The role of the magnocellular pathway in serial deployment of visual attention

Normal human visual function often demands detection of a target among a number of other objects cluttering the scene, such as when searching for a known face in a crowd. In these and similar tasks, the search performed is a serial one, with an attentional spotlight scanning the objects of the scene. We have investigated whether one of the afferent channels in vision, the colour-blind magnocellular pathway, is essential in such serial searches. We did this by using items that were isoluminant with the background but of a different colour, for which the magnocellular cells would be blind. The search in these conditions required much longer reaction times than when even a very small luminance contrast (2%) was added to the items. Because such luminance contrasts can be detected only by magnocellular cells and not by neurons of the other channels (parvocellular and koniocellular), the magnocellular pathway appears vitally important for serial search. In contrast, in a feature search task, which does not require allocation of attentional resources, the search was as efficient with isoluminance as when luminance contrast was added to the items.     

Parallel interhemispheric processing in hemineglect: Relation to visual field defects

Parallel interhemispheric processing is required to explore our visual environment and to integrate visual information from both hemifields simultaneously. Damage to the right temporo-parietal cortex can disrupt such parallel processes and result in neglect and visual extinction of stimuli in the left contralesional visual space. Neglected or extinguished stimuli can still be processed, yet without reaching the patient's awareness. Such unconscious processing has been attributed to structurally intact primary visual areas in neglect. To study whether unconscious parallel processing depends on visual functional integrity, we compared the performance of neglect patients with visual field defects (VFDs) (n = 11) and hemianopic patients with partial or complete blindness of one visual hemifield (n = 11) on redundant targets effects (RTE). The RTE manifests as faster reaction times to redundant paired (two stimuli, one in each hemifield) than single stimulation (in one hemifield). We found RTEs, i.e., unconscious processing, in neglect patients but not in hemianopic patients. Furthermore, neglect patients showed large crossed–uncrossed differences (CUDs), i.e., faster response times to ipsi- than contralesional hemifield stimulation, reflecting a difference in processing speed for single stimuli in the two hemispheres that were correlated with VFDs and visual extinction. The finding that extinction, but not RTE, was correlated with the CUD suggests that under competitive bilateral stimulus conditions the delayed contralesional visual field input may not be detected by the intact left hemisphere, which presumably mediates the task given the impairment of the right hemisphere. By contrast, unconscious parallel processing of contralesional stimuli (RTE) occurred even when contralesional visual field input is lacking (VFD) or delayed (CUD) and is possibly mediated via subcortical visual pathways.     

Contour-integration deficits on the intact side of the visual field in hemianopia patients

OBJECTIVE: Visual impairments in hemianopia are thought to be exclusively caused by the reduced visual field size. However, the primary lesion may affect the contralateral hemisphere through damage of interhemispheric projections. The question therefore arises if the presumed "intact" hemifield is perceptually impaired. METHODS: Three hemianopia patients and three matched controls carried out a Yes/No figure detection task with their intact side of the visual field. The figure (square) contours were composed of non-contiguous Gabor patches embedded in a random patch array of different background densities (low, delta=2; high, delta=1). Response accuracy and reaction times were recorded. RESULTS: A temporal-parietal patient revealed figure detection impairments, with accuracy rate, 77% (delta=2) and 53% (delta=1), below compared control values. An occipital patient was comparable to his match: 99% (delta=2); 84% (delta=1). Both patients exhibited frequent false alarms to random patterns and required longer presentation times to perform the task. In the third patient, with optic tract lesion, figure detection was nearly normal at low density (92%, delta=2) but impaired at noisy background (62%, delta=1). CONCLUSION: The "intact" visual field in hemianopes is impaired in detection of incomplete figures embedded in a noisy background. This deficit may be caused by damage to higher visual centers and/or loss of interhemispheric interactions.     

Rapid targeting followed by sustained deployment of visual spatial attention

We investigated preparatory attention processes when a spatial discrimination was required at a cued location, by measuring electroencephalography following a central symbolic cue to deploy spatial attention. Electroencephalography activity in response to the cue revealed three cue-related activations: an early-onset positivity following the P1 at posterior scalp sites contralateral to the cued location, followed by cue-related frontal scalp activity and later-onset sustained activity at posterior scalp sites contralateral to the cued location. The early contralateral positivity may reflect rapid targeting of the cued location. Our results also extend the findings of cue-related frontal activity followed by posterior activity contralateral to the cued location, found with nonspatial feature discriminations, to a task requiring a spatial discrimination.     

