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PubMedDate not supplied

The role of ipsilateral and contralateral inputs from primary cortex in responses of area 21a neurons in cats.

Neuronal responses in cat visual area 21a were analyzed when the primary visual cortex (areas 17 and 18) was deactivated by cooling. Ipsilateral and contralateral cortices were deactivated separately. Results established that (1) cooling the ipsilateral primary cortex diminished the activity of all area 21a cells and, in 30%, blocked responsiveness altogether, and (2) cooling the contralateral primary cortex initially increased activity in area 21a cells but, with further cooling, reduced it to below the original level although only 9% of cells ceased responding. These findings were then compared to earlier results in which bilateral deactivation of the primary cortex greatly reduced and, in most cases, blocked the activity of area 21a cells (Michalski et al., 1993). Despite the response attenuation following cooling of the primary visual cortex (either ipsilateral or contralateral), neurons of area 21a retained their original orientation specificity and sharpness of tuning (measured as the half-width at half-height of the orientation tuning curve). Direction selectivity also tended to remain unchanged. We concluded that for area 21a cells (1) the ipsilateral primary cortex provides the main excitatory input; (2) the contralateral primary cortex supplies a large inhibitory input; and (3) the nature of orientation specificity, sharpness of orientation tuning, and direction selectivity are largely unaffected by removal of the ipsilateral hemisphere excitatory input or the contralateral hemisphere inhibitory input.

Animals↗
PubMed1985

Localization of glutaminase-like and aspartate aminotransferase-like immunoreactivity in neurons of cerebral neocortex.

The distribution of glutaminase (GLNase)- and aspartate aminotransferase (AATase)-immunoreactive cells was examined in the cerebral neocortex of rat and guinea pig and in the somatic sensorimotor and primary visual cortex of the Macaca fascicularis monkey. These enzymes are involved in the metabolism of glutamate and aspartate, two amino acids thought to be excitatory amino acid transmitters for cortical neurons. In each of the species examined a large percentage of layer V and VI pyramidal neurons have pronounced glutaminase-like immunoreactivity (GLNase IR). In contrast, neurons in layers I, II, and IV show little GLNase IR. Layer III in the rat and guinea pig contains only a few, densely labeled GLNase-like-immunoreactive (GLNase-Ir) pyramidal neurons, whereas in the monkey the number of GLNase-Ir cells in layer III varies between cytoarchitectonic fields. Area 3b of the primary somatic sensory cortex and area 17 (primary visual cortex) contain few GLNase-Ir cells in layer III. However, layer III contains moderate numbers of GLNase IR in cells in areas 3a, 1, 2, 5, and in the primary motor cortex. Within the motor cortex the largest pyramidal ("Betz") cells are not labeled. In marked contrast to the results with antibody to GLNase, antibody to AATase labels cells that appear nonpyramidal in form, and these cells are in all cortical layers in each of the species examined. This distribution is roughly similar throughout all areas of rodent neocortex, but in monkey visual cortex AATase-immunoreactive neurons are more numerous in layers II-III, IVc, and VI. When combined with the findings of other studies, our results suggest that GLNase IR marks pyramidal neurons that use an excitatory amino acid transmitter. Antibody to AATase appears to mark intrinsic cortical neurons. The AATase immunoreactivity of these cells could indicate that they use an excitatory amino acid transmitter. However, their form and distribution in cortex suggest that this antibody labels GABAergic neurons.

Animals↗
PubMed1993

Development of orientation selectivity in ferret visual cortex and effects of deprivation.

The orientation selectivity of cells in ferret primary visual cortex was studied during normal development and in animals deprived of vision or of visual cortical activity. In normal animals from the age when visual responses were first recorded (postnatal day 23) through postnatal week 5, only about 25% of cells showed orientation-selective responses. By postnatal week 7, cortical responses had matured to an adult-like state, with approximately 75% of cells clearly selective for orientation. This development of orientation selectivity was not merely a reflection of the development of cortical cell responsiveness: at all ages studied, there was no correlation between responsiveness and selectivity. Infusion of TTX into visual cortex to silence neuronal activity completely blocked the maturation of orientation selectivity. Visual deprivation by bilateral lid suture impaired but did not completely block the normal development of orientation selectivity. We conclude that the maturation of orientation-selective responses in ferret primary visual cortex requires cortical neuronal activity, and that normal development requires visually driven activity.

