Recognition of visual images by separate hemispheres in conditions of masked blocking of interhemispheric transmission

The subjects learned to recognize three figures presented in the left visual hemifield and three figures presented in the right visual hemifield. During presentation of a stimulus, the contralateral hemifield was overlapped by a mask. After the training, recognition of all six figures presented in the right and left visual hemifields, was compared. Each hemisphere recognizes figures which were learned in the corresponding visual hemifield, but the recognition of figures learned in the opposite visual hemifield was poor. Thus, the ability of the hemispheres to act separately in recognizing different sets of visual images, was established.     

Model approach to neurological variants of visuo-spatial neglect

Neglect is a neurological disorder of spatial attention with reduced awareness of visual stimuli in the hemifield contralateral to an acute temporo-parietal lesion mainly of the right hemisphere. There is a close association of multisensory orientation centers (MSO) and vestibular tonus imbalance. A lesion of the dominant right MSO causes a left-sided neglect due to a lack of ipsilateral activation of the visual cortex, which is further enhanced by increasing inhibition from the contralateral visual cortex. The nondominant MSO in the left hemisphere might be involved in the manifestation of the less frequent and more transient right-sided neglect and in the plastic mechanisms of gradual recovery from left-sided neglect or extinction. There is evidence that a vestibular tonus inbalance due to peripheral or central vestibular pathway lesions may also induce a neglect. In a first model approach using an attractor network and assuming that there is only one MSO in the right hemisphere, it is possible to simulate attentional shifts into a visual hemifield and to induce a neglect. The neural network model consists of four layers of neurons: retina, MSO, visual cortex V1, and superior colliculus. The superior colliculus layer is modeled as a recurrent attractor network with one inhibitory interneuron and synaptic weights chosen to implement a winner-take-all network that centers the hill of activity on the strongest input. We are well aware of the simplifications used in the conceptual drawings and the computational model, but nevertheless hope that they will serve as an inspiration for further modeling and clinical studies.     

Abnormal attentional modulation of retinotopic cortex in parietal patients with spatial neglect

Brain regions beyond visual cortex are thought to be responsible for attention-related modulation of visual processing [1, 2], but most evidence is indirect. Here, we applied functional magnetic resonance imaging (fMRI), including retinotopic mapping of visual areas, to patients with focal right-parietal lesions and left spatial neglect [3, 4]. When attentional load at fixation was minimal, retinotopic areas in right visual cortex showed preserved responses to task-irrelevant checkerboards in the contralateral left hemifield, analogously to left visual cortex for right-hemifield checkerboards, indicating a "symmetric" pattern in both hemispheres with respect to contralateral stimulation under these conditions. But when attentional load at fixation was increased, a functional asymmetry emerged for visual cortex, with contralateral responses in right visual areas being pathologically reduced (even eliminated for right V4/TEO), whereas left visual areas showed no such reduction in their contralateral response. These results reveal attention-dependent abnormalities in visual cortex after lesions in distant (parietal) regions. This may explain otherwise puzzling aspects of neglect [5, 6], as confirmed here by additional behavioral testing.     

Eye position affects orienting of visuospatial attention

The ability to detect an incoming visual stimulus is enhanced by knowledge of stimulus location (orienting of visuospatial attention). Although the brain mechanisms at the basis of this enhancement are not yet fully clarified, there is evidence that orienting of attention is accompanied by the activation of oculomotor circuits. It remains unclear, however, whether this oculomotor activity is an epiphenomenon or is functionally related to the attentional process. Attentional benefits are usually measured by the classical Posner paradigm. When subjects fixate centrally and are requested to detect a visual stimulus that could appear in an attended or unattended location, they react faster to stimuli appearing in the attended one. Here, we demonstrate that in monocular vision visuospatial attention was significantly modulated by the position of the eye in the orbit. When the screen was placed 40 degrees to the right or to the left of subjects' sagittal plane, attentional benefits for stimuli appearing in subjects' temporal spatial hemifield dramatically decayed, even if the retinal stimulation was exactly the same as in the classical paradigm. The finding that eyes and attention show a common limit stop point supports their close functional coupling.     

