Surround suppression sharpens orientation tuning in the cat primary visual cortex

In the primary visual cortex (V1), the response of a neuron to stimulation of its classical receptive field (CRF) is suppressed by concurrent stimulation of the extraclassical receptive field (ECRF), a phenomenon termed 'surround suppression'. It is also known that the orientation tuning of V1 neurons becomes sharper as the size of the stimulus increases beyond the CRF. However, there have been few quantitative investigations of the relationship between sharpening of orientation tuning and surround suppression. We examined this relationship in 73 V1 neurons recorded from anesthetized and paralysed cats using sinusoidal grating patches as stimuli. We found that sharpening of orientation tuning was significantly correlated with the strength of surround suppression for large stimuli that cover both CRF and ECRF. Furthermore, simulation analysis using a variety of tuning widths and most suppressive orientation of orientation-tuned surround suppression demonstrated that broadly orientation-tuned surround suppression sharpens orientation tuning for large gratings without shift in optimal orientation. Our findings suggest that one of the functional roles of surround suppression in V1 is enhancement of orientation discrimination for large and uniformly patterned objects.     

Surround suppression by high spatial frequency stimuli in the cat primary visual cortex

Surround suppression is a phenomenon whereby stimulation of the extraclassical receptive field suppressively modulates the visual responses of neurons in the primary visual cortex (V1) (also known as area 17). It is known that surround suppression tunes to spatial frequencies (SFs) that are much lower and broader than the frequencies to which the classical receptive field tunes. In this study, we tested the effects of varying SFs on surround suppression by using a circular sinusoidal grating patch that covered both the classical receptive field and the extraclassical receptive field. Using area-summation tuning curves, we found high-SF-tuned surround suppression in the cat V1. This high-SF-tuned surround suppression causes the SF tuning to shift to low SF for large stimuli. By simulating a model neuron lacking a suppressive surround mechanism, we confirmed that these preferred SF shifts do not occur in the absence of surround suppression. We surmise that the high-SF-tuned suppression, which shifts the preferred SF according to size, functionally contributes to the scale-invariant processing of visual images in V1.     

Contrast-dependent, contextual response modulation in primary visual cortex and lateral geniculate nucleus of the cat

In the primary visual cortex (V1), the responses of neurons to stimuli presented in their classical receptive fields (CRFs) are modulated by another stimulus concurrently presented in their surround (receptive field surround, SRF). We studied the nature of the modulatory effects of SRF stimulation with respect to stimulus contrast in cat V1. In 51 V1 neurons studied, large SRF stimuli (40 degreesx30 degrees ) induced only the suppression of responses to CRF stimulation and the suppressive effects became stronger as the contrast for SRF stimulation increased. The contrast sensitivity of SRF suppression did not correlate with that of CRF responses. By independently controlling contrast of CRF and SRF stimuli, we studied whether SRF effects vary with CRF response magnitude. Increasing contrast for CRF stimulation caused an upward shift of the range of effective contrasts for SRF stimulation, indicating that a high contrast for SRF stimulation is required for suppressing strong responses to CRF stimulation at high contrasts. To assess the possible origin of the suppressive SRF effect on V1 neurons, we also investigated the contrast dependency of SRF effects in 28 neurons from the lateral geniculate nucleus. Our results suggest that SRF effects obtained at the subcortical level strongly contribute to those in V1. Taken together, we conclude that along the thalamocortical projections, SRF modulation exhibits a gain-control mechanism that scales the suppressive SRF effect depending on the contrast for CRF stimulation. In addition, SRF effects can be facilitatory at low stimulus contrasts potentially due to the enlargement of the summation field.     

Spike synchronization in cat primary visual cortex depends on similarity of surround‐suppression magnitude

In the primary visual cortex (V1), the spike synchronization seen in neuron pairs with non‐overlapping receptive fields can be explained by similarities in their preferred orientation (PO). However, this is not true for pairs with overlapping receptive fields, as they can still exhibit spike synchronization even if their POs are only weakly correlated. Here, we investigated the relationship between spike synchronization and suppressive modulation derived from classical receptive‐field surround (surround suppression). We found that layer 4 and layer 2/3 pairs exhibited mainly asymmetric spike synchronization that had non‐zero time‐lags and was dependent on both the similarity of the PO and the strength of surround suppression. In contrast, layer 2/3 and layer 2/3 pairs showed mainly symmetric spike synchronization that had zero time‐lag and was dependent on the similarity of the strength of surround suppression but not on the similarity in POs. From these results, we propose that in cat V1 there exists a functional network that mainly depends on the similarity in surround suppression, and that in layer 2/3 neurons the network maintains surround suppression that is primarily inherited from layer 4 neurons.     