Animals↗
PubMed1988

The effects of ablation of visual cortex in neonatal rabbits on the organization of retinothalamic and retinopretectal projections.

Primary visual cortex was ablated unilaterally in neonatal rabbits. Following a survival of 2-4 months, retrograde degeneration of the dorsal lateral geniculate nucleus (LGd) was assessed, and reorganization of retinofugal pathways was studied using methods of anretrograde transport of [3H]proline or of horseradish peroxidase. A complete lesion of primary visual cortex resulted in complete retrograde degeneration of the LGd with no sparing of any class of neurons. The terminations of retinofugal axons in the pretectum and thalamus were compared with those observed in normal animals. No major reorganization of ipsilateral retinofugal projections was observed in either the thalamus and pretectum ipsilateral to the ablated cortex, or in the thalamus and pretectum contralateral to the ablated cortex. However, contralateral retinofugal projections to the thalamus and to the pretectum ipsilateral to the ablated cortex were significantly different from normal. In the thalamus, the projections to the lateral posterior nucleus were expanded in area and increased in density. In the pretectum, the projections to the rostral pretectal areas were greatly increased in area, especially in the region of the olivary pretectal nucleus and posterior pretectal nucleus. However, the density of these projections was not increased relative to normal. Consideration of these results in relation to other published data on the anatomical consequences of neonatal visual cortex lesions, both in mammals which show behavioral sparing following neonatal visual cortex lesions and in mammals which, like the rabbit, show no behavioral sparing, suggests that: (1) behavioral sparing may correlate with patterns of survival or death of neurons in the thalamus and retina; and (2) reorganization of retinofugal pathways is not necessarily associated with behavioral sparing.

Animals↗
PubMed1989

Intraocular injections of tetrodotoxin reduce transiently expressed acetylcholinesterase activity in developing rat visual cortex.

Geniculo-recipient layers of primary visual cortex in the rat display a transient pattern of acetylcholinesterase (AChE) activity during the second postnatal week of life. Previous work has demonstrated that neonatal enucleations markedly reduce the transient AChE activity in visual cortex. The present studies were undertaken to determine the effects of reduced afferent neural activity on expression of the transient pattern of AChE activity. Rat pups received intraocular injections of tetrodotoxin (TTX) on postnatal days (PND) 3, 5, 7, 9 and 11 and were sacrificed on PND 12. Some animals were enucleated on PND 3. Brain sections were processed for AChE histochemistry and analyzed by optical densitometry. These experiments show that uniocular injections result in a markedly decreased level of AChE activity in layer IV of the medial part of cortical area 17 contralateral to the injected eye. The degree of reduction of AChE activity from repeated TTX injections was similar to the degree of reduction following enucleation on PND 3. Binocular injections of TTX result in a reduction of AChE activity in layer IV throughout cortical area 17, similar to the effects of binocular enucleation on PND 3. Experiments combining injection of horseradish peroxidase along with TTX on PND 11 demonstrate that retinal ganglion cells of TTX injected eyes are still capable of anterograde axonal transport. These data demonstrate that normal innervation and afferent activity are necessary for the transient expression of AChE activity by geniculocortical neurons.

Acetylcholinesterase↗
PubMed1990

Lateral interactions in visual cortex.

The findings presented in these studies have brought out different ideas concerning the mechanisms of processing in primary visual cortex than were held at the outset. Rather than thinking of receptive fields as being restricted in their extent, with the process of integration of the components of an image occurring at a much later stage along the visual pathway, we have shown that the integrative process is a progressive one, beginning in the primary visual cortex (or perhaps even earlier) and building up in a cascading series of converging and diverging connections. Rather than thinking of the filter characteristics of a cell as being fixed, it is apparent that they are dynamic and can be modified by the context in which features are presented. Finally, rather than a cortex with a functional architecture that is fixed after a critical period ending in infancy, we find that perturbing the system can lead to long-term topographical reorganization. Other examples of contextual interactions have been demonstrated in the submodalities of motion, where a cell's directional selectivity is modulated by the presence of movement in the surround (Allman et al. 1985; Tanaka et al. 1986; Gulyas et al. 1987; Orban et al. 1987). In the domain of color, the phenomenon of color constancy, reported for cells in visual area V4 (Zeki 1983), also requires lateral interactions in visual space, comparing the wavelength distribution of light coming from surfaces in different parts of the visual field. The influences presented in these studies, as in our own work in the domain of orientation, are modulatory. The long-term changes in cortical topography following removal of somatosensory input (Merzenich et al. 1984, 1988) or by retinal lesions suggest that with the appropriate manipulations the lateral interactions can be enhanced to the point of activating the postsynaptic cells. Although retinal lesions clearly represent an abnormal disruption of sensory input, they may nevertheless be representative of long-term reorganizations of neural networks occurring under normal circumstances, such as those required for memory.