Distractor-target interactions during directed visual attention

The spatial extent of directed visual attention (DVA) was examined in a series of experiments using precuing in a suprathreshold luminance detection (reaction time) paradigm. Previous findings (Hughes, H. C. and Zimba, L. D. J. Exp. Psychol.; Human Percept Perf., 1985, 11, 409-430) indicated that, in an empty visual field, the effects of DVA were primarily manifest as a uniform elevation of response times to all probe targets in the hemifield contralateral to the observer's expectancy. The present experiments were designed to determine whether increased spatial selectivity could be found when luminous markers indicated the exact location of the expected visual target. To maintain equivalent states of adaptation in both hemifields, luminous markers were also present at the same location in the contralateral hemifield. In general, hemifield effects were again obtained, but with two notable exceptions. First, marking locations in the unattended hemifield produced a local increase (enhanced interference) in RTs above the level characteristic of other locations within that hemifield. Second, when multiple locations were indicated with identical luminous markers, graded costs were obtained in both hemifields. However, scaling the markers according to estimates of cortical magnification factor (M) substantially reduced the slope of these inhibitory gradients, and the results once again approached those characteristic of an unstructured visual field. The findings suggest that when attention is directed to a marked location along the horizontal meridian, a transition in performance typically occurs at the vertical meridian. In addition, irrelevant stimuli some distance from the attentional focus interfere with detection times to unexpected targets that appear in the same vicinity. This interference may relate to an enhanced susceptibility to spatial interactions between the distractors and target away from the attentional focus. The interference appears to extend over a constant area of visual cortex, since it is reduced when the markers are M-scaled.     

Bilateral field advantage in visual enumeration

A number of recent studies have demonstrated superior visual processing when the information is distributed across the left and right visual fields than if the information is presented in a single hemifield (the bilateral field advantage). This effect is thought to reflect independent attentional resources in the two hemifields and the capacity of the neural responses to the left and right hemifields to process visual information in parallel. Here, we examined whether a bilateral field advantage can also be observed in a high-level visual task that requires the information from both hemifields to be combined. To this end, we used a visual enumeration task--a task that requires the assimilation of separate visual items into a single quantity--where the to-be-enumerated items were either presented in one hemifield or distributed between the two visual fields. We found that enumerating large number (>4 items), but not small number (<4 items), exhibited the bilateral field advantage: enumeration was more accurate when the visual items were split between the left and right hemifields than when they were all presented within the same hemifield. Control experiments further showed that this effect could not be attributed to a horizontal alignment advantage of the items in the visual field, or to a retinal stimulation difference between the unilateral and bilateral displays. These results suggest that a bilateral field advantage can arise when the visual task involves inter-hemispheric integration. This is in line with previous research and theory indicating that, when the visual task is attentionally demanding, parallel processing by the neural responses to the left and right hemifields can expand the capacity of visual information processing.     

Spatial attention and neglect: parietal, frontal and cingulate contributions to the mental representation and attentional targeting of salient extrapersonal events.

The syndrome of contralesional neglect reflects a lateralized disruption of spatial attention. In the human, the left hemisphere shifts attention predominantly in the contralateral hemispace and in a contraversive direction whereas the right hemisphere distributes attention more evenly, in both hemispaces and both directions. As a consequence of this asymmetry, severe contralesional neglect occurs almost exclusively after right hemisphere lesions. Patients with left neglect experience a loss of salience in the mental representation and conscious perception of the left side and display a reluctance to direct orientating and exploratory behaviours to the left. Neglect is distributed according to egocentric, allocentric, world-centred, and object-centred frames of reference. Neglected events can continue to exert an implicit influence on behaviour, indicating that the attentional filtering occurs at the level of an internalized representation rather than at the level of peripheral sensory input. The unilateral neglect syndrome is caused by a dysfunction of a large-scale neurocognitive network, the cortical epicentres of which are located in posterior parietal cortex, the frontal eye fields, and the cingulate gyrus. This network coordinates all aspects of spatial attention, regardless of the modality of input or output. It helps to compile a mental representation of extrapersonal events in terms of their motivational salience, and to generate 'kinetic strategies' so that the attentional focus can shift from one target to another.     

