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Contrast constancy in natural scenes in shadow or direct light: A proposed role for contrast-normalisation (non-specific suppression) in visual cortex.

The range of contrasts in natural scenes is generally thought to far exceed the limited dynamic ranges of individual contrast-encoding neurons in the primary visual cortex. The visual system may employ gain-control mechanisms (Ohzawa et al. 1985) to compensate for the mismatch between the range of natural contrast energies and the limited dynamic range of visual neurons; one proposed mechanism is contrast normalisation or non-specific suppression (Heeger 1992a). This paper aims to evaluate the role of contrast normalisation in human contrast perception, using a computer model of primary visual cortex. The model uses orthogonal pairs of Gabor patches to simulate simple-cell receptive-fields to calculate local, band-limited contrast in a series of 50 digitised photographs of natural scenes. The average range of contrast energies in each image was 2.29 log units, while the "lifetime range" each model simple cell would see across all images was 2.98 log units. These ranges are greater than the dynamic range of real mammalian simple cells. Contrast normalisation (dividing contrast responses by the summed responses of all nearby neurons) reduces contrast ranges, perhaps sufficiently to match them to neurons' limited dynamic ranges. Comparison of images taken under diffuse and direct lighting conditions showed that contrast normalisation can sometimes match these conditions effectively. This may lead to perceptual contrast constancy in the face of spurious changes in contrast caused by natural environmental conditions.

Computer Simulation↗

Dissociation between two modes of spatial processing by a visual form agnosic.

We report a dissociation between two aspects of visuospatial processing in a patient with a profound impairment in the visual perception of objects ('visual form agnosia'). The orientation-in-depth of a visual field ('visual pitch') was found to systematically influence the elevation at which she perceived her own eye level, just as it does in normal individuals; but at the same time, she was unable to discriminate perceptually the orientation-in-depth of the same visual field, a trivial task for individuals with normal vision. These results suggest that, in the normal brain, the processes that integrate orientation information from the visual field with extraretinal information about eye position are separable from those supporting the perception of the orientation of the visual field itself. The pattern of brain damage in D.F., in conjunction with the reported dissociation, suggests that the former set of processes maps onto the stream of information flowing from primary visual cortex to the posterior parietal cortex, the so-called dorsal stream, whereas the latter involves the projections from primary visual cortex to the inferotemporal cortex, the so-called ventral stream.

Adult↗

Contrast adaptation in striate cortical neurons of the nocturnal primate bush baby (Galago crassicaudatus).

It has been argued that in order for the visual system to detect edges accurately under a range of conditions, the visual system needs to adapt to the local contrast level to preserve sensitivity (Blakemore & Campbell, 1969). Cells in the primary visual cortex of cats adapt to stimuli with low to moderate contrast. Curiously, macaque monkey neurons in primary visual cortex (V1) do not show evidence for similar adaptation. To address the question of whether this differential sensitivity in contrast adaptation might be due to phylogenetic variation between cats and primates or to specializations for visual niche (e.g. nocturnal vs. diurnal), contrast adaptation to temporally and spatially optimized gratings was examined in 30 V1 cells of three nocturnal primate bush babies (Galago crassicaudatus). A second objective was to examine the relationship between the degree of contrast adaptation and cell classification or cell location relative to cortical layers or compartments [i.e. cytochrome-oxidase (CO) blobs and interblobs]. All cells were classified (simple vs. complex) and anatomically localized relative to cortical layers and cytochrome-oxidase (CO) blob and interblob compartments. Two independent measures of contrast adaptation were used. In the first test, contrast was sequentially increased from 3-56% and then decreased. The contrast required to maintain a half-maximum response amplitude in the 30 cells tested increased an average of 0.24 (+/- 0.12) log units during the sequential decrements in contrast. For the second test, four sets of five interleaved contrasts within +/- 1 octave of a central adapting contrast (10%, 14%, 20%, and 28%, respectively) were presented. The cells produced a mean adaptation index of 0.57 (+/- 0.47) which is very similar to that exhibited by cat cortical neurons (0.54 +/- 0.41). Interestingly, cells in interblobs showed a trend toward greater adaptation than did blob cells. Moreover, cells in the supragranular layers exhibited greater adaptation than cells in the infragranular layers. No significant differences in adaptation were found to correlate with other cell classification indices. Taken together, our results suggest that contrast adaptation may be more important for maintaining sensitivity in nocturnal species (primates or cats) than in diurnal species (macaque monkeys), and that in the nocturnal bush baby, cells in cortical layers and compartments may be differentially specialized for contrast adaptation.