Animals↗
PubMed1977

[Types of receptive fields of neurons in different laminae of the rabbit visual cortex].

Neurons of the primary rabbit visual cortex were classified into 7 large groups according to features of their receptive fields. The neurons with receptive fields were mostly revealed in the layers IV and VI, those with uniform directional receptive fields--in layer V, those with simple I-in layer VI, those with simple II--in layer II + III, and those with hypercomplex--in layer IV. The neurons with concentric receptive fields and those without responses to visual stimuli were equally distributed over the layers. The data obtained in rabbits, cats, and monkeys suggest that the tendency towards function stratification of primary visual cortex is successive in an evolutionary order of mammals.

Animals↗
PubMed1990

Quantitative analysis of a vulnerable subset of pyramidal neurons in Alzheimer's disease: II. Primary and secondary visual cortex.

In this study we investigated the primary and secondary visual areas of normal and Alzheimer's disease brains by using the SMI32 antibody. It is known that in Alzheimer's disease primary sensory areas are usually less devastated than association cortices, although visual symptomatology has been documented early in the course of the disease. In area 17, the SMI32 antibody primarily labeled the perikarya and dentritic tree of the large Meynert cells and cells in layer IVB. Smaller neurons in layers III, V, and VI were also immunoreactive (ir). In area 18, very large SMI32-ir pyramidal neurons in layers III and V were observed. In both areas, staining intensity was correlated with cell size, the largest neurons being the most intensely stained. Only a few changes were observed in the Alzheimer's disease cases. The only statistically significant differences in SMI32-ir neuron counts between control and Alzheimer's disease brains occurred in layer IVB cells and Meynert cells in area 17, and in layer III cells in area 18. In contrast with association cortices, there were no changes in staining intensity in the visual areas. There were fewer neurofibrillary tangles and neuritic plaques in these areas than in prefrontal and inferior temporal cortex, and a correlation between neurofibrillary tangle counts and SMI32-ir neuron loss was only observed in layer III of area 18. These observations show that in the primary and secondary visual cortex, SMI32 also labeled a distinct subset of pyramidal cells that are known from data obtained in the monkey brain to furnish long corticocortical as well as subcortical projections. Interestingly, although there is much less cell and/or neurofibrillary tangle formation in these occipital regions than in prefrontal and temporal association areas, there is significant loss within key subsets of pyramidal cells. The selective loss of this particular subpopulation of pyramidal neurons will disrupt association pathways linking primary visual cortex with areas involved in higher level visual processing. The partial disconnection of such pathways may be relevant to the visual symptomatology frequently observed in Alzheimer's disease patients. These data further support the hypothesis that subtypes of pyramidal neurons with specific anatomical and molecular profiles may display a differential vulnerability in Alzheimer's disease.

Aged↗
PubMed1988

Coherent oscillations: a mechanism of feature linking in the visual cortex? Multiple electrode and correlation analyses in the cat.

Primary visual coding can be characterized by the receptive field (RF) properties of single neurons. Subject of this paper is our search for a global, second coding step beyond the RF-concept that links related features in a visual scene. In recent models of visual coding, oscillatory activities have been proposed to constitute such linking signals. We tested the neurophysiological relevance of this hypothesis for the visual system. Single and multiple spikes as well as local field potentials were recorded simultaneously from several locations in the primary visual cortex (A17 and A18) using 7 or 19 individually advanceable fiber-microelectrodes (250 or 330 microns apart). Stimulus-evoked (SE)-resonances of 35-85 Hz were found in these three types of signals throughout the visual cortex when the primary coding channels were activated by their specific stimuli. Stimulus position, orientation, movement direction and velocity, ocularity and stationary flicker caused specific SE-resonances. Coherent SE-resonances were found at distant cortical positions when at least one of the primary coding properties was similar. Coherence was found 1) within a vertical cortex column, 2) between neighbouring hypercolumns, and 3) between two different cortical areas. We assume that the coherence of SE-resonances is mediated by recurrent excitatory intra- and inter-areal connections via phase locking between assemblies that represent the linking features of the actual visual scene. Visually related activities are, thus, transiently labelled by a temporal code that signalizes their momentary association.