Effects of Multimodal Load on Spatial Monitoring as Revealed by ERPs

While the role of selective attention in filtering out irrelevant information has been extensively studied, its characteristics and neural underpinnings when multiple environmental stimuli have to be processed in parallel are much less known. Building upon a dual-task paradigm that induced spatial awareness deficits for contralesional hemispace in right hemisphere-damaged patients, we investigated the electrophysiological correlates of multimodal load during spatial monitoring in healthy participants. The position of appearance of briefly presented, lateralized targets had to be reported either in isolation (single task) or together with a concurrent task, visual or auditory, which recruited additional attentional resources (dual-task). This top-down manipulation of attentional load, without any change of the sensory stimulation, modulated the amplitude of the first positive ERP response (P1) and shifted its neural generators, with a suppression of the signal in the early visual areas during both visual and auditory dual tasks. Furthermore, later N2 contralateral components elicited by left targets were particularly influenced by the concurrent visual task and were related to increased activation of the supramarginal gyrus. These results suggest that the right hemisphere is particularly affected by load manipulations, and confirm its crucial role in subtending automatic orienting of spatial attention and in monitoring both hemispaces.     

The Spatial and Temporal Deployment of Voluntary Attention across the Visual Field

Several studies have addressed the question of the time it takes for attention to shift from one position in space to another. Here we present a behavioural paradigm which offers a direct access to an estimate of voluntary shift time by comparing, in the same task, a situation in which subjects are required to re-engage their attention at the same spatial location with a situation in which they need to shift their attention to another location, all other sensory, cognitive and motor parameters being equal. We show that spatial attention takes on average 55 ms to voluntarily shift from one hemifield to the other and 38 ms to shift within the same hemifield. In addition, we show that across and within hemifields attentional processes are different. In particular, attentional spotlight division appears to be more difficult to operate within than across hemifields.     

Visual hemifield differences in recognition of kanji and hiragana and its relation to hemispheric cerebral asymmetries

Visual hemifield differences in recognition of kanji and hiragana were studied on forty male right handers. A letter of kanji or hiragana was presented unilaterally to the right or left visual hemifield on a CRT display for 123 msec. A hundred and twenty recognition trials were performed for each subject using 20 well-acquainted kanji, 20 unfamiliar kanji and 20 hiragana. Kanji was more accurately recognized in the left visual hemifield than in the right hemifield. This tendency was more prominent in unfamiliar kanji compared with well-acquainted kanji. There were no visual hemifield differences in recognition of hiragana. Learning effects were observed for the right hemifield on kanji and both hemifields on hiragana. The results were discussed in relation to cerebral asymmetries of function. Kanji might be processed in the right cerebral hemisphere as geometric forms. The results on hiragana may be explained by mental set. It is suggested that modes of processing may be different between kanji and hiragana.     

