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Activation of human primary visual cortex during visual recall: a magnetic resonance imaging study.

The degree to which the process involved in visual perception and visual imagery share a common neuroanatomical substrate is unclear. Physiological evidence for localization of visual imagery early in the visual pathways would have important bearing on current theories of visual processing. A magnetic resonance imaging technique sensitive to regional changes in blood oxygenation was used to obtain functional activation maps in the human visual cortex. During recall of a visual stimulus, focal increases in signal related to changes in blood flow were detected in V1 and V2 cortex in five of seven subjects. These experiments show that the same areas of the early visual cortex that are excited by visual stimulation are also activated during mental representation of the same stimulus. Some of the processes used in topographically mapped cortical areas during visual perception may also be utilized during visual recall.

Adult↗

Task-dependent influences of attention on the activation of human primary visual cortex.

There has been a good deal of controversy over whether attention influences area V1-the first cortical area onto which information from the retina is projected. Attention to motion has been found to modulate monkey area MT and the human homolog of MT/MST. Here we show that activation of V1 by attention to motion is task dependent. Our stimulus consisted of a group of translating random dots superimposed over another group of random dots executing expansion motion. Subjects were instructed to pay attention selectively to the translation, expansion, or neither in particular (passive condition). The activity in the human MT/MST homolog measured by functional magnetic resonance imaging (fMRI) was significantly higher in both the translation and the expansion conditions than in the passive condition, while the activity in area V1 was significantly higher only in the translation condition. These results show that attention to motion modulates area V1, and more interestingly that high-level cognitive processing such as attention may directly or indirectly determine the retroactive extent of feedback within the motion pathway in a manner dependent on the type of motion attended.

Cognition↗

Functional MRI reveals spatially specific attentional modulation in human primary visual cortex.

Selective visual attention can strongly influence perceptual processing, even for apparently low-level visual stimuli. Although it is largely accepted that attention modulates neural activity in extrastriate visual cortex, the extent to which attention operates in the first cortical stage, striate visual cortex (area V1), remains controversial. Here, functional MRI was used at high field strength (3 T) to study humans during attentionally demanding visual discriminations. Similar, robust attentional modulations were observed in both striate and extrastriate cortical areas. Functional mapping of cortical retinotopy demonstrates that attentional modulations were spatially specific, enhancing responses to attended stimuli and suppressing responses when attention was directed elsewhere. The spatial pattern of modulation reveals a complex attentional window that is consistent with object-based attention but is inconsistent with a simple attentional spotlight. These data suggest that neural processing in V1 is not governed simply by sensory stimulation, but, like extrastriate regions, V1 can be strongly and specifically influenced by attention.

Attention↗

Layer IVA of rhesus monkey primary visual cortex.

Layer IVA of rhesus monkey striate cortex contains pyramidal cells arranged in distinct groups. Their cell bodies are in a configuration of flat cones, each with an average diameter of 60 microns, and their apical dendrites aggregate into bundles that ascend toward the pial surface. Nissl-stained sections suggest that these pyramidal cell cones have their bases in layer IVB, with their tops extending into layer IVA. The neurons in the cones are readily apparent in MAP2 antibody-stained material, and in cytochrome oxidase-reacted tissue it is evident that the pyramidal cell cones occupy the pale spaces that are surrounded by the darkly reactive honeycomb lattice. This lattice of neuropil around the cones contains some axons and boutons that are immunoreactive for parvalbumin, and it is within the lattice that other investigators have shown afferents from the parvocellular (P)-layers of the dLGN to terminate. Because of this input, it is likely that the pyramidal cell cones of layer IVA are involved with color and form perception. The relationship between the layer IVA cones of neurons and the underlying system of previously described pyramidal cell modules (Peters and Sethares, 1991) is discussed, as well as the possibility that the pyramidal cell cones might represent aggregations of neurons, which receive input from basic sets of P-like afferents originating from color-responsive ganglion cells of the retina, as described by Schein and de Monasterio (1987).

Animals↗

Moving illusory contours activate primary visual cortex: an fMRI study.