Efficacy of visuo-spatial training in right-brain damaged patients with spatial hemineglect and attention disorders

Several rehabilitative approaches have been used to reduce neglect disorders. Some studies tried to demonstrate that hemineglect can be ameliorated by using tasks promoting attentional activation towards the neglected hemispace (Robertson et al., 1995, 2001). As a consequence, a functional link between level of attention and disorders of space exploration has been proposed. For this reason we tried to explore the possible role of attentional deficits on the efficacy of a standard neglect treatment based on visuo spatial rehabilitation. In this study we then examined the performances of a selected group of seven right brain damaged patients, suffering from both severe hemineglect and attentional deficits (of both phasic and tonic components of attention), to several tasks before and after a visuo-spatial training (VST) whose efficacy has been already demonstrated (Pizzamiglio et al., 1992). Results showed that VST induces a significant remission of hemineglect symptoms independently from disorders of attention. Moreover, after visuo-spatial rehabilitation, no improvement of attentional deficits is detectable.     

A distortion of perceived space in patients with right-hemisphere lesions and visual hemineglect.

In twelve patients with left visuospatial hemineglect following acute right hemisphere cerebrovascular accident (CVA), and in twelve normal controls, we used a matching task to investigate judgement of the length of lines. Their task was to choose which of three lines of different length matched a separate single line. On each trial, the lines were all vertical or all horizontal. The set of three lines was presented either to the left or the right of the single line. When the lines were vertical, the choices made both by normal subjects and patients were nearly always correct. However, when the lines were horizontal and the set of comparison lines was on the left, patients mostly selected a line that was longer than the sample on the right, and never chose shorter lines, suggesting that the length of horizontal lines on the left was underestimated. When the set of three lines was on the right, patients chose the correct line more often, but made errors in both directions, i.e. longer or shorter, although shorter lines were chosen more often than longer lines. The rare errors made by normal subjects were in the same directions as those observed in the patients. The results provide further evidence for a non-Euclidean distortion of perceptual space in patients with left sided visuospatial neglect, and suggest that this distortion could be a gross exaggeration of normal performance or a product of diminished spatial attention, or both.     

Game theoretical mapping of white matter contributions to visuospatial attention in stroke patients with hemineglect

White matter bundles linking gray matter nodes are key anatomical players to fully characterize associations between brain systems and cognitive functions. Here we used a multivariate lesion inference approach grounded in coalitional game theory (multiperturbation Shapley value analysis, MSA) to infer causal contributions of white matter bundles to visuospatial orienting of attention. Our work is based on the characterization of the lesion patterns of 25 right hemisphere stroke patients and the causal analysis of their impact on three neuropsychological tasks: line bisection, letter cancellation, and bells cancellation. We report that, out of the 11 white matter bundles included in our MSA coalitions, the optic radiations, the inferior fronto‐occipital fasciculus and the anterior cingulum were the only tracts to display task‐invariant contributions (positive, positive, and negative, respectively) to the tasks. We also report task‐dependent influences for the branches of the superior longitudinal fasciculus and the posterior cingulum. By extending prior findings to white matter tracts linking key gray matter nodes, we further characterize from a network perspective the anatomical basis of visual and attentional orienting processes. The knowledge about interactions patterns mediated by white matter tracts linking cortical nodes of attention orienting networks, consolidated by further studies, may help develop and customize brain stimulation approaches for the rehabilitation of visuospatial neglect.     

A unilateral defect of short-term visual memory in left hemineglect

Normal subjects and right- and left-brain-damaged patients with and without neglect performed a task of immediate visual memory and a task of same-different judgment on visual items. In both tasks, stimull consisted in geometrical patterns, which differed either on the right or on the left side. Only right-brain-damaged patients with neglect showed an impaired performance in these tasks. They performed worse on items differing on the left side than on items differing on the right side. This impairment was more severe on the memory task than on the same-different task, suggesting that a lateralized visual memory deficit contributes to the behavioural manifestations of left hemineglect.     

