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A hexagonal orthogonal-oriented pyramid as a model of image representation in visual cortex.

Retinal ganglion cells represent the visual image with a spatial code, in which each cell conveys information about a small region in the image. In contrast, cells of primary visual cortex employ a hybrid space-frequency code in which each cell conveys information about a region that is local in space, spatial frequency, and orientation. Despite the presumable importance of this transformation, we lack any comprehensive notion of how it occurs. Here we describe a mathematical model for this transformation. The hexagonal orthogonal-oriented quadrature pyramid (HOP) transform, which operates on a hexagonal input lattice, employs basis functions that are orthogonal, self-similar, and localized in space, spatial frequency, orientation, and phase. The basis functions, which are generated from seven basic types through a recursive process, form an image code of the pyramid type. The seven basis functions, six bandpass and one low-pass, occupy a point and a hexagon of six nearest neighbors on a hexagonal sample lattice. The six bandpass basis functions consist of three with even symmetry, and three with odd symmetry. The three even kernels are rotations of 0, 60, and 120 degrees of a common kernel; likewise for the three odd kernels. At the lowest level, the inputs are image samples. At each higher level, the input lattice is provided by the low-pass coefficients computed at the previous level. At each level, the output is subsampled in such a way as to yield a new hexagonal lattice with a spacing square root 7 larger than the previous level, so that the number of coefficients is reduced by a factor of seven at each level. In the biological model, the input lattice is the retinal ganglion cell array. The resulting scheme provides a compact, efficient code of the image and generates receptive fields that resemble those of the primary visual cortex.

Computer Simulation↗

Transient activity in the human calcarine cortex during visual-mental imagery: an event-related fMRI study.

Although it is largely accepted that visual-mental imagery and perception draw on many of the same neural structures, the existence and nature of neural processing in the primary visual cortex (or area V1) during visual imagery remains controversial. We tested two general hypotheses: The first was that V1 is activated only when images with many details are formed and used, and the second was that V1 is activated whenever images are formed, even if they are not necessarily used to perform a task. We used event-related functional magnetic resonance imaging (ER-fMRI) to detect and characterize the activity in the calcarine sulcus (which contains the primary visual cortex) during single instances of mental imagery. The results revealed reproducible transient activity in this area whenever participants generated or evaluated a mental image. This transient activity was strongly enhanced when participants evaluated characteristics of objects, whether or not details actually needed to be extracted from the image to perform the task. These results show that visual imagery processing commonly involves the earliest stages of the visual system.

Acoustic Stimulation↗

Cortico-cortical and cortico-amygdaloid projections of the rat occipital cortex: a Phaseolus vulgaris leucoagglutinin study.

The efferent projections of the occipital cortex of the rat were investigated using the Phaseolus vulgaris leucoagglutinin anterograde tract tracing technique. Particular attention was focused on projections to the amygdala and amygdalopetal cortical areas. The primary visual cortex had projections to the medial and lateral portions of occipital area 2 and other cortical regions, but no projections to the amygdala or amygdalopetal cortical areas. The only occipital area that had direct projections to the amygdala was the most ventral portion of lateral occipital area 2, located just dorsal to temporal area 2. This occipitotemporal junction region, which received projections from secondary visual cortical areas but not from the primary visual cortex, had projections to the lateral nucleus, magnocellular basal nucleus, and lateral capsular subdivision of the central nucleus of the amygdala. Occipital area 2 had projections to seven amygdalopetal cortical regions: temporal area 2, temporal area 3, frontal area 2, ventrolateral orbitofrontal area, occipitotemporal junction region, lateral entorhinal area, and the perirhinal cortex. Projections to the perirhinal cortex targeted regions located adjacent to the parietal cortex and caudal temporal cortex, but not regions adjacent to the rostral temporal cortex. Other cortical regions receiving projections from medial and lateral portions of occipital area 2 included the presubiculum, retrosplenial areas, and caudal portions of the parietal cortical areas 1 and 2. The results of the present investigation, in conjunction with previous anatomical and neurobehavioral studies, support the concept that rodent cortical visual pathways, like those of primates, consist of a dorsal system involved with visuospatial functions and a ventral system involved with object recognition. As in primates, the ventral pathway projects to the temporal-perirhinal region in a cascading manner; only highly processed information from tertiary visual cortical areas reaches the amygdala. Unlike primates, however, cortical areas in the rat brain that receive highly processed visual information appear to be regions of multisensory convergence.