Identifying the cortical areas activated by illusory contours provides valuable information on the mechanisms of object perception. We applied functional magnetic resonance imaging to identify the visual areas of the human brain involved in the perception of a moving Kanizsa-type illusory contour. Our results indicate that, in addition to other cortical regions, areas V5 and V1 are activated. Activity in area V1 was particularly prominent.

Adult↗

Numerical relationships between geniculocortical afferents and pyramidal cell modules in cat primary visual cortex.

An analysis has been made of the quantitative data available on the number of pyramidal cell modules of layer IV neurons, and of geniculocortical axons and their synapses in cat striate cortex. It is found that the convergence of geniculocortical afferents upon any one pyramidal cell module is enormous, since in any one location there is overlap between 360-540 X-axons and 300-540 Y-axons. In total, the X- and Y-axonal arbors provide some 1640 x 10(6) synapses to area 17, which is equivalent to a ratio of 160-200 synapses per layer IV neuron. These values assume that geniculocortical terminals synapse only with the spiny stellate cells of layer IV. The values are reduced to 100-125 per spiny stellate cell when account is taken of the synapses that involve the dendrites that enter layer IV from neurons with cell bodies in other layers. Since each layer IV neuron receives some 2500 asymmetric synapses, this means that only 5% of the total excitatory input to a layer IV neuron seems to be provided by the geniculocortical afferents. Further, if the boutons in the geniculocortical axonal arbors are distributed homogeneously across layer IV, each axon could only provide one synapse to about one in four of the layer IV neurons encompassed by its plexus. It may be, however, that instead of being spread evenly, boutons in individual arbors converge upon individual neurons to supply a number of synapses to them. But even so, it seems unlikely that any individual geniculate axon could dominate the activity of a particular cortical neuron.

Animals↗

Regulation of calcium-binding protein immunoreactivity in GABA neurons of macaque primary visual cortex.

Monocular deprivation produces an imbalance in visual drive from the two eyes, which in adult macaque V1 leads to marked changes in the neurochemistry of GABA interneurons. Such changes were further examined by studying immunoreactivity for calbindin, calretinin, and parvalbumin, three calcium-binding proteins that mark distinct subpopulations of GABA neurons, in macaques that had been monocularly deprived by intravitreal injection of tetrodotoxin. Deprivation for 5 d or longer produced a reversal in the normal pattern of calbindin immunostaining in layer III, from one in which intense neuronal immunostaining surrounded the cytochrome oxidase-rich puffs to one in which it occupied the puffs. Over the same period, calbindin immunostaining in other layers was reduced across the entire width of deprived-eye columns or extended into flanking regions of normal-eye columns. In contrast, reduction in parvalbumin immunostaining occurred only in deprived-eye columns and included only terminals with short periods of deprivation (up to 17 d) but both terminals and somata with longer periods. No change in calretinin immunoreactivity was observed. These findings demonstrate that GABA neurons of macaque V1 vary in their response to monocular deprivation according to their neurochemistry and position, suggesting that the weight of inputs from the two eyes and the intrinsic characteristics of each GABA population determine how a neuron responds to a change in visual input.

Animals↗

Corticocortical connections between area 21a and primary visual cortex in the cat.

We investigated the corticocortical connections between area 21a and ipsilateral areas 17 and 18 in the cat. The anterograde/retrograde fluorescent tracer tetramethyl-rhodamine conjugated to dextran (Fluoro-Ruby) was injected into area 21a of the anaesthetized cat. Cell bodies labelled retrogradely from area 21a were consistently observed in both areas 17 and 18, primarily located in the supragranular layers of cortex where they formed discrete patches of cells. Similar numbers of cell bodies were labelled retrogradely in areas 17 and 18 of each animal. Our data are also consistent with previous reports of a reciprocal projection from area 21a back to areas 17 and 18 terminating principally in infragranular cortical layers.

Animals↗

Gating of neuronal responses in macaque primary visual cortex by an attentional spotlight.

Neuronal responses were recorded from the striate cortex of monkeys trained to perform visual discrimination at locations in the visual field to which their attention was drawn. A subset of neurons showed vigorous responses to visual stimuli for trials in which the monkey was directing its attention to the respective receptive field location. In trials where attention is directed elsewhere, responses to the same stimuli were significantly reduced. In some cells the early response component was not modulated by attention, but later components were affected by the locus of attention. The results suggest the operation of a feedback in the paradigm that spotlights a topographically restricted area of V1 for further processing at higher levels.

