Prestriate Cortex Research Articles

Prediction, Suppression of Visual Response, and Modulation of Visual Perception: Insights From Visual Evoked Potentials and Representational Momentum.

When a visual object changes its position along with certain sequential regularities, the visual system rapidly and automatically forms a prediction regarding the future position of the object based on the regularities. Such prediction can drastically alter visual perception. A phenomenon called representational momentum (RM: a predictive displacement of the perceived final position of a visual object along its recent regular pattern) has provided extensive evidence for the predictive modulation of visual perception. The purpose of the present study was to identify neural effects that could explain individual differences in the strength of the predictive modulation of visual perception as measured by RM. For this purpose, in two experiments with a conventional RM paradigm where a bar was discretely presented in a regular rotation manner (with a step of 18° in Experiment 1 and a step of 20° in Experiment 2), visual evoked potentials (VEPs) in response to the regularly rotated bar were measured, and correlations between the magnitudes of RM and VEPs were examined. The results showed that the magnitudes of RM and central P2 were negatively correlated, consistently in both experiments; participants who showed a smaller central P2 tended to exhibit greater RM. Together with a previous proposal that central P2 would represent delayed reactivation of lower visual areas around the striate and prestriate cortices via reentrant feedback projections from higher areas, the present results suggest that greater suppression of delayed reactivation of lower visual areas (as indicated by smaller central P2) may underlie stronger predictive modulation of visual perception (as indicated by greater RM).

Early visual cortical responses produced by checkerboard pattern stimulation

Visual evoked potentials have been traditionally triggered with flash or reversing checkerboard stimuli and recorded with electroencephalographic techniques, largely but not exclusively in clinical or clinically related settings. They have been crucial in determining the healthy functioning or otherwise of the visual pathways up to and including the cerebral cortex. They have typically given early response latencies of 100ms, the source of which has been attributed to V1, with the prestriate cortex being secondarily activated somewhat later. On the other hand, magnetoencephalographic studies using stimuli better tailored to the physiology of individual, specialized, visual areas have given early latencies of <50ms with the sources localized in both striate (V1) and prestriate cortex. In this study, we used the reversing checkerboard pattern as a stimulus and recorded cortical visual evoked magnetic fields with magnetoencephalography, to establish whether very early responses can be traced to (estimated) in both striate and prestriate cortex, since such a demonstration would enhance considerably the power of this classical approach in clinical investigations. Our results show that cortical responses evoked by checkerboard patterns can be detected before 50ms post-stimulus onset and that their sources can be estimated in both striate and prestriate cortex, suggesting a strong parallel input from the sub-cortex to both striate and prestriate divisions of the visual cortex.

Differentiation of neuronal operations in latent components of event‐related potentials in delayed match‐to‐sample tasks

The goal of this study was to decompose ERPs into latent components associated with hypothetical processes of category discrimination, comparison to working memory and action-related operations. In five variants of the delayed match-to-sample s1-s2 task, instructions were varied for manipulation of the processes. The blind source separation was applied to the collection of ERPs. The category discrimination operation is attributed to three latent components with peak latencies of 130 to 170 ms, which are generated in different parts of the prestriate cortex. The comparison to working memory operation is attributed to a latent component that is generated in the temporal cortex and manifested in a positive deflection with a peak latency of 250 ms after s2. The category discrimination and comparison to working memory effects were dissociated spatially and temporally from attention and action selection effects.

Parallelism in the brain's visual form system.

We used magnetoencephalography (MEG) to determine whether increasingly complex forms constituted from the same elements (lines) activate visual cortex with the same or different latencies. Twenty right-handed healthy adult volunteers viewed two different forms, lines and rhomboids, representing two levels of complexity. Our results showed that the earliest responses produced by lines and rhomboids in both striate and prestriate cortex had similar peak latencies (40 ms) although lines produced stronger responses than rhomboids. Dynamic causal modeling (DCM) showed that a parallel multiple input model to striate and prestriate cortex accounts best for the MEG response data. These results lead us to conclude that the perceptual hierarchy between lines and rhomboids is not mirrored by a temporal hierarchy in latency of activation and thus that a strategy of parallel processing appears to be used to construct forms, without implying that a hierarchical strategy may not be used in separate visual areas, in parallel.

Functional specialization and generalization for grouping of stimuli based on colour and motion.