Action Potentials↗
PubMed1993

Relationship between intrinsic connections and functional architecture revealed by optical imaging and in vivo targeted biocytin injections in primate striate cortex.

In primate primary visual cortex, neurons sharing similar response properties are clustered together forming functional domains that appear as a mosaic of patches or bands, often traversing the entire cortical depth from the pia to the white matter. Similarly, each cortical site connects laterally through an extensive network of intrinsic projections that are organized in multiple clusters (patches) and reach distances of up to a few millimeters. The relationship between the functional domains and these laterally connected patches has remained a controversial issue despite intensive research efforts. To investigate this relationship, we obtained high-resolution functional maps of the cortical architecture by in vivo optical imaging. Subsequently, extracellular injections of the sensitive anterograde tracer biocytin were targeted into selected functional domains. Within the ocular dominance system, we found that long-range intrinsic connections tended to link the monocular regions of same-eye ocular dominance columns. Furthermore, we discovered that binocular domains formed a separate set of connections in area V1; binocular regions were selectively connected among themselves but were not connected to strictly monocular regions, suggesting that they constitute a distinct columnar system. In the other subsystem subserving orientation preference, patches of intrinsic connections tended to link domains sharing similar orientation preferences. Analyses of the precision of these connections indicated that in both functional subsystems, < 15% of the connections were between domains having orthogonal response properties. However, their selectivity was limited; approximately 30% +/- 10% of the interconnected patches contained neurons exhibiting orientation tuning that differed from those found at the injection sites by at least 45 degrees. At short range (up to 400 microns from the injection site), this casual trend seemed markedly accentuated; the local, synaptic-rich axonal and dendritic arbors crossed freely through columns of diverse functional properties. These complex sets of connections can endow cortical neurons with a rich diversity of response properties and broad tuning.

Animals↗
PubMed1982

Single unit receptive fields in rabbit primary binocular cortex.

The receptive fields of 125 single units recorded from the binocular region of rabbit primary visual cortex have been analysed. The population of 43% radially symmetric, 23% directional, and 23% orientation selective units is similar to that of rabbit monocular visual cortex. The relative scarcity of orientation selective units and the absence of orientation columns differentiates rabbit from cat primary visual cortex. However, the majority of binocular units had similar receptive fields in each eye and very unconventional receptive fields were not encountered. Tested binocular units demonstrated summation upon simultaneous stimulation of both receptive fields. In conjunction with findings reported elsewhere, these results suggest that rabbit and cat possess a similar provision for binocular vision in spite of some differences in their cortical organisation.

Animals↗
PubMed1988

Imaging of cerebral blood flow and metabolism in amblyopia by positron emission tomography.

We used positron emission tomography to study monocular visual activation of various brain regions in four amblyopic and two normally sighted adults. Imaging of relative cerebral blood flow using the tracer H2(15)O showed reduced activation of primary visual cortex by the amblyopic as compared with the sound eye. Imaging of relative cerebral glucose metabolism using the tracer [18F]-2-deoxyglucose showed equal activation of primary visual cortex by either eye in the control subject, but reduced activation of primary and accessory visual cortex by the amblyopic as compared with the sound eye in two amblyopic subjects. Relative glucose metabolism was consistently higher in the frontal and temporal lobes contralateral to the viewing eye, both in normal and amblyopic subjects.

Amblyopia↗
PubMed1991

Stereopsis in the cat: behavioral demonstration and underlying mechanisms.