Animals↗

Geometrical and topological relationships between multiple functional maps in cat primary visual cortex.

The mammalian striate cortex is organized such that the receptive field properties of neighboring neurons change gradually across the cortical surface, forming so-called cortical maps. The presence of such maps has been demonstrated in different species of mammals for several parameters characterizing the visual space: retinotopy, ocular dominance, orientation, direction of motion and spatial frequency. In this study we used the optical imaging of intrinsic signals to simultaneously record the multiple functional maps in the same animal in order to obtain a comprehensive set of rules that govern mutual dependencies among the functional maps. Our results indicate that while orientation, direction and ocular dominance are represented on the cortex in a mutually dependent manner, the representation of spatial frequency is independent of the other types of cortical representations. The presence and/or absence of mutual dependence among the multiple functional maps are suggested to provide an important clue for the understanding of the development of visual cortical information representation in neonatal animals.

Animals↗

Direct mapping of ocular dominance columns in human primary visual cortex.

Functional magnetic resonance imaging at 2.0T was employed to identify columnar structures in human visual cortex. Sagittal sections (4 mm thickness) covering the calcarine cortex were acquired with use of a multiecho low flip angle gradient-echo sequence at 4.0 s temporal resolution and 0.25 x 0.25 mm2 spatial resolution. Extending earlier attempts based on a differential paradigm contrasting left vs right eye stimulation, this work presents the first direct mapping of human ocular dominance columns by measuring separate activation maps with left and right eye stimulation. The resulting individual maps reveal patterns of ocular dominance as spots or bands of altered activity in calcarine cortex. Their superposition shows only little spatial overlap of eye-specific encoding which strongly supports the genuineness of these functional units.

Adult↗

Visual field defects in Alzheimer's disease patients may reflect differential pathology in the primary visual cortex.

The aim of this study was to test the hypothesis that differences in density of senile plaques (SP) and neurofibrillary tangles (NFT) in the cuneal and lingual gyri of area V1 of the visual cortex could explain the predominantly inferior visual field defects seen in patients with Alzheimer's disease (AD). The density of SP and NFT was measured in the cuneal and lingual gyri of 18 AD patients. In 7/18 (39%) patients, the density of SP and/or NFT was significantly greater in the cuneal compared with the lingual gyri. In 3/18 (17%) patients, densities were greater in the lingual than the cuneal gyri and in 8/18 (44%) patients there were no significant differences among gyri. The data suggest that pathological differences between cuneal and lingual gyri could contribute to the reported visual field defects in some AD patients.

Aged↗

Independent component filters of natural images compared with simple cells in primary visual cortex.

Properties of the receptive fields of simple cells in macaque cortex were compared with properties of independent component filters generated by independent component analysis (ICA) on a large set of natural images. Histograms of spatial frequency bandwidth, orientation tuning bandwidth, aspect ratio and length of the receptive fields match well. This indicates that simple cells are well tuned to the expected statistics of natural stimuli. There is no match, however, in calculated and measured distributions for the peak of the spatial frequency response: the filters produced by ICA do not vary their spatial scale as much as simple cells do, but are fixed to scales close to the finest ones allowed by the sampling lattice. Possible ways to resolve this discrepancy are discussed.

Animals↗

Development of cortical afferents and cortico-tectal efferents of the mammalian (rat) primary visual cortex.

At the time when the fibres from the striate cortex (area 17) begin to innervate the superficial layers of the superior colliculus of the young rat (postnatal days 4 and 5) a high degree of specificity in the organization of this newly formed cortico-tectal projection is already apparent. Thus, in young rats, as in adult mammals of virtually all species studied so far, the somata of cortico-tectal neurones are confined to lamina V of the ipsilateral cortex. However, this high degree of laminar (radial) specificity in young animals is accompanied by a substantial degree of exuberance as indicated by a tangential distribution of the cortico-tectal cells which is wider than that in the adult. The exuberant projections are pruned during the second postnatal week. The cortico-cortical associational and commissural fibres start to enter the grey matter of the rat striate cortex after postnatal day 7. Again a high degree of specificity in the laminar distribution of those newly established projections is apparent. However, the cortico-cortical projection, at the time when cortico-cortical fibres enter the cortical laminae, is clearly exuberant since the tangential spread of cortical cells projecting to the striate cortex is wider than that in the adult. Pruning of these excessive projections takes place some time after postnatal day 14. It is believed that understanding the mechanism(s) underlying the development of connections of the rat visual cortex might be of general importance in understanding developmental abnormalities in the pattern of interconnections of the visual cortices of other mammalian orders.