Adaptation, Ocular↗

A model for the estimate of local image velocity by cells in the visual cortex.

Some computational theories of motion perception assume that the first stage en route to this perception is the local estimate of image velocity. However, this assumption is not supported by data from the primary visual cortex. Its motion sensitive cells are not selective to velocity, but rather are directionally selective and tuned to spatio-temporal frequencies. Accordingly, physiologically based theories start with filters selective to oriented spatio-temporal frequencies. This paper shows that computational and physiological theories do not necessarily conflict, because such filters may, as a population, compute velocity locally. To prove this point, we show how to combine the outputs of a class of frequency tuned filters to detect local image velocity. Furthermore, we show that the combination of filters may simulate 'Pattern' cells in the middle temporal area (MT), whereas each filter simulates primary visual cortex cells. These simulations include three properties of the primary cortex. First, the spatio-temporal frequency tuning curves of the individual filters display approximate space-time separability. Secondly, their direction-of-motion tuning curves depend on the distribution of orientations of the components of the Fourier decomposition and speed of the stimulus. Thirdly, the filters show facilitation and suppression for responses to apparent motions in the preferred and null directions, respectively. It is suggested that the MT's role is not to solve the aperture problem, but to estimate velocities from primary cortex information. The spatial integration that accounts for motion coherence may be postponed to a later cortical stage.

Animals↗

Extra-classical receptive field effects measured in striate cortex with fMRI.

The aim of this study was to measure the contextual influence of globally coherent motion on visual cortical responses using functional magnetic resonance imaging. Our motivation was to test a prediction from representational theories of perception (i.e. predictive coding) that primary visual responses should be suppressed by top-down influences during coherent motion. We used a sparse stimulus array such that each element could not fall within the same classical receptive field of primary visual cortex neurons (i.e. precluding lateral interactions within V1). This enabled us to attribute differences, in striate cortex responses, to extra-classical receptive field effects mediated by backward connections. In accord with theoretical predictions we were able to demonstrate suppression of striate cortex activations to coherent relative to incoherent motion. These results suggest that suppression of primary visual cortex responses to coherent motion reflect extra-classical effects mediated by backward connections.

Adult↗

Preliminary FMRI evidence of visual system dysfunction in Parkinson's disease patients with visual hallucinations.

Using functional magnetic resonance imaging (fMRI), the authors examined visual cortex function in Parkinson's disease patients who did and did not experience visual hallucinations. Patients with visual hallucinations demonstrated increased activation in the visual association cortex and deficits in the primary visual cortex, suggesting that visual hallucinations are associated with an abnormality of visual-cortex function.

Aged↗

Functional plasticity in extrastriate visual cortex following neonatal visual cortex damage and monocular enucleation.

Neonatal lesions of primary visual cortex (areas 17, 18 and 19; VC) in cats lead to significant changes in the organization of visual pathways, including severe retrograde degeneration of retinal ganglion cells of the X/beta class. Cells in posteromedial lateral suprasylvian (PMLS) cortex display plasticity in that they develop normal receptive-field properties despite these changes, but they do not acquire the response properties of striate neurons that were damaged (e.g., high spatial-frequency tuning, low contrast threshold). One possibility is that the loss of X-pathway information, which is thought to underlie striate cortical properties in normal animals, precludes the acquisition of these responses by cells in remaining brain areas following neonatal VC damage. Previously, we have shown that monocular enucleation at the time of VC lesion prevents the X-/beta-cell loss in the remaining eye. The purpose of the present study was to determine whether this sparing of retinal X-cells leads to the development of striate-like response properties in PMLS cortex. We recorded the responses of PMLS neurons to visual stimuli to assess spatial-frequency tuning, spatial resolution, and contrast threshold. Results indicated that some PMLS cells in animals with a neonatal VC lesion and monocular enucleation displayed a preference for higher spatial frequencies, had higher spatial resolution, and had lower contrast thresholds than PMLS cells in cats with VC lesion alone. Taken together, these results suggest that preserving X-pathway input during this critical period leads to the addition of some X-like properties to PMLS visual responses.