The neural substrates subserving stereopsis were investigated behaviorally and electrophysiologically in the cat. In one set of studies, we examined behaviorally the ability of normal cats to perceive depth on the sole basis of spatial disparity using random-dot stereograms. Results showed that the animals were able to carry out this discrimination. We then evaluated the contribution of the optic chiasm, the corpus callosum and the primary visual cortex to this function. Results indicated that: (1) chiasma transection drastically reduced the ability of the animals to solve the random-dot problem; (2) a callosal split had little or no effect on their ability to relearn the same discrimination; (3) a section of both the corpus callosum and optic chiasm abolished this ability; and (4) bilateral lesions of areas 17-18 also abolished it. In another set of studies, we examined electrophysiologically the properties of neurons in the various visual cortical areas where disparity-based depth discrimination processes are presumed to take place. We recorded from areas 17, 18 and 19 of normal and split-chiasm cats. Results showed that: (1) the primary visual cortex of the normal cat contained cells sensitive to stimulus disparity; (2) these disparity sensitive neurons were also present in area 19 although in a much lower proportion and were more widely tuned than those in areas 17-18; and (3) following the section of the optic chiasm, there was a significant decrease in the number of disparity sensitive cells in areas 17-18, whereas in area 19 they were nearly completely absent. The results obtained from the lesion studies and from the single unit recording experiments indicate that stereoscopic depth perception is highly dependent in the cat upon the integrity of the through-the-chiasm geniculo-striate pathway and its target primary visual cortex.

Animals↗
PubMed1982

Facilitatory effects of intermittent photic stimulation on visual cortical kindling.

There are few reports on kindling preparation by daily electrical stimulation to the primary visual cortex. In our previous studies on cats, it was demonstrated that repeated stimulation to the primary visual cortex brought a similar progress in seizure development as in amygdaloid kindling, although it took more trials. In this study, we examined the effects of intermittent photic stimulation (IPS) (2.5 Hz) on the visual cortical kindling preparation to clarify more underlying mechanisms of photosensitivity. Under IPS, afterdischarges synchronizing to IPS were seen and a duration of afterdischarge was extremely prolonged, even in the early stage of the kindling process. The threshold of stimulus intensity inducing afterdischarge was significantly reduced and the generalized convulsive seizure was obtained with fewer times of stimulation, as compared to visual cortical kindling without IPS. In visual cortical kindling myoclonic responses were sometimes caused by IPS for a few minutes immediately after cessation of the electrically induced generalized seizure. Such phenomena were never seen in amygdaloid kindling. These results indicate that IPS facilitates a seizure development in all stages of visual cortical kindling.

Animals↗
PubMed1989

Identification and localization of 5-hydroxytryptamine receptor sites in macaque visual cortex.

The two main receptor subtypes for 5-hydroxytryptamine (5HT) were measured and localized in visual cortical areas of macaque monkey. [3H]5HT was used to label all 5HT-1 receptor subtypes and [3H]ketanserin was used to label 5HT-2 receptors. Both receptor types could be demonstrated in membranes prepared from macaque primary visual cortex. The specificity of these ligands for 5HT-1 or 5HT-2 receptors was demonstrated by the pharmacological profile of inhibitors of the specific binding. 5HT-1A receptor sites were detected by displacement experiments and by direct labeling with [3H]8-hydroxy-2(di-n-propylamino) tetralin 8OH-DPAT. Receptor autoradiography showed that the distribution of these receptor subtypes varied from one part of visual cortex to another. 5HT-1 receptors, labeled with [3H]5HT were present in several bands through layer IV of primary visual cortex with the densest band seen in and above layer IVA: another band was in lower layer VI. The band in layer VI was predominantly 5HT-1A sites. There were two main bands of 5HT-2 receptor sites, the most prominent around the IV/V boundary, and the other extending from layer IVA upwards. Adjacent areas showed 5HT receptors in a broad band corresponding to layer IV. 5HT-1A sites were found in superficial layers of adjacent areas, except V2. These layering patterns did not correspond precisely with cytoarchitectonic layering, nor with the pattern of 5HT-containing presynaptic fibres in published reports. It is important, therefore, in considering the role of the 5HT-containing neurons in cortical function to take account not only of the anatomy of the presynaptic terminals, but also of the postsynaptic receptors upon which the released transmitter will act, and their location within the cortex.

8-Hydroxy-2-(di-n-propylamino)tetralin↗
PubMedDate not supplied

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↗
PubMed1990

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↗
PubMedDate not supplied

[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↗