This study was undertaken to learn whether the principle of functional specialization that is evident at the level of the prestriate visual cortex extends to areas that are involved in grouping visual stimuli according to attribute, and specifically according to colour and motion. Subjects viewed, in an fMRI scanner, visual stimuli composed of moving dots, which could be either coloured or achromatic; in some stimuli the moving coloured dots were randomly distributed or moved in random directions; in others, some of the moving dots were grouped together according to colour or to direction of motion, with the number of groupings varying from 1 to 3. Increased activation was observed in area V4 in response to colour grouping and in V5 in response to motion grouping while both groupings led to activity in separate though contiguous compartments within the intraparietal cortex. The activity in all the above areas was parametrically related to the number of groupings, as was the prominent activity in Crus I of the cerebellum where the activity resulting from the two types of grouping overlapped. This suggests (a) that, the specialized visual areas of the prestriate cortex have functions beyond the processing of visual signals according to attribute, namely that of grouping signals according to colour (V4) or motion (V5); (b) that the functional separation evident in visual cortical areas devoted to motion and colour, respectively, is maintained at the level of parietal cortex, at least as far as grouping according to attribute is concerned; and (c) that, by contrast, this grouping-related functional segregation is not maintained at the level of the cerebellum.

Retinotopic mapping of the primary visual cortex - a challenge for MEG imaging of the human cortex

Magnetoencephalography (MEG) can be used to reconstruct neuronal activity with high spatial and temporal resolution. However, this reconstruction problem is ill-posed, and requires the use of prior constraints in order to produce a unique solution. At present there are a multitude of inversion algorithms, each employing different assumptions, but one major problem when comparing the accuracy of these different approaches is that often the true underlying electrical state of the brain is unknown. In this study, we explore one paradigm, retinotopic mapping in the primary visual cortex (V1), for which the ground truth is known to a reasonable degree of accuracy, enabling the comparison of MEG source reconstructions with the true electrical state of the brain. Specifically, we attempted to localize, using a beanforming method, the induced responses in the visual cortex generated by a high contrast, retinotopically varying stimulus. Although well described in primate studies, it has been an open question whether the induced gamma power in humans due to high contrast gratings derives from V1 rather than the prestriate cortex (V2). We show that the beanformer source estimate in the gamma and theta bands does vary in a manner consistent with the known retinotopy of V1. However, these peak locations, although retinotopically organized, did not accurately localize to the cortical surface. We considered possible causes for this discrepancy and suggest that improved MEG/magnetic resonance imaging co-registration and the use of more accurate source models that take into account the spatial extent and shape of the active cortex may, in future, improve the accuracy of the source reconstructions.

Motion processing, directional selectivity, and conscious visual perception in the human brain

Several human and monkey studies have demonstrated a close relationship between motion perception and activation of area V5, leading to the general view that activity in this area correlates with the subjective experience of motion. In the present study, we investigate whether the responses of this area are still governed by the motion percept when the latter is in conflict with the reality of the physical visual stimulation. We simultaneously presented two different, specially designed random-dot kinematograms, one to each eye. These stimuli either both had a single direction of motion and worked in synergy, or had opposite motion directions and thus cancelled each other out perceptually. In this way, we were able to pit the visual stimulus (one vs. two stimulating directions) against the reported perception (directional motion vs. motion noise) of human volunteers during fMRI experiments. We found that a strong motion stimulus that is weakly perceived is more effective in activating V5 (as well as V3) than a weaker motion stimulus, which is nevertheless robustly perceived. Thus, contrary to the prevailing view of perception being the correlate of activity in higher visual areas, we show here that activity is instead dominated by the properties of the physical stimulus, raising the question of whether there is a subpopulation of cells in V5 whose activity is critical for generating the motion percept. In addition, our results provide the first robust evidence for the presence of directionally selective neuronal populations in human prestriate cortex.

A Map for Horizontal Disparity in Monkey V2

The perception of visual depth is determined by integration of spatial disparities of inputs from the two eyes. Single cells in visual cortex of monkeys are known to respond to specific binocular disparities; however, little is known about their functional organization. We now show, using intrinsic signal optical imaging and single-unit physiology, that, in the thick stripe compartments of the second visual area (V2), there is a clustered organization of Near cells and Far cells, and moreover, there are topographic maps for Near to Far disparities within V2. Our findings suggest that maps for visual disparity are calculated in V2, and demonstrate parallels in functional organization between the thin, pale, and thick stripes of V2.

Adaptation to Natural Binocular Disparities in Primate V1 Explained by a Generalized Energy Model

Sensory processing in the brain is thought to have evolved to encode naturally occurring stimuli efficiently. We report an adaptation in binocular cortical neurons that reflects the tight constraints imposed by the geometry of 3D vision. We show that the widely used binocular energy model predicts that neurons dedicate part of their dynamic range to impossible combinations of left and right images. Approximately 42% of the neurons we record from V1 of awake monkeys behave in this way (a powerful confirmation of the model), while about 58% deviate from the model in a manner that concentrates more of their dynamic range on stimuli that obey the constraints of binocular geometry. We propose a simple extension of the energy model, using multiple subunits, that explains the adaptation we observe, as well as other properties of binocular neurons that have been hard to account for, such as the response to anti-correlated stereograms.