Afferent Pathways↗

Spatial resolution and contrast sensitivity of single neurons in area 19 of split-chiasm cats: a comparison with primary visual cortex.

Electrophysiological recordings were carried out in the callosal recipient zone of area 19 in normal and split-chiasm cats and, for comparison purposes, at the border of areas 17 and 18 of split-chiasm cats. The influences of retinothalamic and callosal inputs on a single cortical neurons were thereby evaluated. Extracellular recordings of single cells were made in anaesthetized and paralysed cats in the zone representing the central visual field. Receptive field properties were assessed using sine wave gratings drifting in optimal directions. Results showed that in area 19 and areas 17/18 one-third of the cells were binocularly driven after section of the optic chiasm. In area 19, the spatial resolution and contrast sensitivity of cells driven via the dominant eye were similar in the normal and split-chiasm groups. In areas 17/18 and area 19 of split-chiasm cats, binocular cells showed significant interocular matching of their receptive field properties (spatial resolution and contrast threshold), although small differences were observed. These small interocular differences were related to the cell's ocular dominance rather than to the signal transmission route (thalamic or callosal).

Animals↗

The perceptual grouping criterion of colinearity is reflected by anisotropies of connections in the primary visual cortex.

An important step in the processing of visual patterns is the segmentation of the retinal image. Neuronal responses evoked by the contours of individual objects need to be identified and associated for further joint processing. These grouping operations are based on a number of Gestalt criteria. Here we report that connections in the visual cortex of the cat exhibit a highly significant anisotropy, preferentially linking neurons activated by contours that have similar orientation and are aligned colinearly. These anatomical data suggest a close relation between the perceptual grouping criterion of colinearity and the topology of tangential intracortical connections. We propose that tangential intracortical connections support perceptual grouping by modulating the saliency of distributed cortical responses in a context-dependent way. The present data are compatible with the hypothesis that the criteria for this grouping operation are determined by the architecture of the tangential connections.

Animals↗

Actions of compounds manipulating the nitric oxide system in the cat primary visual cortex.

1. We iontophoretically applied NG-nitro-L-arginine (L-NOArg), an inhibitor of nitric oxide synthase (NOS), to cells (n = 77) in area 17 of anaesthetized and paralysed cats while recording single-unit activity extracellularly. In twenty-nine out of seventy-seven cells (38%), compounds altering NO levels affected visual responses. 2. In twenty-five out of twenty-nine cells, L-NOArg non-selectively reduced visually elicited responses and spontaneous activity. These effects were reversed by co-application of L-arginine (L-Arg), which was without effect when applied alone. Application of the NO donor diethylamine-nitric oxide (DEA-NO) produced excitation in three out of eleven cells, all three cells showing suppression by L-NOArg. In ten cells the effect of the soluble analogue of cGMP, 8-bromo-cGMP, was tested. In three of those in which L-NOArg application reduced firing, 8-bromo-cGMP had an excitatory effect. In six out of fifteen cells tested, L-NOArg non-selectively reduced responses to NMDA and alpha-amino-3-hydroxy-5-methylisoxasole-4-propionic acid (AMPA). Again, co-application of L-Arg reversed this effect, without enhancing activity beyond control values. 3. In a further subpopulation of ten cells, L-NOArg decreased responses to ACh in five. 4. In four out of twenty-nine cells L-NOArg produced the opposite effect and increased visual responses. This was reversed by co-application of L-Arg. Some cells were also affected by 8-bromo-cGMP and DEA-NO in ways opposite to those described above. It is possible that the variety of effects seen here could also reflect trans-synaptic activation, or changes in local circuit activity. However, the most parsimonious explanation for our data is that NO differentially affects the activity of two populations of cortical cells, in the main causing a non-specific excitation.

Acetylcholine↗