Amygdala↗

Evolution of specialized pyramidal neurons in primate visual and motor cortex.

The neocortex of primates contains several distinct neuron subtypes. Among these, Betz cells of primary motor cortex and Meynert cells of primary visual cortex are of particular interest for their potential role in specialized sensorimotor adaptations of primates. Betz cells are involved in setting muscle tone prior to fine motor output and Meynert cells participate in the processing of visual motion. We measured the soma volumes of Betz cells, Meynert cells, and adjacent infragranular pyramidal neurons in 23 species of primate and two species of non-primate mammal (Tupaia glis and Pteropus poliocephalus) using unbiased stereological techniques to examine their allometric scaling relationships and socioecological correlations. Results show that Betz somata become proportionally larger with increases in body weight, brain weight, and encephalization whereas Meynert somata remain a constant proportion larger than other visual pyramidal cells. Phylogenetic variance in the volumetric scaling of these neuronal subtypes might be related to species-specific adaptations. Enlargement of Meynert cells in terrestrial anthropoids living in open habitats, for example, might serve as an anatomical substrate for predator detection. Modification of the connectional and physiological properties of these neurons could constitute an important evolutionary mode for species-specific adaptation.

Aged↗

Delayed maturation of receptive field center/surround mechanisms in V2.

Neurons in the adult visual cortex are capable of integrating signals over a large area that surrounds their classic receptive field (RF), and this ability of cortical neurons is thought to be intimately involved in perceptual binding. It is not known, however, at what age these long-range signal interactions emerge. Here, we report that qualitatively adult-like center/surround interactions are already present in the primary visual cortex as early as postnatal day 14 in macaque monkeys. However, the RF surrounds of visual area 2 (V2) neurons were largely absent until 4 weeks of age and, as late as 8 weeks of age, center/surround signal interactions in V2 neurons were immature. Our results suggest that the cortical circuits underlying the RF center/surround of individual neurons mature considerably later in V2 than in the primary visual cortex and give critical evidence for the hypothesis that the functional maturation of the primate visual brain proceeds in a hierarchical manner.

Age Factors↗

Spiral K-space MR imaging of cortical activation.

Brain function can be mapped with magnetic resonance (MR) imaging sensitized to regional changes in blood oxygenation due to cortical activation. Several MR imaging methods, including conventional imaging and echo-planar imaging, have been successfully used for this purpose. The authors investigated spiral k-space MR imaging, implemented with an unmodified 1.5-T clinical imager, for imaging of cortical activation. A gradient-echo, spiral k-space imaging method was used to measure activation in the primary visual cortex (number sequence task), primary motor cortex (fist-clenching task), and prefrontal cortex (verbal fluency task). Comparison of conventional and spiral k-space imaging in the visual and motor cortex, in which signal-to-noise ratio, voxel size, and imaging time were matched, showed that artifacts were reduced with the spiral k-space method, while the area and degree of activation were similar. The number of sections that could be imaged in a fixed time interval was increased by a factor of four with this implementation of spiral k-space imaging compared with conventional imaging.

Brain Mapping↗

fMRI reveals a common neural substrate of illusory and real contours in V1 after perceptual learning.

Perceptual learning involves the specific and relatively permanent modification of perception following a sensory experience. In psychophysical experiments, the specificity of the learning effects to the trained stimulus attributes (e.g., visual field position or stimulus orientation) is often attributed to assumed neural modifications at an early cortical site within the visual processing hierarchy. We directly investigated a neural correlate of perceptual learning in the primary visual cortex using fMRI. Twenty volunteers practiced a curvature discrimination on Kanizsa-type illusory contours in the MR scanner. Practice-induced changes in the BOLD response to illusory contours were compared between the pretraining and the posttraining block in those areas of the primary visual cortex (V1) that, in the same session, had been identified to represent real contours at corresponding visual field locations. A retinotopically specific BOLD signal increase to illusory contours was observed as a consequence of the training, possibly signaling the formation of a contour representation, which is necessary for performing the curvature discrimination. The effects of perceptual training were maintained over a period of about 10 months, and they were specific to the trained visual field position. The behavioral specificity of the learning effects supports an involvement of V1 in perceptual learning, and not in unspecific attentional effects.

Adult↗

Spatial invariance of visual receptive fields in parietal cortex neurons.

