Effects of visual cortex lesions upon the visual fields of monocularly deprived cats.
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[Neurons of the rabbit visual cortex with simple and complex visual fields].
The receptive filelds of orientation-selective neurons were studied in the rabbit visual cortex. Two mechanisms of the selectivity were found: the mutual inhibition between on- and off-regions of the receptive field (simple type) and the orientation-dependent inhibition within the uniform region of the receptive field. The non-selective neurons could exhibit the lateral inhibition within the uniform region of the receptive field also. It is supposed that both "simple" and "complex" cells originate from the non-selective units.
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[Heterogeneity of a neuron population with complex receptive fields in the visual cortex of costs].
Visual responses of the striate complex cells to stimuli orientation, direction and velocity of movement were studied in awakening, unanesthetized cats. "Complex" cells were divided into four groups according to the response characteristics which were obtained using a stationary slit, moving light spot and moving oriented stimulus. The first group units response characteristics suggest the presence of the orientation selectivity mechanism in their receptive field organization, the fourth group--the direction selectivity mechanisms, the second and third groups--the presence of both mechanisms. It is supposed that there are two separate mechanisms for coding the orientation and direction of the stimulus movement in the neuronal structures of the visual cortex.
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Physiological consequences for the cat's visual cortex of effectively restricting early visual experience with oriented contours.
1. The early visual experience of nine cats was restricted to viewing horizontal or vertical lines inside opaque goggles. 2. When the kittens were 3-4 mo old, extracellular recordings were made in the primary visual cortex. To obtain a representative sample of cortical cells, units were studied at regularly spaced intervals along the course of electrode penetrations traveling oblique to the cortical surface. An automated assessment of preferred orientation using a computer-driven optical display was employed, and during the recording session the experimenters did not know which orientation(s) each animal had viewed in early life. 3. In the cats that viewed horizontal lines with one eye and vertical lines with the other during rearing, two major findings of previous workers (14) were confirmed. First, a majority of units were not selective for orientation. Second, units with preferred orientations near vertical tended to be activated exclusively by the eye that had viewed vertical, and likewise for horizontal. 4. In cats that viewed lines of the same orientation with both eyes during rearing, a substantially smaller proportion of units were selective for orientation; the preferred orientations of these units also tended to match the orientation to which the cats had been exposed. 5. Portions of some electrode penetrations showed an orderly arrangement of cells according to preferred orientation similar to that seen in normal cats, but with regions over which only nonselective cells were found. Many penetrations appeared less orderly. 6. The results are consistent with a role for early visual experience in maintaining the responsiveness and innate selectivity of cortical neurons, although they cannot entirely rule out the possibility that experience may alter or determine the preferred orientation of some cells.
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Mapping the representation of the visual field by electrical stimulation of human visual cortex.
Electrical stimulation of human visual cortex produces punctuate phosphenes in the visual field. This phenomenon, which is being explored as the basis for a visual prosthesis for the blind, also provides the first electrophysiological information about the retinocortical map in man. Stimulation of points clustered on the surface of the visual cortex produces phosphenes clustered in visual space. However, adjacent surface electrodes located on opposite sides of a sulcus can produce widely separated phosphenes, because the intervening cortex is buried and inaccessible to stimulation. Such electrodes can also produce multiple phosphenes by simultaneously stimulating both banks of the sulcus. Electrodes which are widely spaced on the brain can produce phosphenes close together in visual space providing they stimulate cortex corresponding to overlapping maps in areas 17 and 18. Analysis of the phosphene map indicates that successive stimulation of points further from the tip of the occipital pole produces phosphenes progressively more distant from the fixation point. Successive stimulation of points along the orthogonal dorsoventral dimension produces a progressive change in phosphene bearing. These results confirm the general view of cortical organization derived from field defect studies in man, and from anatomical and electrophysiological studies in monkeys, and provide a new tool for more detailed study of retinotopic projections in man.
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[Responses of rabbit visual cortex neurons to non-visual stimuli and their complexes with light].
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Postnatal development of interhemispheric connections of the cat visual cortex.
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Influence of association cortex on penicillin discharges in the primary visual cortex.
The influence of visual association cortex on the pattern of neuronal activity in the lateral geniculate body, during penicillin-induced discharges in visula cortex, was studied. In a majority of those geniculate cells which exhibited a pre-penicillin discharge increment in activity, that increment was abolished or significantly reduced during focal hypothermia of the visual association cortex. The frequency of penicillin discharges in visual cortex was, in most cases, reduced during cooling of association cortex. These findings are briefly discussed in terms of corticothalamic interactions.
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Postnatal dendritic development in the rabbit visual cortex.