Temporal properties of colour opponent receptive fields in the cat lateral geniculate nucleus

The primordial form of mammalian colour vision relies on opponent interactions between inputs from just two cone types, ‘blue’ (S‐) and ‘green’ (ML‐) cones. We recently described the spatial receptive field structure of colour opponent blue‐ON cells from the lateral geniculate nucleus of cats. Functional inputs from the opponent cone types were spatially coextensive and equally weighted, supporting their high chromatic and low achromatic sensitivity. Here, we studied relative cone weights, temporal frequency tuning and visual latency of cat blue‐ON cells and non‐opponent achromatic cells to temporally modulated cone‐isolating and achromatic stimuli. We confirmed that blue‐ON cells receive equally weighted antagonistic inputs from S‐ and ML‐cones whereas achromatic cells receive exclusive ML‐cone input. The temporal frequency tuning curves of S‐ and ML‐cone inputs to blue‐ON cells were tightly correlated between 1 and 48 Hz. Optimal temporal frequencies of blue‐ON cells were around 3 Hz, whereas the frequency optimum of achromatic cells was close to 10 Hz. Most blue‐ON cells showed negligible response to achromatic flicker across all frequencies tested. Latency to visual stimulation was significantly greater in blue‐ON than in achromatic cells. The S‐ and ML‐cone responses of blue‐ON cells had on average, similar latencies to each other. Altogether, cat blue‐ON cells showed remarkable balance of opponent cone inputs. Our results also confirm similarities to primate blue‐ON cells suggesting that colour vision in mammals evolved on the basis of a sluggish pathway that is optimized for chromatic sensitivity at a wide range of spatial and temporal frequencies.     

Visual response properties of ventral lateral geniculate nucleus cells projecting to the pretectum and superior colliculus in the cat

The visual properties of cells in the cat ventral lateral geniculate nucleus (LGV) identified antidromically from the pretectum and/or superior colliculus (projection cells) were studied in comparison with those of LGV neurons which could not be activated antidromically (non-projection cells). ON-phasic receptive fields (RFs) were relatively predominant in 27 projection cells, whereas ON-tonic RFs were found more commonly in the non-projection group. The distribution of the RF centers revealed a centroperipheral gradient of the visual field representation within the LGV that the central visual field was more densely organized.     

Synaptic reorganization in the dorsal lateral geniculate nucleus following damage to visual cortex in newborn or adult cats

We have studied the effects of making large lesions of visual cortex on the synaptic organization of the dorsal lateral geniculate nucleus (LGN) in the cat. Visual cortex was removed at birth in one group of cats and during adulthood in a second group. Following survival periods of 6 months to 2 years, the organization of synapses made by afferents from the retina in the LGN was investigated quantitatively with the electron microscope. In single thin sections we determined the percentage of retinal axon terminals that made synapses in the LGN, the average number of synapses made by each retinal axon terminal, and the identity of each postsynaptic process. These measurements were made separately for retinogeniculate connections in the A and C laminae of the LGN. For comparison, similar sets of measurements were made in adult cats that had been reared normally. When single thin sections from the A or C laminae of the LGN in normal cats are examined, about 60% of the axon terminals from the retina are seen to make at least one synaptic contact. These contacts can be with dendrites or F profiles or both. On average, each retinogeniculate terminal makes approximately 1.4 synapses in the plane of a single section and contacts dendrites three times as often as F profiles. In the A laminae of the LGN in cats that received a visual cortex lesion at birth or in adulthood, the percentage of retinal terminals that make synapses is the same as in normal cats. Similarly, the average number of synaptic contacts made by each retinogeniculate terminal is not changed by a lesion of visual cortex. In contrast, the number of contacts made with dendrites is reduced markedly, by about 29% after a lesion at birth and 53% after a lesion as an adult. However, these reductions are offset by compensatory increases in the number of contacts made with F profiles, and thus the mean number of contacts made by each retinogeniculate terminal is stabilized at a normal value. In the C laminae of the LGN, retinogeniculate terminals also reapportion their synaptic contacts. In cats with a lesion during adulthood, the redistribution of synapses is compensatory, as in the A laminae. When a lesion is made at birth, however, the number of new retinal contacts made with F profiles exceeds the number of dendritic contacts that are lost. As a result, each retinogeniculate terminal makes about 26% more synapses, in total, than normal.(ABSTRACT TRUNCATED AT 400 WORDS)     