Spatial information is conveyed to the primary visual cortex in retinal coordinates. Movement trajectory programming, however, requires a transformation from this sensory frame of reference into a frame appropriate for the selected part of the body, such as the eye, head or arms. To achieve this transformation, visual information must be combined with information from other sources: for instance, the location of an object of interest can be defined with respect to the observer's head if the position of the eyes in the orbit is known and is added to the object's retinal coordinates. Here we show that in a subdivision of the monkey parietal lobe, the ventral intraparietal area (VIP), the activity of visual neurons is modulated by eye-position signals, as in many other areas of the cortical visual system. We find that individual receptive fields of a population of VIP neurons are organized along a continuum, from eye to head coordinates. In the latter case, neurons encode the azimuth and/or elevation of a visual stimulus, independently of the direction in which the eyes are looking, thus representing spatial locations explicitly in at least a head-centred frame of reference.

Animals↗

Binocular interactions and disparity coding in area 21a of cat extrastriate visual cortex.

We have examined, using both qualitative and quantitative techniques, binocular interactions of extracellularly recorded single neurons in the extrastriate cortical area 21a of anaesthetized and paralysed cats. Consistent with previous reports we have found that: (a) all area 21a neurons were orientation-selective, with about 65% of them preferring orientations within 30 degrees of the vertical; and (b) over 75% of area 21a cells could be activated through either eye. Furthermore, a significant minority (4 cells; about 10%) of a subpopulation of 39 neurons in which binocular interactions were examined quantitatively, were "obligatory binocular neurons", that is, they responded very weakly, if at all, to the monocular stimuli presented through either eye but responded vigorously to simultaneous stimulation through both eyes. Almost 70% (27/39) of neurons tested quantitatively for binocular interaction have shown significant modulation (over 50%) of their peak responses in relation to binocular positional retinal disparities. The majority of neurons sensitive to binocular positional disparities resembled either "tuned excitatory" (22 cells; 56.5% of the sample) or "tuned inhibitory" (2 cells; 5% of our sample) cells. In particular, they gave, respectively, maximal or minimal responses to optimally oriented, moving photic stimuli when the receptive fields plotted through each eye completely or partially overlapped. Although neurons recorded in area 21a have relatively large receptive fields (mean width 3.3 +/- 1.1 degrees; range 2.0-5.6 degrees), the mean width of the disparity tuning curve (2.8 +/- 1.0 degrees; range 1.3-4.8 degrees) for our sample of area 21a neurons was similar to those of neurons with significantly smaller receptive fields, recorded in areas 17 and 18 of cat's primary visual cortex. We conclude that area 21a of the cat, like areas 17 and 18 of primary visual cortex, is likely to play an important role in binocular depth discrimination and might constitute a "higher order" area for stereoscopic binocular vision.

Animals↗

Retinotopic organization of striate and extrastriate visual cortex in the hooded rat.

The visuotopic organization of the primary visual cortex (area 17) and the extrastriate visual regions surrounding it (areas 18a and 18) has been studied in gray rats using standard microelectrode mapping techniques. The results confirm and extend previous observations in the rat. Apart from the representation of the contralateral visual field (VF) in area 17, in which the upper VF is represented caudally and the nasal VF laterally, there are additional representations of the VF in the extrastriate cortex. In lateral extrastriate cortex (area 18a) there are at least 4 such representations, namely lateromedial (LM), anterolateral (AL), laterointermediate (LI) and laterolateral (LL). In LM (second visual area) the upper VF is represented caudally and the nasal VF medially, being thus a mirror image of V1. In AL (third visual area) the upper VF is represented rostrally and the nasal VF, medially, being thus a mirror image of LM. In LI, the upper VF is medial and the nasal VF, lateral, being thus a mirror image of LM, or a reduced copy of V1. In medial extrastriate cortex (area 18) there are two representations of the temporal VF, labeled anteromedial (AM) and posteromedial (PM). In AM, the upper temporal VF is medial and the lower temporal VF, lateral, the extreme temporal field being rostral. The 30 degrees azimuth provides the boundary between AM and PM. Thus, AM is organized as a counter-clockwise rotation by 90 degrees of the V1 representation. In PM, the upper lower VF topography is like in AM, but the extreme temporal VF is caudal, being thus a mirror image of AM.

Animals↗

A direct projection from area V1 to area V3A of rhesus monkey visual cortex.

Small cortical lesions were made in regions of the primary visual cortex (V1) representing different retinal eccentricities. It was found that, whereas all parts of V1 project to visual areas V2, V3 and the motion area of the superior temporal sulcus, only parts of V1 representing peripheral eccentricities (in excess of 30 degrees) project directly to visual area V3A.