Cerebral Asymmetry of fMRI-BOLD Responses to Visual Stimulation

Hemispheric asymmetry of a wide range of functions is a hallmark of the human brain. The visual system has traditionally been thought of as symmetrically distributed in the brain, but a growing body of evidence has challenged this view. Some highly specific visual tasks have been shown to depend on hemispheric specialization. However, the possible lateralization of cerebral responses to a simple checkerboard visual stimulation has not been a focus of previous studies. To investigate this, we performed two sessions of blood-oxygenation level dependent (BOLD) functional magnetic resonance imaging (fMRI) in 54 healthy subjects during stimulation with a black and white checkerboard visual stimulus. While carefully excluding possible non-physiological causes of left-to-right bias, we compared the activation of the left and the right cerebral hemispheres and related this to grey matter volume, handedness, age, gender, ocular dominance, interocular difference in visual acuity, as well as line-bisection performance. We found a general lateralization of cerebral activation towards the right hemisphere of early visual cortical areas and areas of higher-level visual processing, involved in visuospatial attention, especially in top-down (i.e., goal-oriented) attentional processing. This right hemisphere lateralization was partly, but not completely, explained by an increased grey matter volume in the right hemisphere of the early visual areas. Difference in activation of the superior parietal lobule was correlated with subject age, suggesting a shift towards the left hemisphere with increasing age. Our findings suggest a right-hemispheric dominance of these areas, which could lend support to the generally observed leftward visual attentional bias and to the left hemifield advantage for some visual perception tasks.     

Amplitude asymmetry of hemifield pattern reversal VEPs in healthy subjects

The amplitudes of transient and steady-state visual evoked potentials (VEPs) were measured during hemifield stimulation of the left eye in 10 healthy adults. Pattern reversal of a checkerboard was produced at 4 stimulation frequencies: 1, 5, 10 and 15 Hz. The amplitudes of pattern VEPs were evaluated using the paired t test to determine significant differences between right and left hemifields. The transient VEP amplitudes from midoccipital, midparietal, ipsilateral occipital and contralateral occipital electrodes were significantly greater with right hemifield stimulation. The steady-state VEP amplitudes from the midoccipital electrode during 15 Hz stimulation were significantly greater with right hemifield stimulation. Our neurophysiological data may be compatible with neuroanatomical asymmetries of the occipital lobes in humans.     

Modelling Visual Neglect: Computational Insights into Conscious Perception

Background

Visual neglect is an attentional deficit typically resulting from parietal cortex lesion and sometimes frontal lesion. Patients fail to attend to objects and events in the visual hemifield contralateral to their lesion during visual search.

Methodology/Principal Finding

The aim of this work was to examine the effects of parietal and frontal lesion in an existing computational model of visual attention and search and simulate visual search behaviour under lesion conditions. We find that unilateral parietal lesion in this model leads to symptoms of visual neglect in simulated search scan paths, including an inhibition of return (IOR) deficit, while frontal lesion leads to milder neglect and to more severe deficits in IOR and perseveration in the scan path. During simulations of search under unilateral parietal lesion, the model''s extrastriate ventral stream area exhibits lower activity for stimuli in the neglected hemifield compared to that for stimuli in the normally perceived hemifield. This could represent a computational correlate of differences observed in neuroimaging for unconscious versus conscious perception following parietal lesion.

Conclusions/Significance

Our results lead to the prediction, supported by effective connectivity evidence, that connections between the dorsal and ventral visual streams may be an important factor in the explanation of perceptual deficits in parietal lesion patients and of conscious perception in general.     

Spring‐autumn variations in hemispheric asymmetry

Abstract

48 right‐handed subjects performed verbal and spatial hemifield tachistoscopic tasks, half of them during spring and the other half during autumn. Results showed increased visual field advantage both in performance speed and accuracy in the right visual field during spring and in the left visual field during autumn. This suggests a seasonal‐hemispheric shift favouring the left hemisphere during spring and the right hemisphere during autumn, and provides a clue to understanding seasonal variations in mood and affective states as related to variations in cognitive style.     

Modality-specific control of strategic spatial attention in parietal cortex

The neural basis of selective spatial attention presents a significant challenge to cognitive neuroscience. Recent neuroimaging studies have suggested that regions of the parietal and temporal cortex constitute a "supramodal" network that mediates goal-directed attention in multiple sensory modalities. Here we used transcranial magnetic stimulation (TMS) to determine which cortical subregions control strategic attention in vision and touch. Healthy observers undertook an orienting task in which a central arrow cue predicted the location of a subsequent visual or somatosensory target. To determine the attentional role of cortical subregions at different stages of processing, TMS was delivered to the right hemisphere during cue or target events. Results indicated a critical role of the inferior parietal cortex in strategic orienting to visual events, but not to somatosensory events. These findings are inconsistent with the existence of a supramodal attentional network and instead provide direct evidence for modality-specific attentional processing in parietal cortex.     