Animals↗

Disruption of orientation tuning in visual cortex by artificially correlated neuronal activity.

In the primary visual cortex, the development of orientation selectivity is influenced by patterns of neural activity. The introduction of artificially correlated activity into the visual pathway (through synchronous activation of retinal ganglion cell axons in the optic nerve) substantially weakens the orientation selectivity of neurons in superficial and deep cortical layers. This is consistent with activity having an instructive role in shaping cortical neuron receptive field tuning properties.

Action Potentials↗

An amplitude equation approach to contextual effects in visual cortex.

A mathematical theory of interacting hypercolumns in primary visual cortex (V1) is presented that incorporates details concerning the anisotropic nature of long-range lateral connections. Each hypercolumn is modeled as a ring of interacting excitatory and inhibitory neural populations with orientation preferences over the range 0 to 180 degrees. Analytical methods from bifurcation theory are used to derive nonlinear equations for the amplitude and phase of the population tuning curves in which the effective lateral interactions are linear in the amplitudes. These amplitude equations describe how mutual interactions between hypercolumns via lateral connections modify the response of each hypercolumn to modulated inputs from the lateral geniculate nucleus; such interactions form the basis of contextual effects. The coupled ring model is shown to reproduce a number of orientation-dependent and contrast-dependent features observed in center-surround experiments. A major prediction of the model is that the anisotropy in lateral connections results in a nonuniform modulatory effect of the surround that is correlated with the orientation of the center.

Animals↗

[Relationship between level of vigilance and changes in the receptive fields of the cat visual cortex].

The activity of 118 neurones of the primary visual cortex (17th field) was studied in unanesthetized cats, immobilized with d-tubocurarine, in a state of calm wakefulness and in strained attention, alarm. The strained attention was elicited by an air-puff directed to the corner of the closed eye, not used for the photic stimulation. Considerable rearrangments of the receptive field of neurones (94%) were observed in the state of strained attention. In the majority of the studied cortical cells (75%) during 10-30 minutes the excitatory centres of their receptive fields became narrower while their inhibitory periphery widened; in the cells with the inhibitory centres of the receptive fields they were larger in 80% of the cases. It is shown that with dark, mesopic and scotopic adaptation the major characteristics of the changes in the receptive fields remain unaltered. It is assumed that these effects have a behavioural value for the organism, because the created rise in the level of alertness is accompanied by a sharpening of the receptive fields, i.e. by a greater ability to a fine analysis and recognition of visual images.

Adaptation, Physiological↗

Neural networks for Braille reading by the blind.

To explore the neural networks used for Braille reading, we measured regional cerebral blood flow with PET during tactile tasks performed both by Braille readers blinded early in life and by sighted subjects. Eight proficient Braille readers were studied during Braille reading with both right and left index fingers. Eight-character, non-contracted Braille-letter strings were used, and subjects were asked to discriminate between words and non-words. To compare the behaviour of the brain of the blind and the sighted directly, non-Braille tactile tasks were performed by six different blind subjects and 10 sighted control subjects using the right index finger. The tasks included a non-discrimination task and three discrimination tasks (angle, width and character). Irrespective of reading finger (right or left), Braille reading by the blind activated the inferior parietal lobule, primary visual cortex, superior occipital gyri, fusiform gyri, ventral premotor area, superior parietal lobule, cerebellum and primary sensorimotor area bilaterally, also the right dorsal premotor cortex, right middle occipital gyrus and right prefrontal area. During non-Braille discrimination tasks, in blind subjects, the ventral occipital regions, including the primary visual cortex and fusiform gyri bilaterally were activated while the secondary somatosensory area was deactivated. The reverse pattern was found in sighted subjects where the secondary somatosensory area was activated while the ventral occipital regions were suppressed. These findings suggest that the tactile processing pathways usually linked in the secondary somatosensory area are rerouted in blind subjects to the ventral occipital cortical regions originally reserved for visual shape discrimination.