Animals↗

Kindling of the visual cortex in cats: comparison with amygdaloid kindling.

Kindling of the primary visual cortex (VC) was compared with that of the amygdala in cats. VC kindling was basically similar to kindling of the amygdala in that daily electrical stimulation can lead to the development of a generalized convulsion in most subjects, a growth of afterdischarges in their configuration and duration, and a reduction of the afterdischarge threshold. The kindling response of the VC differed from that of the amygdala in a number of respects, i.e., a high afterdischarge threshold, a different pattern of behavioral seizure development, an abrupt growth of electroclinical seizures coincident with the onset of a generalized convulsion, an intersubject variability in seizure susceptibility, and a marked seizure instability. In VC kindling the afterdischarge propagation into the amygdala was not observed until the generalized convulsion developed, and the early involvement of afterdischarge was seen in the pulvinar, lateral geniculate body, and superior colliculus. These data suggest that a neural mechanism different from amygdaloid kindling may participate in VC kindling, and that the subcortical structures of the visual system are involved in the preferential pathway for a seizure generalization from the VC.

Amygdala↗

Mechanisms of inhibition in cat visual cortex.

1. Neurones from layers 2-6 of the cat primary visual cortex were studied using extracellular and intracellular recordings made in vivo. The aim was to identify inhibitory events and determine whether they were associated with small or large (shunting) changes in the input conductance of the neurones. 2. Visual stimulation of subfields of simple receptive fields produced depolarizing or hyperpolarizing potentials that were associated with increased or decreased firing rates respectively. Hyperpolarizing potentials were small, 5 mV or less. In the same neurones, brief electrical stimulation of cortical afferents produced a characteristic sequence of a brief depolarization followed by a long-lasting (200-400 ms) hyperpolarization. 3. During the response to a stationary flashed bar, the synaptic activation increased the input conductance of the neurone by about 5-20%. Conductance changes of similar magnitude were obtained by electrically stimulating the neurone. Neurones stimulated with non-optimal orientations or directions of motion showed little change in input conductance. 4. These data indicate that while visually or electrically induced inhibition can be readily demonstrated in visual cortex, the inhibition is not associated with large sustained conductance changes. Thus a shunting or multiplicative inhibitory mechanism is not the principal mechanism of inhibition.

Action Potentials↗

Noradrenergic modulation of functional selectivity in the cat visual cortex: an in vivo extracellular and intracellular study.

In vitro intracellular studies have shown that norepinephrine modulates cellular excitability and synaptic transmission in the cortex. Based on these effects, norepinephrine has been proposed to enhance the signal-to-noise ratio and to improve functional selectivity by potentiating strong synaptic responses and reducing weak ones. Here we have studied the functional effects of iontophoretic applications of norepinephrine during in vivo extracellular and intracellular recordings from neurons of the primary visual cortex of kittens and adult cats. Analysis of extracellular data concentrated on norepinephrine-induced changes in spontaneous and evoked activities, in signal-to-noise ratio, and in orientation and direction selectivity. Analysis of the intracellular data concentrated on actions of norepinephrine on spike firing accommodation, which has been shown to be reduced by norepinephrine in vitro, and on synaptic responses. Application of norepinephrine resulted in a depression of both spontaneous and evoked spiking activity. However, no systematic change in signal-to-noise ratio was observed. The suppressive effect of norepinephrine was exerted with no significant sharpening of direction or orientation selectivity tuning. The overall reduction in visual activity by norepinephrine affected the orientation tuning curves in a way compatible with a divisive effect, that is a normalization or gain control with no change in tuning width. Norepinephrine applied during intracellular recordings reduced the visually evoked depolarizing potentials whereas no change in the responsiveness of the cell to current-induced depolarizations was observed. In conditions of optimal visual stimulation which produced large depolarizations of several hundreds of milliseconds and sustained repetitive firing comparable to that obtained by direct current injection, we were unable to observe a facilitation of the evoked responses by norepinephrine as it would be expected from the well-documented increase in excitability induced by norepinephrine in vitro. In conclusion, from these results we suggest that norepinephrine released in the primary visual cortex primarily reduces the level of cortical activation by afferent signals, without affecting the cortical functional selectivity nor increasing the signal-to-noise ratio.

Action Potentials↗

Electrophysiological correlates of lateral interactions in human visual cortex.

