial attention may be limited to analysis of information in the visual periphery.     

Evidence of minimal cerebral asymmetries for the processing of English words and American sign language in the congenitally deaf

Previous investigators reported that auditory association cortex is usually larger within the left than within the right hemisphere, even in the newborn. They suggested that this asymmetry could play a causal role in the predominant tendency for language processes to lateralize in the left rather than in the right hemisphere. For congenitally deaf persons, however, the role of auditory association area asymmetry presumably would be nullified. This question was addressed by comparing the performances of congenitally deaf college students and hearing persons on lateralized tachistoscopic tasks thought to reflect hemispheric language and spatial processing efficiencies. The results indicated that indicated that congenitally deaf Ss showed minimal half-field asymmetries for both English word stimuli and for American Sign Language stimuli (drawings of hand configurations representing letters and words for the deaf Ss). Results were interpreted as supporting auditory experience as a major determinant of cerebral functional asymmetries and as contradictory to clinical reports that had suggested that the cerebral organization of “communicative” functions were entirely comparable in deaf and hearing persons.     

Visual contrast sensitivity in deaf versus hearing populations: exploring the perceptual consequences of auditory deprivation and experience with a visual language

Early deafness in humans provides a unique opportunity to examine the perceptual consequences of altered sensory experience. In particular, visual perception in the deaf may be altered as a result of their auditory deprivation and/or because the deaf rely heavily upon a visual language (American Sign Language, or ASL, in the US). Recently, we found that deaf, but not hearing, subjects exhibit a right visual field/left hemisphere advantage on a low-level direction of motion task, a finding that has been attributed to the deaf's experience with ASL [Psychol. Sci. 10 (1999) 256; Brain Res. 405 (1987) 268]. In order to determine whether this visual field asymmetry generalizes to other low-level visual functions, in this study we measured contrast sensitivity in deaf and hearing subjects to moving stimuli over a range of speeds (0.125-64 degrees /s). We hypothesized that if ASL use drives differences between hearing and deaf subjects, such differences may occur over a restricted range of speeds most commonly found in ASL. In addition, we tested a third group, hearing native signers who learned ASL early from their deaf parents, to further assess whether potential differences between groups results from ASL use. These experiments reveal no overall differences in contrast sensitivity, nor differences in visual field asymmetries, across subject groups at any speed tested. Thus, differences previously observed between deaf and hearing subjects for discriminating the direction of moving stimuli do not generalize to contrast sensitivity for moving stimuli, a result that has implications for the neural level at which plastic changes occur in the visual system of deaf subjects.     

Comparing the effects of auditory deprivation and sign language within the auditory and visual cortex

To investigate neural plasticity resulting from early auditory deprivation and use of American Sign Language, we measured responses to visual stimuli in deaf signers, hearing signers, and hearing nonsigners using functional magnetic resonance imaging. We examined "compensatory hypertrophy" (changes in the responsivity/size of visual cortical areas) and "cross-modal plasticity" (changes in auditory cortex responses to visual stimuli). We measured the volume of early visual areas (V1, V2, V3, V4, and MT+). We also measured the amplitude of responses within these areas, and within the auditory cortex, to a peripheral visual motion stimulus that was attended or ignored. We found no major differences between deaf and hearing subjects in the size or responsivity of early visual areas. In contrast, within the auditory cortex, motion stimuli evoked significant responses in deaf subjects, but not in hearing subjects, in a region of the right auditory cortex corresponding to Brodmann's areas 41, 42, and 22. This hemispheric selectivity may be due to a predisposition for the right auditory cortex to process motion; earlier studies report a right hemisphere bias for auditory motion in hearing subjects. Visual responses within the auditory cortex of deaf subjects were stronger for attended than ignored stimuli, suggesting top-down processes. Hearing signers did not show visual responses in the auditory cortex, indicating that cross-modal plasticity can be attributed to auditory deprivation rather than sign language experience. The largest effects of auditory deprivation occurred within the auditory cortex rather than the visual cortex, suggesting that the absence of normal input is necessary for large-scale cortical reorganization to occur.     

Dyslexic adolescents: evidence of impaired visual and auditory language processing associated with normal lateralization and visual responsivity.