Role of GABAergic inhibition in shaping the spatial frequency tuning of neurons and its contrast dependency in the dorsal lateral geniculate nucleus of cat

To understand how GABAergic inhibition contributes to the elaboration of spatial frequency (SF) tuning of neurons in the lateral geniculate nucleus, we examined the effects of the microiontophoretic administration of bicuculline methiodide (BIC) and gabazine, both antagonists of GABA_A receptors, on visual responses to grating stimuli with high or low contrasts. BIC administration changed the shape of the SF tuning curve of the spike response from band‐pass to low‐pass. We took the tuning curve obtained under the BIC condition as an estimated excitatory contribution to the control tuning curve and then estimated the difference between tuning curves recorded with and without BIC as the tuning curve of the estimated GABAergic inhibitory contribution. The SF tuning profile of estimated inhibition (Estimated‐Inh) varied widely from cell to cell, as did estimated excitation (Estimated‐Ex). Nonetheless, the relationship that Estimated‐Inh exhibited more low‐pass tuning than did Estimated‐Ex was well conserved in the majority of cells, and the relationship refined the SF tuning of Estimated‐Ex toward the band‐pass tuning of the geniculate output. Lowering the stimulus contrast decreased the response magnitude, but did not change the degree of band‐pass tuning. The GABAergic refinement of the SF tuning was also observed at low stimulus contrast, but was weaker than at high contrast, suggesting that GABAergic inhibition is regulated in coordination with excitatory inputs to keep the degree of the band‐pass tuning constant. We therefore concluded that the degree of band‐pass tuning is conserved contrast invariantly in the lateral geniculate nucleus on the basis of the dynamic regulatory action of GABAergic inhibition.     

The cholinergic influence on the function of the cat dorsal lateral geniculate nucleus (dLGN)

The functional influence of the cholinergic input to cat dLGN has been examined by assessing the action of iontophoretically applied acetylcholine (ACh) on the visual responses of cells in layers A and A1. Iontophoretically applied pulses of ACh exerted a strong excitatory action on all 113 cells studied within these layers. In the presence of a sustained application of ACh, the excitatory responses to an optimal stimulus such as a spot of light located within the receptive field centre were greatly facilitated, but at the same time stimulus-specific inhibitory influences were also enhanced. The action of ACh on the stimulus-specific inhibitory influences had the consequence that the responses to non-optimal stimuli were not facilitated to the same extent as those to optimal stimuli and in some cases even diminished. The stimulus-specific inhibitory effects seen in the presence of ACh were very powerful and frequently resulted in complete suppression of the elevated background discharge. We suggest that the ACh directly excites both the relay cells and the Golgi type II inhibitory interneurones within the dLGN. The facilitation of the stimulus-specific inhibition may follow from a direct action on the presynaptic dendrites of the Golgi type II cells which arborize within the dendritis field of the relay cell. Supplementary observations on cells in the perigeniculate nucleus confirm previous findings showing that ACh has an inhibitory effect on these cells. We suggest a tripartite action for the cholinergic influence on the dLGN, involving direct facilitation of relay cells, enhancement of stimulus-specific inhibition via the Golgi type II cells, and disinhibition of the non-specific inhibitory influence from the perigeniculate nucleus.     

Subdivisions of the dorsal raphe nucleus projecting to the lateral geniculate nucleus and primary visual cortex in the Mongolian gerbil

The mammalian dorsal raphe nucleus (DRN) is composed of sub-divisions with different anatomical and functional properties. Using cholera toxin subunit B as a retrograde tracer, DRN subdivisions projecting to the lateral geniculate nucleus and to the primary visual cortex were examined in the Mongolian gerbil. DRN neurons projecting to the lateral geniculate nucleus were observed in the lateral DRN (rostrally) and in the ventromedial DRN (caudally), while DRN cells projecting to the primary visual cortex were observed at all rostral-caudal levels in the ventromedial DRN. These results demonstrate a significant overlap between the DRN projections to the lateral geniculate and superior colliculus, and show that only the caudal ventromedial DRN projects to all three major visual targets: the lateral geniculate nucleus, primary visual cortex, and superior colliculus. Since the DRN is involved in depression and other neuropsychiatric disorders, as well as is affected by many psychotropic substances, these data may help to develop new treatments and therapies targeting specific DRN subdivisions.     