Detection thresholds for visually presented targets can be influenced by the nature of information in adjacent regions of the visual field. For example, detection thresholds for low-contrast Gabor patches decrease when flanked by patches that are oriented collinearly rather than orthogonally with the target. Such results are consistent with the known microanatomy of primary visual cortex, where long-range horizontal connections link cortical columns with common orientation preferences. To investigate the neural bases of collinearity effects, we recorded event-related brain potentials (ERPs) together with psychophysical measures for targets flanked by collinear vs. orthogonal gratings. Human volunteers performed a contrast discrimination task on a target grating presented at a perifoveal location. For targets flanked by collinear stimuli, we observed an increased positive polarity voltage deflection in the occipital scalp-recorded ERPs between 80 to 140 ms after stimulus onset. Such a midline occipital scalp voltage distribution of this ERP collinearity effect is consistent with a generator in primary visual cortex. Two later negative voltage ERP deflections (latencies of 245-295 and 300-350 ms) were focused at lateral occipital scalp sites, a pattern consistent with activity in extrastriate visual cortex. These ERP effects were correlated with improved contrast discrimination for central targets presented with collinear flanks. These results demonstrate that the integration of local flanking elements with a central stimulus can occur as early as 80 ms in human visual cortex, but this includes processes occurring at longer latencies and appears to involve both striate and extrastriate visual areas.

Adolescent↗

Inhibition of ocular dominance column formation by infusion of NT-4/5 or BDNF.

During the development of the visual system of higher mammals, axons from the lateral geniculate nucleus (LGN) become segregated into eye-specific patches (the ocular dominance columns) within their target, layer 4 of the primary visual cortex. This occurs as a consequence of activity-dependent synaptic competition between axons representing the two eyes. The possibility that this competition could be mediated through neurotrophin-receptor interactions was tested. Infusion of neurotrophin-4/5 (NT-4/5) or brain-derived neurotrophic factor (BDNF) into cat primary visual cortex inhibited column formation within the immediate vicinity of the infusion site but not elsewhere in the visual cortex. Infusion of nerve growth factor, neurotrophin 3 (NT-3), or vehicle solution did not affect column formation. These observations implicate TrkB, the common receptor for BDNF and NT-4/5, in the segregation of LGN axons into ocular dominance columns in layer 4. Moreover, they suggest that in addition to their better known roles in the prevention of cell death, neurotrophins may also mediate the activity-dependent control of axonal branching during development of the central nervous system.

Animals↗

Adult cortical dynamics.

There are many influences on our perception of local features. What we see is not strictly a reflection of the physical characteristics of a scene but instead is highly dependent on the processes by which our brain attempts to interpret the scene. As a result, our percepts are shaped by the context within which local features are presented, by our previous visual experiences, operating over a wide range of time scales, and by our expectation of what is before us. The substrate for these influences is likely to be found in the lateral interactions operating within individual areas of the cerebral cortex and in the feedback from higher to lower order cortical areas. Even at early stages in the visual pathway, cells are far more flexible in their functional properties than previously thought. It had long been assumed that cells in primary visual cortex had fixed properties, passing along the product of a stereotyped operation to the next stage in the visual pathway. Any plasticity dependent on visual experience was thought to be restricted to a period early in the life of the animal, the critical period. Furthermore, the assembly of contours and surfaces into unified percepts was assumed to take place at high levels in the visual pathway, whereas the receptive fields of cells in primary visual cortex represented very small windows on the visual scene. These concepts of spatial integration and plasticity have been radically modified in the past few years. The emerging view is that even at the earliest stages in the cortical processing of visual information, cells are highly mutable in their functional properties and are capable of integrating information over a much larger part of visual space than originally believed.

Adult↗

The mystery of Louis Verrey (1854-1916).

In 1888, Louis Verrey, a Swiss ophthalmologist, stated emphatically that there is a "centre for the chromatic sense" in the human brain and that it is located in the lingual and fusiform gyri. He did not, however, consider the "colour centre" to be a separate area but a large sub-division of the primary visual cortex. His evidence was quickly dismissed and forgotten. It was not to be taken seriously again until after the experimental discovery of functional specialization in the monkey brain. This paper considers why it is that Verrey did not consider the "colour centre" to be a separate cortical area, distinct from the primary visual cortex, why his evidence was so quickly and effectively dismissed, and why it is that Verrey did not pursue the logic of his findings.

Animals↗