Golgi preparations of rabbit visual cortex aged 1-25 days, as well as similar tissues from adults, were examined for the growth of the dendritic arbor, and in particular the development of dendritic spines. The layer 5 pyramidal neurons and layer 4 stellate neuron were chosen as representatives of larger classes of neurons in the visual cortex. It was determined that the growth of the dendritic arbor, determined by counts of total number of dendritic and total dendritic length, is quite similar for pyramidal and stellate neurons. Dendritic spine development, however, is more rapid in pyramidal neurons than in stellate. This disparity in the rate of dendritic spine development is discussed in the light of physiologic studies on the development of receptive field properties in the rabbit visual cortex.
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Retinotopic organization of the guinea pig's visual cortex.
Under anaesthesia the right visual cortex of the guinea pig was investigated with 1.5 M potassium citrate-filled glass microelectrodes. Single unit and small unit cluster response showed an organized three-part representation of the contralateral visual field on the animal's visual cortex. The major central division (V1) had the representation of a nasotemporally elongated visual field. This visual field extended for about 110 degrees horizontally and for about 70 degrees vertically. The nasal visual field was represented laterally on the cortex and the lower visual field anteriorly. The visual field representation was not uniformly magnified in V1; the magnification of the nasal field was higher than the rest of the visual field. On either side of V1, the guinea pig's cortex had two additional visually responsive strips. The strip V2L, situated lateral to V1, had a condensed representation of about 40 degrees of the animal's nasal visual field and mirror-imaged the visual field representation of the adjacent V1. The strip V2M, situated medial to V1, had a similar representation of about 40 degrees of the animal's temporal visual field and mirror-imaged the visual field representation of the area V1 adjacent to it. A binocularly responsive zone of cortex was observed on either side of the boundary between the areas V1 and V2L, representing between 10 degrees and 15 degrees of the nasal visual field on each side of the boundary line. Investigation of the retinal ganglion cell distribution of the animal showed a nasotemporally oriented 'visual streak' with a high ganglion cell density per unit area of the retina, near the optic disc. Away from the 'visual streak' the density of the ganglion cells of the retina diminished progressively. The configuration of the 'visual streak' and the general pattern of the ganglion cell distribution in the rest of the retina coincided with the asymmetry in the magnification of representation of the animal's visual field on the cortex.
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Neurophysiological mechanisms of recovery from visual cortex damage in cats: properties of lateral suprasylvian visual area neurons following behavioral recovery.
Damage to visual cortical areas 17, 18, and 19 in the cat produces severe and long-lasting deficits in performance of form and pattern discriminations. However, with extensive retraining the animals are able to recover their ability to discriminate form and pattern stimuli. Recent behavioral experiments from this laboratory have shown that a nearby region of cortex, the lateral suprasylvian visual area (LS area), plays an important role in this recovery (Wood et al., 1974; Baumann and Spear, 1977b). The present experiment investigated the underlying neurophysiological mechanisms of the recovery by recording from single neurons in the LS area of cats which had recovered from long-term visual cortex damage. Five adult cats received bilateral removal of areas 17, 18, and 19. They were then trained to criterion on two-choice brightness, form, and pattern discriminations. Recording from LS area neurons was carried out after the behavioral training, from 3 to 7 months after the visual cortex lesions. The properties of these neurons were compared to those of LS area neurons in normal cats (Spear and Baumann, 1975) and in cats with acute or short-term visual cortex damage and no behavioral recovery (Spear and Baumann, 1979). The results showed that all of the changes from normal which were produced by acute visual cortex damage were also present after the behavioral recovery. Moreover, all of the response properties of LS area neurons which remain after acute visual cortex damage were present in similar form after the behavioral recovery. There was no evidence for any functional reorganization in the LS area concomitant with its role in the behavioral recovery. These results suggest that functional reorganization plays little or no role in recovery from visual cortex damage in adult cats. Rather, the recovery of form and pattern discrimination ability appears to be based upon the functioning of residual neural processes in the LS area which remain after the visual cortex damage.
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Synaptic development in the rabbit superior colliculus and visual cortex.
The development of synapses in the visual cortex (VC) and superior colliculus (SC) of the rabbit has been examined with the electron microscope. In both areas, the number of synapses reaches adult levels by 20--25 days of postnatal age, but the development in the visual cortex is delayed in comparison to that in the superior colliculus. When S synapses (spheroidal vesicles, asymmetric thickening) are compared with F synapses (flattened vesicles, symmetric thickening), even greater differences are seen. In both the VC and SC, S synapses develop earlier than F synapses, though there is considerable overlap. Of interest is that fact that synapses in the visual cortex seem to overshoot their adult levels late in development, suggesting that an excess of synapses may be formed in this system. Multiple synapses, probably of retinal origin, increase in the first 3 weeks of synaptic development in the SC, but never are present in significant proportions in the VC. Synapse formation most often is characterized by formation of a junction and a postsynaptic thickening, followed by acquisition of synaptic vesicles. After 15 days, there is only a small number of such "non-vesicle synapses" in either the SC or VC.