Temporal interactions in the cat visual system. II. Suppressive and facilitatory effects in the lateral geniculate nucleus

Extracellular responses were recorded from single neurons in the lateral geniculate nucleus (LGN) of the cat during presentation of pairs of brief visual stimuli identical to those that produce orientation-selective paired-pulsed suppression in the visual cortex. LGN neurons also show paired-pulse suppression, but the suppression is not orientation selective, and it occurs only for short interstimulus intervals (ISIs; usually less than 200 msec). At longer ISIs, most LGN neurons show a period of facilitation. Thus, the paired-pulse suppression in the LGN cannot account for that seen in the visual cortex. Paired-pulse suppression in the LGN was found to be enhanced by stimulation of the receptive field surround. LGN neurons also showed a second type of suppression, termed "offset suppression," which consisted of a more long-lasting suppression of spontaneous activity following the offset of an excitatory visual stimulus. The suppression of spontaneous activity was accompanied by a reduction of the antidromic excitability, assessed by stimulating LGN axons within the cortex or optic radiation. Unlike paired-pulsed suppression, offset suppression was not enhanced by increased stimulation of the receptive field surround. Paired-pulse suppression and offset suppression are most likely due to different mechanisms because they have different time courses and depend differently on the spatial properties of the stimuli. Functionally, paired-pulse suppression may be related to the reduced visual sensitivity that accompanies eye movements, while offset suppression may serve to enhance temporal contrast.     

Distribution of neurofilament proteins in the lateral geniculate nucleus, primary visual cortex, and area MT of adult Cebus monkeys

We investigated the distribution pattern of SMI-32-immunopositive cells in the lateral geniculate nucleus (LGN) and in the primary (V1) and middle temporal (MT) cortical visual areas of the adult New World monkey Cebus apella. In the LGN, the reaction for SMI-32 labeled cells in both the magnocellular (M) and parvocellular (P) layers. However, the cellular label was heavier in M layers, which also showed a more intense labeling in the neuropil. In V1, the reaction showed a lamination pattern, with the heaviest labeling occurring in layer 4B and upper layer 6 (layers that project to area MT). Area MT shows a dense band of labeled neuropil and large pyramidal neurons in layer 3, large darkly labeled but less densely packed neurons in layer 5, and a population of small, lightly labeled cells in layer 6. These results resemble those found in other New and Old World monkeys, which suggest that the preferential labeling of projection neurons associated with fast-conducting pathways to the extrastriate dorsal stream is a common characteristic of simian primates. In the superficial layers of V1 in Cebus monkeys, however, SMI-32-labeled neurons are found in both cytochrome oxidase blobs and interblob regions. In this aspect, our results in Cebus are similar to those found in the Old World monkey Macaca and different from those described for squirrel monkey, a smaller New World Monkey. In Cebus, as well as in Macaca, there is no correlation between SMI-32 distribution and the blob pattern.     

Influence of the visual cortex upon receptive field organization of lateral geniculate cells in rabbits

In anesthetized rabbits, the receptive fields of lateral geniculate cells were mapped prior to and following the interruption of the corticogeniculate feed-back. Visual cortex (V.C.) was depressed by a focal application of 3 M KCl. The responsiveness of the V.C. was verified by monitoring the visually evoked potentials. In off- and on-center cells, the surround excitatory responses were remarkably reduced and even fully abolished in most units. In contrast, the center excitation remained unmodified. These effects were reversible. In some on-center units the center response had also decreased, and was replaced by an evoked inhibitory response. Relay cells and interneurons which yielded on and off responses over the entire area of the receptive field exhibited a loss of only one of the evoked discharges. It is concluded that the V.C. exerts mostly a specific desinhibitory action upon the geniculate network. This action affects either the center or the surround responses differentially. The results are compared with those obtained from cats.     