The questions of whether chronically dyslexic adolescent suffer any deficits of simple language stimulus processing or are less left hemisphere dominant than normal reading controls were addressed. The dyslexics were chosen for clarity of their specific reading problem and were older than dyslexics previously studied with lateralizing tests. Tasks administered in Experiment I were unilateral and bilateral tachistoscopic work recognitions and a tachistoscopic recognition report-time task for single lateralized letter stimuli. Experiment II, conducted a year later, readministered these tasks with modifications, and added dichotic digits and motor reaction time-stimulus detection tasks. It was concluded that right handed, chronic dyslexics: (1) possess left hemisphere language specialization; (2) show normal interhemispheric processing delays for single letter stimuli; (3) are, unlike nondyslexis but equally poor-reading Ss, clearly impaired in their efficiency of visual and auditory processing of simple language stimuli; (4) possess clear auditory memory deficits for verbal material; and (5) may possess an additional deficit of left hemisphere visual association area function.     

Altered spatial distribution of visual attention in near and far space after early deafness

Early deafness results in a redistribution of more attentional resources to the visual periphery in near space, specifically under conditions of selective attention, probably to compensate for the loss of auditory alertness to potentially dangerous stimuli from outside the current attentional focus. It remains poorly understood, however, whether spatial distribution of attention in far space is altered by early deafness as well. In the present study, we investigated whether and how early deafness alters the distribution of visuospatial attention in far space, compared to hearing controls. We asked deaf individuals and hearing controls to perform a flanker task with either peripheral or central distractors, either in near or far space. Sizes of compatibility effect were used to assess the amount of attentional resources received by the peripheral and central distractors. In near space, peripheral distractors induced significantly larger compatibility effect in deaf individuals than in hearing controls while central distractors induced significantly larger compatibility effect in hearing controls than in deaf individuals. On the other hand in far space, although peripheral distractors induced equivalent sizes of compatibility effect in the deaf and hearing groups, central distractors caused significant compatibility effect only in deaf individuals, but not in hearing controls. Our results suggest that early deafness results in a redistribution of visuospatial attention not only in near space but also in far space, with enhanced peripheral attention in near space and enhanced central attention in far space.     

Perception of static eye gaze direction facilitates subsequent early visual processing.

OBJECTIVE: Using event-related potentials (ERPs), it has been recently shown that a reflexive shift of attention following the observation of a dynamic eye gaze cue enhances and speeds up early visual processing of a target presented at the gazed-at location. Here we investigate whether similar early sensory modulations are also elicited by static gaze cues, or if previously described attentional effects were caused mainly by visual motion cues and not by eye gaze direction per se. Furthermore, we explore if these possible attentional orienting effects reflect facilitation of the processing of cued stimuli, inhibition of the unattended stimuli, or both. METHODS: Subjects were presented with a face looking to the right or left visual field (VF), or straight away, before the occurrence of a lateralized target to detect. There were 3 conditions in this nonpredictive cueing task: (1) target presented in the VF indicated by the eye gaze direction (congruent); (2) opposite to the eye gaze direction (incongruent); or (3) preceded by a straight gazing face (neutral). RESULTS: Subjects were faster at detecting congruently than incongruently and neutrally cued targets. Facilitation effects were observed on early ERP components: the occipital P1 and occipito-temporal N1 components were speeded up as early as approximately 100 ms following stimulus onset (P1), and enhanced (P1 and N1) in response to congruent trials, particularly in the right hemisphere. CONCLUSIONS: Spatial attention triggered by static eye gaze direction produces response facilitations - predominantly lateralized to the right hemisphere - from the early sensory stages of visual processing. Significance: This study provides the first evidence of a speeding up and amplification of early visual processing following attention triggered by static eye gaze perception.     

Hemispheric asymmetry in processing visual half-field pattern-reversal stimuli assessed by reaction time and evoked potentials

Reversing checkerboard patterns, like those used to elicit clinical visual evoked potentials (VEPs), were adapted to the study of cerebral lateralization of visual processing in a divided visual field experiment. Seventeen right-handed subjects (9 male, 8 female) provided left- and right-handed responses to pattern-reversal stimuli presented in left, right and both visual half-fields in pseudorandom order. Simple unwarned RTs were shorter for left- compared with right-field stimuli under left-hand response conditions. Parietal VEPs obtained from 9 of the subjects showed larger amplitudes for left compared with right field stimuli at the right hemisphere. The results were consistent with an efficiency model of cerebral dominance incorporating both interhemispheric transfer time and a reduced efficiency for left hemisphere processing of our patterned visual stimuli.     