Sleep wakefulness cycle

Abstract

48 right‐handed subjects performed verbal and spatial hemifield tachistoscopic tasks, half of them in spring and the other half in autumn. Significant differences were detected showing increased right visual field (left hemisphere) advantage in spring and increased left visual field (right hemisphere) advantage in autumn. Results may help to explain the reported seasonal propensity towards elevated mood in spring and towards depressed mood in autumn as related to the relative role of both hemispheres.     

A preference for contralateral stimuli in human object- and face-selective cortex

Visual input from the left and right visual fields is processed predominantly in the contralateral hemisphere. Here we investigated whether this preference for contralateral over ipsilateral stimuli is also found in high-level visual areas that are important for the recognition of objects and faces. Human subjects were scanned with functional magnetic resonance imaging (fMRI) while they viewed and attended faces, objects, scenes, and scrambled images in the left or right visual field. With our stimulation protocol, primary visual cortex responded only to contralateral stimuli. The contralateral preference was smaller in object- and face-selective regions, and it was smallest in the fusiform gyrus. Nevertheless, each region showed a significant preference for contralateral stimuli. These results indicate that sensitivity to stimulus position is present even in high-level ventral visual cortex.     

Crossmodal spatial influences of touch on extrastriate visual areas take current gaze direction into account

Recent results indicate that crossmodal interactions can affect activity in cortical regions traditionally regarded as "unimodal." Previously we found that combining touch on one hand with visual stimulation in the anatomically corresponding hemifield could boost responses in contralateral visual cortex. Here we manipulated which visual hemifield corresponded to the location of the stimulated hand, by changing gaze direction such that right-hand touch could now arise in either the left or right visual field. Crossmodal effects on visual cortex switched from one hemisphere to the other, depending on gaze direction, regardless of whether the hand was seen. This indicates that crossmodal influences of touch upon visual cortex depend on spatial alignment for the multimodal stimuli, with gaze posture taken into account.     

Saccade latency during probability presentation of the visual targets in man

The latent periods of saccadic eye movements in response to peripheral visual stimuli were measured in 8 right-handed healthy subjects using Posner's paradigm "COST-BENEFIT". In 6 subjects, the saccade latency in response to visual target presented in expected location in valid condition was shorter than that in neutral condition ("benefit"). Increase in saccade latency in response to the visual target presented in unexpected location in valid condition versus neutral condition took place only in 4 subjects ("cost"). A decrease in left-directed saccade latency in response to expected target presented in the left hemifield and increase in saccade latency in response to unexpected left target in comparison with analogous right-directed saccades were observed in valid condition. This phenomenon can be explained by the dominance of the right hemisphere in the processes of spatial orientation and "disengage" of attention.     

Transfer of Learning between Hemifields in Multiple Object Tracking: Memory Reduces Constraints of Attention

Many tasks involve tracking multiple moving objects, or stimuli. Some require that individuals adapt to changing or unfamiliar conditions to be able to track well. This study explores processes involved in such adaptation through an investigation of the interaction of attention and memory during tracking. Previous research has shown that during tracking, attention operates independently to some degree in the left and right visual hemifields, due to putative anatomical constraints. It has been suggested that the degree of independence is related to the relative dominance of processes of attention versus processes of memory. Here we show that when individuals are trained to track a unique pattern of movement in one hemifield, that learning can be transferred to the opposite hemifield, without any evidence of hemifield independence. However, learning is not influenced by an explicit strategy of memorisation of brief periods of recognisable movement. The findings lend support to a role for implicit memory in overcoming putative anatomical constraints on the dynamic, distributed spatial allocation of attention involved in tracking multiple objects.