Patterns of fiber degeneration in the dorsal lateral geniculate nucleus of the cat following lesions in the visual cortex

Golgi preparations show a group of relatively fine axons ending in the dorsal lateral geniculate nucleus. These, the type I axons, are characterized by many short collateral terminal branches and form a well oriented plexus within the lateral geniculate nucleus. Individual axons distribute terminals to more than one geniculate lamina and some can be seen to come from the region of the internal capsule. Four or more days after damage to the visual cortex, the Nauta method shows relatively fine degenerating fibers that approach the lateral geniculate nucleus from a rostral direction. Some pass straight into lamina a, While others enter the three major laminae via the central interlaminar nucleus. The appearance of this degeneration suggests that the type I axons are cortico-genciulate, as does the sequence of the diencephalic degenerative changes. Thus, relatively coarse corticofugal fibers to the lateral thalamus and tectum show heavy degeneration within three days. The fine axons in the lateral geniculate nucleus show slight, early degeneration after three days, extensive degeneration after four days. They are possibly collaterals of the coarser corticofugal fibers. A second group of coarse fibers which runs laterally from the lateral geniculate nucleus shows early degeneration after four days but does not show consistent and clear degeneration until five or more days. Since retrograde cell changes are already recognizable after four days the third group is regarded as geniculo-cortical.     

The ventral lateral geniculate nucleus in the cat: thalamic and commissural connections revealed by the use of WGA-HRP transport

The present investigation was carried out to clarify the topographical details of both the origin and terminal site of the thalamic projections and the commissural connections of the ventral lateral geniculate nucleus (LGNv) in the cat by using bidirectional transport of wheat germ agglutinin conjugated to horseradish peroxidase (WGA-HRP). Thalamic projections: Unilateral injections of WGA-HRP into the LGNv produced orthograde labeling in the intralaminar nuclei bilaterally and in the lateralis posterior (LP) and the pulvinar (Pul) nucleus ipsilaterally. In the intralaminar nuclei the rostral part of the nucleus centralis lateralis (CL) was most densely labeled by orthogradely transported material, particularly in its dorsal and lateral large-celled portion. Other intralaminar nuclei--such as the nucleus paracentralis, centralis medialis, and centralis dorsalis--also were labeled bilaterally with ipsilateral predominance, but no labeling was detected in the caudal portion of the CL and the centromedian and parafascicular nuclei. In the Pul, labeling of terminal ramifications was found to be concentrated in a region just medial to the so-called retinorecipient zone of the Pul as a slim band of labeling inclining dorsoventrally. In the LP, fine labeled fibers were located in the lateral portion of the LP. Commissural connections: Commissural fibers crossed in the dorsal part of the posterior commissure and reached the most caudal part of the contralateral LGNv. Labeling in the contralateral LGNv was concentrated in the dorsomedial part of the medial zone that extends medially to the middle portion of the cerebral peduncle. Origins of the commissural connections arose mostly from the medial zone that roughly corresponds to the commissural terminal zone and partly from aberrant cells dispersed among optic tract fibers. From these results, together with the previous studies, it is concluded that although the cat's LGNv has connections with diverse structures in the central nervous system, the origin and terminal site of the connections are partially segregated within the nucleus, which suggests that the LGNv may contain functional subsystems.     

Development of the projections from the dorsal lateral geniculate nucleus to the lateral suprasylvian visual area of cortex in the cat.

In the study reported in the preceding paper, we used retrograde labeling methods to show that the enhanced projection from the thalamus to the posteromedial lateral suprasylvian (PMLS) visual area of cortex that is present in adult cats following neonatal visual cortex damage arises at least partly from surviving neurons in the dorsal lateral geniculate nucleus (LGN). In the C layers of the LGN, many more cells than normal are retrogradely labeled by horseradish peroxidase (HRP) injected into PMLS cortex ipsilateral to a visual cortex lesion. In addition, retrogradely labeled cells are found in the A layers, which normally have no projection to PMLS cortex in adult cats. The purpose of the present study was to investigate the mechanisms of this enhanced projection by examining the normal development of projections from the thalamus, especially the LGN, to PMLS cortex. Injections of HRP were made into PMLS cortex on the day of birth or at 1, 2, 4, or 8 weeks of age. Retrogradely labeled neurons were present in the lateral posterior nucleus, posterior nucleus of Rioch, pulvinar, and medial interlaminar nucleus, as well as in the LGN, at all ages studied. Within the LGN of the youngest kittens, a small number of retrogradely labeled cells was present in the interlaminar zones and among the cells in the A layers that border these zones. Such labeled cells were virtually absent by 8 weeks of age, and they are not found in normal adult cats. Sparse retrograde labeling of C-layer neurons also was present in newborn kittens.(ABSTRACT TRUNCATED AT 250 WORDS)     