Signed words in the congenitally deaf evoke typical late lexicosemantic responses with no early visual responses in left superior temporal cortex

Congenitally deaf individuals receive little or no auditory input, and when raised by deaf parents, they acquire sign as their native and primary language. We asked two questions regarding how the deaf brain in humans adapts to sensory deprivation: (1) is meaning extracted and integrated from signs using the same classical left hemisphere frontotemporal network used for speech in hearing individuals, and (2) in deafness, is superior temporal cortex encompassing primary and secondary auditory regions reorganized to receive and process visual sensory information at short latencies? Using MEG constrained by individual cortical anatomy obtained with MRI, we examined an early time window associated with sensory processing and a late time window associated with lexicosemantic integration. We found that sign in deaf individuals and speech in hearing individuals activate a highly similar left frontotemporal network (including superior temporal regions surrounding auditory cortex) during lexicosemantic processing, but only speech in hearing individuals activates auditory regions during sensory processing. Thus, neural systems dedicated to processing high-level linguistic information are used for processing language regardless of modality or hearing status, and we do not find evidence for rewiring of afferent connections from visual systems to auditory cortex.     

Altered visual-evoked potentials in congenitally deaf adults

Visual-evoked potentials recorded from the scalp of congenitally deaf adults were significantly larger over both auditory and visual cortical areas than in normal hearing adults. Over temporal and frontal areas peripheral stimuli presented at long intervals elicited N150 components which were larger in deaf than in hearing subjects. Over occipital and parietal areas peripheral and foveal stimuli elicited larger P230 components in deaf than in hearing subjects. These results imply that early auditory experience influences the organization of the human brain for visual processing.     

Orienting and maintenance of spatial attention in audition and vision: an event-related brain potential study

We examined the effects of orienting and maintenance of attention on performance and event-related brain potentials (ERPs) in audition and vision. Our subjects selectively attended to sounds or pictures in one location (Maintenance of attention) or alternated the focus of their auditory or visual attention between left and right locations (Orienting of attention) in order to detect and press a response button to infrequent targets among the attended stimuli. Reaction times were longer in the Auditory Orienting condition and hit rates were lower and false alarm rates higher in the Visual Orienting condition than in the corresponding Maintenance conditions. Comparison of ERPs to the attended and unattended stimuli in the Auditory and Visual Orienting and Maintenance conditions revealed attention-related modulations of ERPs. In each modality, ERPs to attended stimuli were negatively displaced in relation to unattended stimuli at 100-250 ms from stimulus onset. These negative differences (Nds) showed modality-specific distributions and they were larger over the hemisphere contralateral to the attended sounds and pictures than over the ipsilateral hemisphere. Moreover, the Nd was larger in the Auditory Orienting condition than in the Auditory Maintenance condition, while no such difference was observed in the visual modality. In addition to the Nd, attended visual stimuli elicited a late positive response (LPR) in both Orienting and Maintenance conditions. In contrast to our recent functional magnetic resonance imaging (fMRI) study employing the same experimental paradigm and indicating orienting-related activity in the frontal and parietal cortices, no ERP responses specifically related to orienting were found in either modality.     

Watching where you look: modulation of visual processing of foveal stimuli by spatial attention

Two experiments investigated the effect of sustained selective spatial attention upon the perceptual analysis of stimuli within the center of gaze. Spatial attention has typically been studied in relation to peripheral stimuli, and its relevance to the processing of central stimuli has remained relatively ignored. Here, behavioral measures in normal human volunteer participants showed that focused spatial attention can also influence responses to central stimuli, over and beyond the advantage conferred by their foveation. Event-related potentials (ERPs) showed that the action of attention upon foveal stimuli included the modulation of perceptual processing in extrastriate visual areas. Surprisingly, the visual modulation revealed an intriguing and consistent pattern of hemispheric asymmetry, in both experiments. These findings show that in addition to the established right hemisphere dominance of the brain areas that direct attention, the consequences of directed attention may also be asymmetrical.     

Neural mechanisms of global/local processing of bilateral visual inputs: an ERP study.