Adult↗

Plasticity in the adult visual cortex: implications for the diagnosis of visual field defects and visual rehabilitation.

It has recently been shown that, contrary to long-held beliefs, sensory and motor maps are not stable in the adult cerebral cortex. Alteration of input from the periphery results in changes in topography in the cortex, including the primary visual cortex. Mechanisms involved consist mainly of reshaping the receptive field of cortical cells and increasing the sensitivity of deprived cells in the visual cortex. Cortical plasticity allows the brain to adapt to background modifications or to damage of the nervous system. It also underlies learning and attention processes. Cortical changes occurring after focal visual differentiation modify visual perception by filling in visual field defects with information from the area surrounding the scotoma. This modification causes affected subjects to ignore or underestimate their defects. With visual field defects, cortical plasticity also causes distortion in spatial perception. Although the effects of cortical plasticity are prominent in neuro-ophthalmological daily practice, they are usually unrecognized or greatly underestimated. These effects cause delay in recognizing visual field defects, and hence in receiving therapy, while affecting the results of some procedures for testing the visual field. Affected individuals who are unaware of their defects may have increased difficulty in coping with activities in everyday life. Up to now, phenomena related to plasticity in the visual system have been investigated mainly by psychophysicists and neurophysiologists. It is essential to start considering the various effects of cortical reorganization in clinical practice. It is especially important to introduce into clinics the concept of dissociation between actual and perceived defects in the visual field, resulting from the filling-in process, and the need to measure it. This dissociation should also be demonstrated to the affected subjects.

Adult↗

[Examination of the visual system with transcranial magnetic stimulation].

The influence of transcranially applied magnetic stimuli on the function of the afferent (sensory) and efferent (motor) parts of the visual system have been discussed. Excitatory (positive) phenomena are subjective photic sensations (phosphenes) which can be elicited by transcranial magnetic stimulation over occipital parts of the skull. The phosphenes appear on the left or right side of the visual field depending upon the direction of the coil currents, which determines whether the visual cortex of the right or the left hemisphere is activated. The configuration of the phosphene fields hints at an excitation of the primary visual cortex (Brodmann's area 17). However, magnetic brain stimulation also produces inhibitory (negative) phenomena. When strong magnetic field pulses are applied over the primary visual cortex, foveally presented visual stimuli cannot be identified even when no phosphenes are perceived at the same time. Depending on the position of the stimulation coil, this suppression of perception can be restricted to visual stimuli presented on the right, or left of, above or below the fixation point. No generation or disturbance of eye movements by transcranial magnetic stimulation has been reported before, except for a delay of saccades within a reaction time paradigm.

Animals↗

Calcium-binding protein-containing neuronal populations in mammalian visual cortex: a comparative study in whales, insectivores, bats, rodents, and primates.

This study is focused on comparative analysis of gamma-aminobutyric acid-positive (GABAergic) neuronal populations in primary visual cortex of totally aquatic toothed whales and select terrestrial mammals with different evolutionary histories and various ecological adaptations. The distribution of neuronal populations containing the calcium-binding proteins calbindin and parvalbumin, which are recognized markers for the GABAergic neurons in cerebral cortex, is compared in five species of toothed whales and in representatives (one species each) of insectivores, bats, rodents, and primates. Computerized image analysis has shown that overall quantitative characteristics of GABAergic cortical neurons in toothed whales are similar to those in other mammalian orders. Thus, GABA-positive neurons represent 26% of the total population of cortical neurons in the visual cortex of whales. Some 97% of GABA-positive cells contain calcium-binding proteins, which is numerically similar to these parameters found in primates and other mammals. On the other hand, the typology and laminar distribution of calcium-binding protein-containing neurons in the primary visual cortex of five whale species (Delphinapterus leucas, Globicephala melaena, Phocoena phocoena, Stenella coeruleoalba, and Tursiops truncatus) differ significantly from those of primates (Macaca mulatta) and rodents (Rattus rattus) and are similar to those found in insectivorous bats (Eptesicus fuscus) and hedgehogs (Erinaceus europaeus). In whales, bats, and hedgehogs a significant concentration of calbindin-positive, vertically oriented bipolar and bitufted neurons was found in layers I, II, and IIIc/V with their axons arranged in a three-dimensional network. In primates and rodents they are distributed evenly across all cortical layers and are predominantly multipolar or bitufted neurons found in all cortical layers with their axons oriented along the vertical axis of the cortical plate. The parvalbumin-positive neurons in all mammalian species, including toothed whales, are represented by variously sized multipolar non-pyramidal cells. As opposed to all other mammalian species, the major concentrations of parvalbumin-positive neurons in whales are found in layers IIIc/V and VI, whereas in other cortical layers there are only scattered parvalbumin-positive neurons.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Detection of visual dysfunction in optic atrophy by functional magnetic resonance imaging during monocular visual stimulation.