How do visual instructions influence the motor system?

The paper distinguishes the use of visual cues to guide reaching and grasping, and the ability to learn to associate arbitrary sensory cues with movements. Using positron emission tomography (PET), we have shown that the arbitrary association of visual cues and movements involves the ventral visual system (prestriate, inferotemporal and ventral prefrontal cortex), the basal ganglia and the dorsal premotor cortex. Using functional magnetic resonance imaging (fMRI), we have shown that the evoked haemodynamic responses in the ventral visual system are time-locked to the presentation of the visual cues, that the response in the motor cortex is locked to the time of response, and that the response in the dorsal premotor cortex shows cuerelated, movement-related and set-related components. Using PET we have shown that there are learning-related changes in activation in both the ventral prestriate cortex and the basal ganglia (globus pallidus) when subjects learn a visuomotor associative task. We argue that the basal ganglia may act as a flexible system for learning the association of sensory cues and movements.

Cortical activity in Parkinson's disease during executive processing depends on striatal involvement

Patients with idiopathic Parkinson's disease exhibit impairments in executive processes, including planning and set-shifting, even at the early stages of the disease. We have recently developed a new card-sorting task to study the specific role of the caudate nucleus in such executive processes and have shown, using functional magnetic resonance imaging (fMRI) in young healthy adults, that the caudate nucleus is specifically required when a set-shift must be planned. Here the same fMRI protocol was used to compare the patterns of activation in a group of early-stage Parkinson's disease patients (seven right-handed patients at Hoehn and Yahr stages 1 and 2; mean age 62 years, range 56-70) and matched control subjects. Increased cortical activation was observed in the patients compared with the control group in the condition not specifically requiring the caudate nucleus. On the other hand, decreased cortical activation was observed in the patient group in the condition significantly involving the caudate nucleus. This event-related fMRI study showed a pattern of cortical activation in Parkinson's disease characterized by either reduced or increased activation depending on whether the caudate nucleus was involved or not in the task. This activation pattern included not only the prefrontal regions but also posterior cortical areas in the parietal and prestriate cortex. These findings are not in agreement with the traditional model, which proposes that the nigrostriatal dopamine depletion results in decreased cortical activity. These observations provide further evidence in favour of the hypothesis that not only the nigrostriatal and but also the mesocortical dopaminergic substrate may play a significant role in the cognitive deficits observed in Parkinson's disease.

Visual functions without the occipital lobe or after cerebral hemispherectomy in infancy

This paper investigates whether and to what extent vision with awareness is still possible in the whole visual field after loss of the occipital lobe of one or both cerebral hemispheres or after hemispherectomy in childhood. The visual functions of four children who suffered from unilateral or bilateral loss of the occipital lobe or who had been hemispherectomized were examined. The results show that even after unilateral loss of the striate and prestriate cortex the extent of the visual field may still be in the normal range. The residual visual functions may be mediated by intact extrastriate areas such as V5 and LO of the damaged cerebral hemisphere. It is also shown that even after complete hemispherectomy in early life the visual field may have a normal extent and that conscious visual perception in the whole visual field may be preserved. In hemispherectomized children, the remaining cerebral hemisphere or neural structures in the midbrain, including the superior colliculi and the praetectum, may be able to mediate these visual functions.

Putting the Cheshire Cat Together in Amblyopia. Focus on “Spatial Resolution for Feature Binding Is Impaired in Peripheral and Amblyopic Vision”

A perceptual “object” can be defined as a spatiotemporal coincidence of its different attributes: for example, an object could be a large, red, moving square. If the different attributes occur at different times, or in different places, they do not define an object. The world would be a strange

Functional MRI of visuospatial working memory in schizotypal personality disorder: a region-of-interest analysis