OBJECTIVE: Examine the neural mechanisms of global/local processing of multiple hierarchical stimuli. METHODS: Event-related brain potentials (ERPs) were recorded from adults who selectively attended to the global or local level of two compound letters that were simultaneously presented in the left and right visual fields, respectively. The compound stimuli were either broadband in spatial frequency (SF) spectrum or contrast balanced to remove low SFs. Subjects were asked to detect the presence of a global or local target that might appear in either the left or the right visual field in separate blocks of trials. RESULTS: Attention to the local level of broadband stimuli elicited a positivity over lateral occipital sites at 80-120 ms (P1) with larger amplitude than those in the global attention condition. However, global attention produced an enhanced positivity at 240-320 ms (P2) over lateral occipital sites relative to local attention. Both the P1 and P2 waves in the global condition were of larger amplitudes over the left than right hemispheres. Contrast balancing eliminated the P1 and P2 effects and modulated the hemispheric asymmetry of the long-latency occipital positivity. CONCLUSIONS: The results provide ERP evidence for modulations of neural activities in the visual cortex by global/local attention to concurrently presented multiple compound letters. Moreover, the modulation of brain activities by global/local attention depends upon the presence of low SFs in the compound stimuli. SIGNIFICANCE: The ERP results of this study contribute to the understanding of neural mechanisms of the processing of simultaneously-presented multiple compound stimuli.     

Event-related potentials in children with specific visual cognitive disability

Event-related potentials (ERPs) elicited by novel and well memorized non-verbal visual patterns were recorded over the right and left hemispheres from ten children with Specific Visual Cognitive Disability (SVCD) and ten matched Control subjects (C). ERPs of SVCD children were generally longer in latency (particularly N2) and decreased in amplitude (particularly P3) relative to the ERPs of Cs. No hemisphere differences were observed in the SVCD group, while the latency of N2 in the C group tended to be slightly shorter and the amplitude of P3 was significantly larger over the right than over the left hemisphere. The results are discussed in terms of deficits in visual learning and visual pattern analysis. The different hemispheric pattern in SVCD children as compared to Cs is suggested to be a symptom secondary to basic visual cognitive deficits.     

Auditory selective attention in middle-aged and elderly subjects: an event-related brain potential study.

Reaction times (RTs) and event-related brain potentials (ERPs) were recorded in middle-aged (MA) and elderly (ELD) subjects performing an auditory selective attention task. Subjects attended to tone bursts of a specified pitch and ear of delivery and responded to occasional longer duration target tones (75 vs. 25 msec). Infrequent novel stimuli (computer synthesized sounds and digitized environmental noises) were also included in the stimulus sequence. No significant age-related differences were found in the speed or accuracy of target detection. However, in both groups, RTs were delayed (by more than 300 msec) to targets that followed novel sounds. The prolongation was greater following novel sounds in the attended ear, particularly in the ELD group. The effects of selective attention on ERPs to standard tones were isolated as negative difference waves (Nds) by substracting ERPs to non-attended stimuli from ERPs to the same signals when attended. Nds had similar amplitudes, latencies of onset (60 msec), and distributions in ELD and MA groups. In both groups, Nd waves were more prominent following right ear stimulation, reflecting possible hemispheric asymmetries of generators in posterior temporal regions. The mismatch negativity (MMN) was isolated by subtracting ERPs to standard tones from ERPs to deviant stimuli. MMN amplitudes were reduced in the ELD group. There was also a significant change in MMN distribution with age: the MMN was larger over the right hemisphere for MA subjects but larger over the left for ELD subjects. Elderly subjects showed a trend toward smaller P3 amplitudes and delayed P3 latencies, but group differences did not reach statistical significance.(ABSTRACT TRUNCATED AT 250 WORDS)     

Which aspects of visual attention are changed by deafness? The case of the Attentional Network Test