A psychophysical study of visual extinction: ipsilesional distractor interference with contralesional orientation thresholds in visual hemineglect patients

Visual extinction was investigated in left (n=15) and right (n=25) brain-damaged patients with or without visual neglect, and in normal control subjects (n=14), using a psychophysical paradigm. Orientation discrimination thresholds were determined for both left and right hemifield gratings presented either in isolation or simultaneously with a contralateral distractor grating. To minimize the influence of possible sensory-perceptual deficits, the luminances of both target and distractor gratings were chosen to be 20 times the luminances necessary to discriminate between horizontal and vertical grating orientations. The location of the target grating was always cued, making the distractor grating task irrelevant. Even after equalizing the visibility of left and right hemifield stimuli, neglect patients still displayed an increased interference effect from an ipsilesional distractor (and no interference from a contralesional distractor). Left or right brain-damaged controls did not show this asymmetric interference of irrelevant distractors, even the patients who demonstrated extinction on standard extinction testing. This suggests that visual extinction is a critical component of the visual neglect syndrome and that it involves an attentional deficit.     

The topography of visuospatial attention as revealed by a novel visual field mapping technique

Previously, we and others have shown that attention can enhance visual processing in a spatially specific manner that is retinotopically mapped in the occipital cortex. However, it is difficult to appreciate the functional significance of the spatial pattern of cortical activation just by examining the brain maps. In this study, we visualize the neural representation of the "spotlight" of attention using a back-projection of attention-related brain activation onto a diagram of the visual field. In the two main experiments, we examine the topography of attentional activation in the occipital and parietal cortices. In retinotopic areas, attentional enhancement is strongest at the locations of the attended target, but also spreads to nearby locations and even weakly to restricted locations in the opposite visual field. The dispersion of attentional effects around an attended site increases with the eccentricity of the target in a manner that roughly corresponds to a constant area of spread within the cortex. When averaged across multiple observers, these patterns appear consistent with a gradient model of spatial attention. However, individual observers exhibit complex variations that are unique but reproducible. Overall, these results suggest that the topography of visual attention for each individual is composed of a common theme plus a personal variation that may reflect their own unique "attentional style."     

The contribution of the 'second' visual system to directed visual attention in man

The visual functions of a patient suffering from a brain lesion incorporating the left n. pulvinar was examined in order to assess the contribution of this structure to human vision. With the exception of the following abnormalities visual functions were normal. First, there was a decrease in the critical flicker frequency in the periphery (but not in the parafoveal region) of the right visual hemifield, that is, that contralateral to the pulvinar lesion. However, the second and most striking characteristic was--as shown by presenting visual stimuli bilaterally and simultaneously-a 'neglect' for the periphery of this contralateral visual half-field. This 'neglect' was a function of (a) position in the visual field (eccentricity), (b) stimulus properties (size and luminance), (c) temporal properties (length of presentation and interstimulus interval). In addition to this reduced capacity to detect stimuli appearing in the periphery of the right visual hemifield, there was also prolonged latency of visually evoked saccadic eye movements and a paucity of spontaneous eye movements directed towards the right visual hemifield. These results are interpreted in terms of a contribution of the n. pulvinar to the detection of light stimuli presented in the periphery of the visual field, and support the view that the tectopulvinar extrastriate visual pathway plays an important role in the control of visual attention.     

The gradient of visual attention in developmental dyslexia

This study investigated the gradient of visual attention in 21 children, 11 children with specific reading disorder (SRD) or dyslexia and 10 children with normal reading skills. We recorded reaction times (RTs) at the onset of a small point along the horizontal axis in the two visual fields. In 70% of the cases the target appeared inside a circle acting as focusing cue and in 30% of the cases it appeared outside, allowing us to study the distribution of attentional resources outside the selected area. Normally reading children showed a normal symmetric distribution of attention. Indeed, RTs were directly proportional to the eccentricity of the target, and no visual field effect was observable. In contrast, children with SRD showed an anomalous and asymmetric distribution. The effect of the target eccentricity influenced RTs only when the stimulus was projected in the left visual field, whereas no effect was observable when the stimulus was projected in the right visual field. Findings allowed us to discuss the relation between this anomalous spatial distribution of visual attentional resources and dyslexia. To interpret the visual perceptual difficulties of children with SRD the hypothesis was made of a selective disorder of spatial attention (left inattention and right over-distractibility) related to a right parietal cortex dysfunction.     