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[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.
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Behavioral study of the visual cortex of Galago senegalensis.
An ablation study of the visual cortex of Galago senegalensis was undertaken in the hope of finding clues about the evolution of primate visual cortex. Removal of area 17 resulted in a profound sensory loss manifested by, first, the failure to discriminate between simple patterns; second, a deficit in localizing objects; third, a deficiency in tracking moving objects; and fourth, symptoms attributable to a deficiency in depth perception, such as misreaching and inaccurate jumping. Thus, the effects of ablating area 17 are similar in bushbabies and monkeys. In contrast, minimal sensory loss is produced by ablating area 17 in squirrels or tree shrews. This difference between primates and other mammals may depend on differences in the extent of the cortical target of the tecto-pulvinar path; in Galago and perhaps in all primates, more of the extrastriate visual cortex is entirely dependent on area 17. Removal of the ventral temporal cortex resulted in a loss of learned visual discriminations and in retardation in learning new visual discriminations. These symptoms seem related to the inferotemporal syndrome in monkeys.
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The rabbit and the cat: a comparison of some features of response properties of single cells in the primary visual cortex.
Receptive field characteristics of single cells in primary visual cortex of rabbit were studied. Seventy-two percent of cells were found to be orientation selective, and the remainder had concentric, uniform, movement selective or pure direction selective receptive fields. Single cells were also recorded from primary visual cortex of cat to permit a comparison of visual cortical organization in cats and rabbits. Laminar organization of receptive field types was observed in rabbits which was similar in most respects to that described in the cat. Although the major categories of orientation selective cells (simple, complex, hypercomplex) were similar for both cat and rabbit, many differences emerged: (I) tuning of orientation selectivity was narrower in cats than in rabbits; (II) units which preferred oblique orientations were less frequently represented in rabbits than in cats; (III) orientation preferences appeared to be arranged in clusters in rabbit cortex; in rabbits we found no evidence of the columnar organization of orientation selectivity which characterizes cat visual cortex. A comparison of our data with those previously reported for mouse, rat, hamster and opossum visual cortex suggest that mammals in which a significant proportion of visual cortical cells are not orientation selective have in common certain patterns of cortical organization involving a less precise and less specilized representation of stimulus orientation.
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[Biochemical correlates of functional features of visual cortex neurons].
Changes of protein concentration and of different metabolic ferments activity were studied in the rabbit visual cortex after visual deprivation. Two groups of neurons with different sensitivity to visual deprivation were found. The less sensitive are supposed to have an additional source of activation. These findings corroborate the microelectrode data showing that the activity of about 40% of the visual cortex neurons are modulated by a different nonvisual stimulus.
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Role of visual cortex in interocular alignment.
The role which the visual cortex plays in the development of interocular alignment in the cat was examined by removing this structure bilaterally in 4 groups of subjects. These included (1) kittens 10 to 14 days of age, (2) 10- to 14-day-old kittens in which one eyelid was sutured shut at the same time, (3) normally reared adult cats, and (4) cats dark-reared until 4 months of age. If the cortex is removed in young kittens, interocular alignment appears to develop normally until the kittens are 60 to 80 days of age. At this time, an abrupt change in alignment resulting in incyclotorsion of the optic axes is observed. If binocular vision is prevented in kittens with neonatal visual cortex lesions by suturing one eyelid shut, convergent strabismus and/or incyclotorsion are frequently observed. This characteristic incyclotorsion does not develop if similar lesions are made in adult cats; no significant alterations of eye alignment occur in these animals even after postoperative survival times of more than 6 months. Incyclotorsion characterizes dark-reared cats when they are first brought into the light, but this diminishes with time and may even be replaced by excyclotorsion after the animals spend a few weeks in the light. If dark-reared cats are decorticated on being brought into the light, these changes are largely prevented. Such animals remain permanently incyclotorted relative to normal cats. The results indicate that the visual cortex plays an important role in the development of torsional alignment of the eyes.
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[The rat visual cortex. Cytoarchitecture and stereotactic parameters].
In the present investigation the visual cortex of the albino rat was analyzed. On the basis of cytoarchitectonic criterions we can demarced the area striata (field 17) from a medial (field 18) and lateral (field 18a) lying peristriatic area. There are differences in the cortical structure not only between the striate and peristriate areae but also between field 18 and field 18a. In this publication we also present the visual cortex in stereotaxic coordinates on the basis of the stereotaxic atlas of FIFKOVA and MARSALA. Stereotaxic informations belonging the different fields of the visual cortex of the rat are not yet known, but they are very helpful for experimental investigations in this region of the brain.
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