Functional MRI studies have begun to identify neural networks implicated in visuo-spatial working memory in healthy volunteers and patients with schizophrenia. The study of schizotypal personality disorder (SPD) provides regional analysis in unmedicated patients in the schizophrenia spectrum. Unmedicated patients with SPD by DSM-IV criteria and normal controls were assessed with fMRI while performing a visuo-spatial working-memory task. It required the subjects to retain the location of three dots located on the circumference of an imaginary circle and then respond to a query display in which one dot was presented and the subject required to press a button to indicate whether the probe dot location was previously displayed. Subject groups did not differ significantly in spatial memory scores. The exact Talairach and Tournoux coordinates of brain areas previously reported to show activation with spatial memory tasks were assessed. The majority of these locations showed BOLD response activation significantly less in patients during the memory retention period, including the left ventral prefrontal cortex, superior frontal gyrus, intraparietal cortex and posterior inferior gyrus. Regions in the right middle prefrontal and prestriate cortex showed greater activation at a trend level for patients with SPD than for normal controls. In addition, we replicated the findings of increased activation with the task in healthy volunteers in the premotor areas, ventral prefrontal cortex and parietal cortex. SPD patients show decreased activation compared to healthy volunteers in key frontal regions and we also provided a partial replication of findings reported in healthy subjects.

Functional Brain Mapping of the Macaque Related to Spatial Working Memory as Revealed by PET

To define the cortical areas that subserve spatial working memory in a nonhuman primate, we measured regional cerebral blood flow (rCBF) with [(15)O]H(2)O and positron emission tomography while monkeys performed a visually guided saccade (VGS) task and an oculomotor delayed-response (ODR) task. Both Statistical Parametric Mapping and regions of interest-based analyses revealed an increase of rCBF in the area surrounding the principal sulcus (PS), the superior convexity, the anterior bank of the arcuate sulcus (AS), the lateral orbitofrontal cortex (lOFC), the frontal pole (FP), the anterior cingulate cortex (ACC), the lateral bank of the intraparietal sulcus (lIPS) and the prestriate cortex. In the prefrontal cortex (PS, superior convexity, AS, lOFC and FP), rCBF values correlated positively with ODR task performance scores. From the hippocampus, rCBF values correlated negatively with ODR task performance. From the AS, superior convexity, lOFC, FP, ACC and lIPS, rCBF values of the PS correlated positively with rCBF values and negatively with hippocampus rCBF values. These results suggest that neural circuitry in the prefrontal cortex directly contributes the spatial working memory processes and that, in spatial working memory processes, the posterior parietal cortex and hippocampus have a different role to the prefrontal cortex.

Perceptual grouping correlates with short synchronization in monkey prestriate cortex.

Synchronization in the visual cortex at 35-80 Hz is assumed to support perceptual grouping. We tested this hypothesis in a figure-ground task in which a trained monkey indicated by a key whether he perceived a figure that was composed of the same blobs as the background distractors. The task was sufficiently difficult such that about 25% of responses were incorrect. We recorded population activity with 7 microelectrodes in prestriate cortex (V2). During a short period before the monkey's perceptual response, locations of figure-activated neurons showed increased synchronization (50-80 Hz) in correct compared to incorrect responses, while other signal measures were unrelated to perception. These are first indications that a short synchronous burst in V2 may support perceptual grouping.

Connections of the medial posterior parietal cortex (area 7m) in the monkey

The afferent and efferent cortical and subcortical connections of the medial posterior parietal cortex (area 7m) were studied in cebus (Cebus apella) and macaque (Macaca fascicularis) monkeys using the retrograde and anterograde capabilities of the horseradish peroxidase (HRP) technique. The principal intraparietal corticocortical connections of area 7m in both cebus and macaque cases were with the ipsilateral medial bank of the intraparietal sulcus (MIP) and adjacent superior parietal lobule (area 5), inferior parietal lobule (area 7a), lateral bank of the IPS (area 7ip), caudal parietal operculum (PGop), dorsal bank of the caudal superior temporal sulcus (visual area MST), and medial prestriate cortex (including visual area PO and caudal medial lobule). Its principal frontal corticocortical connections were with the prefrontal cortex in the shoulder above the principal sulcus and the cortex in the shoulder above the superior ramus of the arcuate sulcus (SAS), the area purported to contain the smooth eye movement-related frontal eye field (FEFsem) in the cebus monkey by other investigators. There were moderate connections with the cortex in the rostral bank of the arcuate sulcus (purported to contain the saccade-related frontal eye field; FEFsac), supplementary eye field (SEF), and rostral dorsal premotor area (PMDr). Area 7m also had major connections with the cingulate cortex (area 23), particularly the ventral bank of the cingulate sulcus. The principal subcortical connections of area 7m were with the dorsal portion of the ventrolateral thalamic (VLc) nucleus, lateral posterior thalamic nucleus, lateral pulvinar, caudal mediodorsal thalamic nucleus and medial pulvinar, central lateral, central superior lateral, and central inferior intralaminar thalamic nuclei, dorsolateral caudate nucleus and putamen, middle region of the claustrum, nucleus of the diagonal band, zona incerta, pregeniculate nucleus, anterior and posterior pretectal nuclei, intermediate layer of the superior colliculus, nucleus of Darkschewitsch and dorsomedial parvicellular red nucleus (macaque cases only), dorsal, dorsolateral and lateral basilar pontine nuclei, nucleus reticularis tegmenti pontis, locus ceruleus, and superior central nucleus. The findings are discussed in terms of the possibility that area 7m contains a "medial parietal eye field" and belongs to a neural network of oculomotor-related structures that plays a role in the control of eye movement.