Deficits in peripheral visual attention in patients with optic ataxia

Earlier research has suggested that optic ataxia, a deficit in reaching in peripheral vision, can be isolated from Balint's syndrome as it is primarily a visuomotor disorder, independent of perceptual or attentional deficits. Yet almost no research has examined the attentional abilities of these patients. We examined peripheral visual attention in two patients with unilateral optic ataxia. Results indicated that both patients were slower to respond to targets in their ataxic visual field, irrespective of cuing condition (i.e. validly, invalidly, and no cue conditions), consistent with an overall decrease in the salience of stimuli in the ataxic field. Attentional deficits in peripheral vision are therefore an important factor to consider when examining visuomotor control deficits in optic ataxia.     

Oculographic diagnosis of hemineglect in patients with homonymous hemianopia

Summary Two patients with complete dense left homonymous hemianopia of acute onset are presented. There was a striking difference in visuospatial orientation between them in the early stages of their disease, due to additional left hemineglect in the second patient. While eye movement patterns known to be typical for homonymous hemianopia were found in the first patient by means of infrared reflection oculography, the second showed several findings which could be attributed to hemineglect. Based on these observations and those described in earlier literature, oculographic criteria are listed that enable hemineglect to be detected in the presence of homonymous hemianopia. The practical importance of diagnosing hemineglect with objective and quantitative methods is stressed.     

The visual attention span deficit in dyslexia is visual and not verbal

The visual attention (VA) span deficit hypothesis of dyslexia posits that letter string deficits are a consequence of impaired visual processing. Alternatively, some have interpreted this deficit as resulting from a visual-to-phonology code mapping impairment. This study aims to disambiguate between the two interpretations by investigating performance in a non-verbal character string visual categorization task with verbal and non-verbal stimuli. Results show that VA span ability predicts performance for the non-verbal visual processing task in normal reading children. Furthermore, VA span impaired dyslexic children are also impaired for the categorization task independently of stimuli type. This supports the hypothesis that the underlying impairment responsible for the VA span deficit is visual, not verbal.     

Feature-based attention in visual cortex

Although most studies of visual attention have examined the effects of shifting attention between different locations in the visual field, attention can also be directed to particular visual features, such as a color, orientation or a direction of motion. Single-unit studies have shown that attention to a feature modulates neuronal signals in a range of areas in monkey visual cortex. The location-independent property of feature-based attention makes it particularly well suited to modify selectively the neural representations of stimuli or parts within complex visual scenes that match the currently attended feature. This review is part of the TINS special issue on The Neural Substrates of Cognition.     

Dynamic upper and lower visual field preferences within the human dorsal frontoparietal attention network

Both in nonhuman primates and in humans, behavioral differences between the upper and lower visual field have been identified in distinct subprocesses of attention. Advantages of the lower field have been explained by its higher spatial resolution; those of the upper field by its higher efficiency in attentional shifting. The physiological basis of visual field asymmetries within in the frontoparietal attention network (FPN) remains unclear. This study investigates the physiological correlates of upper and lower field preferences within the FPN using event‐related functional magnetic resonance imaging. The paradigm separated two attentional subprocesses during a visual search task. Whether in the upper or lower field, the attention of subjects was first directed at stationary locations (spatial orienting) and then shifted between locations to search for a target (visual search) in easy or difficult search displays. Depending on the task phase (spatial orienting vs. easy visual search), upper and lower visual field preferences in the FPN changed. The analysis revealed a lower field preference during stationary spatial orienting and an upper field preference during visual search. We conclude that also higher areas represent upper and lower visual field asymmetries depending on distinct subcomponents of visuospatial attentional processing. Hum Brain Mapp, 2011. © 2010 Wiley‐Liss, Inc.