PURPOSE: To evaluate functional magnetic resonance imaging as an objective method for detecting visual dysfunction in various ophthalmologic disorders involving the optic nerve and the chiasm. METHODS: We performed functional magnetic resonance imaging during monocular visual stimulation on seven patients with visual field loss caused by lesions of the optic nerve and the chiasm and on three normal control subjects with no visual field loss. We correlated static threshold perimetry in the seven patients with the results of functional magnetic resonance imaging. RESULTS: In the three normal control subjects, we found good intrasubject similarity in areas of bilateral occipital lobe activation between monocular stimulation of the right and left eyes. In the patients with unilateral optic neuropathy, including glaucoma, stimulation of the affected eye induced no activation of the primary visual cortex in the portion corresponding to the central visual field defects and reduced activity of the associated visual cortex. In the patients with chiasmal compression, monocular stimulation resulted in a marked asymmetry of activation in the primary visual cortex, which corresponded to the visual field abnormality. CONCLUSIONS: Functional magnetic resonance imaging appears to be useful in confirming the clinical diagnosis of optic atrophy because it can objectively disclose visual field loss, even a small defect such as central scotoma.

Adult↗

Activation of striate cortex in the absence of visual stimulation: an fMRI study of synesthesia.

It has been suggested that internally generated visual perception involves the primary visual cortex V1. To test this hypothesis, a functional MRI study was conducted with a female subject with orthographic color-word synesthesia. This subject was selected as she reported clear involuntary visualization of auditorily presented verbal material. Hearing a word resulted in seeing the word in a particular color. fMRI scans were acquired while the subject performed two verbal tasks (passive listening to words and verbal fluency). Significant activity was detected in primary visual cortex, in the absence of external visual stimulation. This finding provides evidence for a role of modulatory feedback connections between associative and primary visual areas in visual experience without direct visual stimulation.

Acoustic Stimulation↗

The hierarchical development of monkey visual cortical regions as revealed by the maturation of parvalbumin-immunoreactive neurons.

The prefrontal cortex is known to be involved in behavioral paradigms requiring decisions based on short-term working memory, and visually related areas of prefrontal cortex represent the final point in a proposed hierarchical sequence of visual signal processing that begins in primary visual cortex. This study asks if the development of at least certain aspects of the circuitry of each region involved in this hierarchy proceeds in a sequential fashion from primary to higher-order areas. The timing and patterns of expression of immunoreactivity for the calcium-binding protein parvalbumin were examined in areas V1, V2, TE, 7a, and 46 in two series of macaque monkeys ranging in age from embryonic day 132 to adult. The number and laminar distribution of parvalbumin-labeled neurons reached adult levels first in area V1 (primary visual cortex), followed by the adjacent visual association area V2, and then by the higher-order regions of the inferior temporal (TE), posterior parietal (7a) and prefrontal (46) cortices. The appearance of parvalbumin immunoreactivity in the axons of the two major classes of local circuit neurons that express this protein, basket and chandelier cells, followed a similar regional pattern. Furthermore, striking differences were present between these two neuronal populations in the laminar pattern and time course of parvalbumin labeling of their axons. These findings demonstrate that at least some aspects of the intrinsic circuitry of the neocortex mature in accordance with a functional hierarchy of cortical regions. In addition, they illustrate the complexity of cortical development in terms of the different timing of expression of even a single protein in different compartments within single neurons, in different cell types, in different laminae within a region, and across different cortical regions.

Animals↗

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.

Adult↗