The neurology of saccades and covert shifts in spatial attention: an event-related fMRI study.

Visual neglect occurs most frequently and persistently after lesions that include the right supramarginal gyrus (SMG), a part of the inferior parietal lobule. Patients with this syndrome make very few saccades to the left, and show abnormal performance on tasks in which they must covertly shift their attention to the left, suggesting that the right SMG is involved in the generation of saccades and attention shifts. Functional imaging studies of saccades and covert attention shifts in the normal brain, however, have shown weak or absent responses in both SMGs. We used event-related functional MRI to re-examine the responses to saccades and attention shifts within a single experiment, and to assess responses to left- and right-sided stimuli independently. When subjects made saccades to peripheral stimuli, the expected responses were seen in striate and prestriate cortex, the superior parietal lobules, the frontal eye fields, the supplementary motor area and the anterior insulae. In addition there was a response in the right SMG but not in the left SMG, as predicted from the clinical literature. When subjects made a covert visual assessment of the peripheral stimulus without any saccade, greater activity was seen in all of the areas in the frontoparietal network. Each area showed a bias towards contralateral stimuli, with two exceptions: the anterior insulae gave mainly ipsilateral responses, whilst the right SMG gave equal responses to right- and left-sided stimuli. These findings are discussed in the context of current theories pertaining to the clinical syndrome of neglect.

AGE-RELATED CHANGES IN THE NEURAL CORRELATES OF DEGRADED AND NONDEGRADED FACE PROCESSING

In order to explore the neural correlates of age-related changes in visual perception of faces, positron emission tomographic scans were obtained on young and old adults while they were engaged in tasks of nondegraded and degraded face matching. Old adults were less accurate than were young adults across all face matching conditions, although the age difference was greatly reduced when degraded performance was adjusted for nondegraded performance. The interaction of age and degree of degradation on performance measures was not significant. Brain activity patterns during nondegraded face matching were similar in the two groups with some differences in parietal and prestriate cortices (greater activity in young adults) and in prefrontal cortex, thalamus, and hippocampus (greater activity in old adults). Increases in activity related to increasing degradation of the faces were seen mainly in prefrontal cortices in both age groups. Despite this similarity in the brain response to face degradation, there were striking differences between groups in the correlations between brain activity and degraded task performance. Different regions of extrastriate cortex were positively correlated with behavioural measures in the two groups (fusiform gyrus in the young adults and posterior occipital regions in old adults). In addition two areas where older adults showed greater activity during nondegraded face matching, thalamus and hippocampus, also showed positive correlations with behaviour during the degraded tasks in this group, but not in the young group. Thus, although the elderly are not more vulnerable to the effects of increasing face degradation, the brain systems involved in carrying out these visual discriminations in young and old adults are not the same. These results are consistent with the idea of functional plasticity in face processing over the life span.

Impaired reading in patients with right hemianopia

A left occipital stroke may result in alexia for two reasons, which may coexist depending on the distribution of the lesion. A lesion of the left lateroventral prestriate cortex or its afferents impairs word recognition (“pure” alexia). If the left primary visual cortex or its afferents are destroyed, resulting in a complete right homonymous hemianopia, rightward saccades during text reading are disrupted (“hemianopic” alexia). By using functional imaging, we showed two separate but interdependent systems involved in reading. The first, subserving word recognition, involved the representation of foveal vision in the left and right primary visual cortex and the ventral prestriate cortex. The second system, responsible for the planning and execution of reading saccades, consisted of the representation of right parafoveal vision in the left visual cortex, the bilateral posterior parietal cortex (left > right), and the frontal eye fields (right > left). Disruption of this distributed neural system was demonstrated in patients with severe right homonymous hemianopia, commensurate with their inability to perform normal reading eye movements. Text reading, before processes involved in comprehension, requires the integration of perceptual and motor processes. We have demonstrated these distributed neural systems in normal readers and have shown how a right homonymous hemianopia disrupts the motor preparation of reading saccades during text reading. Ann Neurol 2000;47:171–178.