Lack of visual suppression in the rabbit lateral geniculate nucleus during blink reflex

Stimulation of the supraorbital branch of the trigeminal nerve (SO) elicited eye blinks in the rabbit, but did not decrease the amplitude of visual cortical evoked potential from stimulation of the optic chiasm (OX). In addition, the SO stimulation neither induced an inhibitory postsynaptic potential (IPSP) in LGN cells, nor activated inhibitory interneurons in the thalamic reticular nucleus (TRN), which proved to mediate both recurrent inhibition and saccadic suppression in the dorsal lateral geniculate nucleus (LGN). All these indicate that there is no visual suppression in the rabbit LGN during blink reflex.     

Relay of visual information to the lateral geniculate nucleus and the visual cortex in albino ferrets

The abnormal organization of the central visual pathways in the albino ferret has been characterized anatomically and physiologically. Recordings in dorsal lateral geniculate nucleus of the albino ferret show that lamina A1, which receives an aberrant projection from the contralateral eye, contains an extensive representation of the ipsilateral visual hemifield with receptive fields located up to 35 degrees from the vertical meridian. This is not the case in pigmented ferrets, for which the vast majority of units, activated through either the contralateral or ipsilateral eye, have receptive fields confined to the contralateral hemifield. The few fields found in the ipsilateral hemifield are driven through the contralateral eye and none is more than 10 degrees from the midline. Cortical topography was studied by making closely spaced electrode penetrations across the area 17/18 border. In pigmented animals, the reversal of topography at the border is characterized by units with receptive fields centered a few degrees into the ipsilateral hemifield. In 22 of 25 albinos, the "Boston" aberrant topography was found: the representation of the vertical meridian is within area 17, rather than at the area 17/18 border. Instead, at the area 17/18 border, there is a reversal in the topographic progression at up to 30 degrees into the ipsilateral hemifield. This pattern was most pronounced in the upper visual field. In agreement with the "Boston" physiology, injections of retrograde tracer made in area 17 usually label neurons in either lamina A or the part of lamina A1 that is aberrantly innervated by the contralateral eye. A column of labeled cells extending through all geniculate layers is rarely seen in albinos, although this is commonly the pattern in pigmented ferrets.     

Structural development of the lateral geniculate nucleus and visual cortex in monkey and man

This study concerns the development of the primary visual pathway of the primate. The lateral geniculate nucleus (LGN) is the principal thalamic relay to the visual cortex (area 17), and its neurons have similar morphological characteristics in both monkey and man, as identified by Golgi impregnation. The commonest neuron is the multipolar with a radiate or tufted dendritic tree; next is the bipolar neuron with two or three diametrically opposed dendritic trunks. Less frequent are neurons with beaded dendrites and others with fine, axon-like dendritic processes, possibly interneurons. The dendritic tree of all neurons remains generally within a lamina, but some dendrites cross interlaminar zones. LGN neurons are identifiable before birth and differ from their adult form by the presence of immature features, especially numerous dendritic and somatic spines, most frequent at birth in monkeys and at about 4 months postnatally in man. They disappear almost completely by 3 months in monkeys and 9 months in man. The human LGN has reached its ‘adult’ volume by this age.Two stages in the development of the human area 17 can be defined. The first is marked by a rapid growth to its ‘adult’ volume by about 4 months, and by intense synaptogenesis beginning in the foetus and reaching a maximum around 8 months. The second stage is one of stabilization in the volume of area 17 and loss of synapses to reach ‘adult’ synaptic density around 11 years, at about 60% of the maximum values.The formation of transitory morphological features in the first weeks or months of life coincides with a period of visual plasticity in infant monkeys and humans. Our observations can be correlated with experimental evidence for visual development in monkeys and with clinical evaluation of visual activity during the human preverbal stage, a period of great importance in the establishment of visual acuity, of stereopsis and of oculomotor function, all very sensitive to the numerous forms of visual deprivation.