The loss of one sensory modality can lead to a reorganization of the other intact sensory modalities. In the case of individuals who are born profoundly deaf, there is growing evidence of changes in visual functions. Specifically, deaf individuals demonstrate enhanced visual processing in the periphery, and in particular enhanced peripheral visual attention. To further characterize those aspects of visual attention that may be modified by deafness, deaf and hearing individuals were compared on the Attentional Network Test (ANT). The ANT was selected as it provides a measure of the efficiency of three neurally distinct subsystems of visual attention: alerting, orienting and executive control. The alerting measure refers to the efficiency with which a temporal cue is used to direct attention towards a target event, and the orienting measure is an indicator of the efficiency with which a spatial cue focuses attention upon that target's spatial location. The executive control measure, on the other hand, is an indicator of the amount of interference from peripheral flankers on processing that central target. In two separate experiments, deaf and hearing individuals displayed similar alerting and orienting abilities indicating comparable attention across populations. As predicted by enhanced peripheral attention, deaf subjects were found to have larger flanker interference effects than hearing subjects. These results indicate that not all aspects of visual attention are modified by early deafness, suggesting rather specific effects of cross-modal plasticity.     

A study of visual processes in a case of interhemispheric disconnexion

A 75-year-old right-handed woman, after a probable cerebral infarct, developed an irregular constriction of the visual fields, a left-sided agraphia, and an anomia for objects in the left hand. Subsequent testing demonstrated an inability to name, though ability to recognize, letters and objects flashed in the homonymous left visual field. An inter-hemispheric disconnexion syndrome was inferred from these findings. The present publication concerns mainly the visual aspects of this disconnexion syndrome.Tasks were devised to test the abilities of the major and minor hemisphere: (a) the left hemisphere demonstrated a complete dominance for language expression and an incomplete dominance for written language comprehension; (b) the right hemisphere appeared to be dominant for some visuo-spatial tasks including number comprehension; (c) when the hemispheres were given contradictory visual informations on a non-verbal task (chimeric stimuli) there was a predominance of the right hemisphere. The right hemisphere appeared able to process complex information. Specialization of functional activities in each hemisphere is briefly discussed.     

Changes in the spatial distribution of visual attention after early deafness

There is much anecdotal suggestion of improved visual skills in congenitally deaf individuals. However, this claim has only been met by mixed results from careful investigations of visual skills in deaf individuals. Psychophysical assessments of visual functions have failed, for the most part, to validate the view of enhanced visual skills after deafness. Only a few studies have shown an advantage for deaf individuals in visual tasks. Interestingly, all of these studies share the requirement that participants process visual information in their peripheral visual field under demanding conditions of attention. This work has led us to propose that congenital auditory deprivation alters the gradient of visual attention from central to peripheral field by enhancing peripheral processing. This hypothesis was tested by adapting a search task from Lavie and colleagues in which the interference from distracting information on the search task provides a measure of attentional resources. These authors have established that during an easy central search for a target, any surplus attention remaining will involuntarily process a peripheral distractor that the subject has been instructed to ignore. Attentional resources can be measured by adjusting the difficulty of the search task to the point at which no surplus resources are available for the distractor. Through modification of this paradigm, central and peripheral attentional resources were compared in deaf and hearing individuals. Deaf individuals possessed greater attentional resources in the periphery but less in the center when compared to hearing individuals. Furthermore, based on results from native hearing signers, it was shown that sign language alone could not be responsible for these changes. We conclude that auditory deprivation from birth leads to compensatory changes within the visual system that enhance attentional processing of the peripheral visual field.     

Electroencephalographic activity in a flanker interference task using Japanese orthography

The neural activities for color word interference effects were investigated using event-related brain potentials (ERPs) recorded in a flanker-type interference task. Kanji words (Japanese morphograms) and kana words (Japanese phonograms) were used as the flanker stimuli to obtain insights about hemispheric specialization for processing two types of Japanese orthographies. Interference effects in reaction time were larger when kanji words were presented in the left visual field and when kana words were in the right visual field. ERPs were modulated by the incongruent flankers, which generated a negative ERP component with the different onset and offset depending on flanker attributes. Consistent with the behavioral data, the interference-related negativity was observed for kanji words presented in the left visual field and for kana words in the right visual field. The negativity distributed maximally over the fronto-central site. The early part of the negativity distributed strongly over the frontal midline area, whereas it extended bilaterally over the frontal area in the late phase. The present results support the view of preferential processing of kanji in the right hemisphere and that of kana in the left hemisphere. The temporal profile of scalp topographies for the interference-related neural activity suggests that the medial and dorsolateral prefrontal regions may be involved in maintaining attentional set and conflict resolution.