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The nature of the visual discrimination impairment after neonatal or adult ablation of superior colliculi in rats.

The superior colliculi were removed in rats at either one or five days of age or in adulthood. Seven months later they were tested on four successively presented two-choice intensity discriminations. The intensity difference between the discriminanda was reduced across the four problems to encourage choice by comparison. The purpose was to establish whether impoverished scanning is a feature of rats with collicular lesions and whether the age at which the lesion is incurred is important. The number of door-push and approach errors made in reaching criterion were used as measures of performance and the number of head-scans during acquisition was counted. The results provide no evidence that either one- or five-day operated rats exhibit sparing or recovery of the ability to scan discriminanda since all operated animals were impaired. Furthermore, novel retinal projections, present in one-day operated animals, fail to mediate such sparing. Finally, the results did not demonstrate a selective increase in approach errors following collicular lesions and were therefore inconsistent with the view that the impairment is one of visually-guided locomotion. It is concluded that visual discrimination learning is impaired following collicular lesions in circumstances where scanning of discriminanda is required for efficient performance.

Age Factors

Scrapie inoculation of mice: light and electron microscopy of the superior colliculi.

Ultrastructural examination of the superior colliculi of mice intraocularly inoculated with the ME7 strain of scrapie showed vacuolation early in the course of infection. Brains were examined between 85-260 days after monocular inoculation with scrapie. The mean incubation period for the development of clinical disease was 302 days. Vacuolation was seen initially in the contralateral superior colliculus and subsequently in the ipsilateral colliculus. In coded trials light microscopical vacuolation was seen from 218 days but ultrastructural examination showed that sparse vacuoles were inconsistently present in either or both of the ipsilateral and contralateral colliculi from 85 days; frequent vacuoles were seen from 190 days. Scrapie-induced vacuoles were differentiated from vacuoles present in control tissue by the presence of loculation or by a limiting double membrane which showed protrusion or proliferation of the innermost lamella. Vacuolation was seen in neuronal perikarya, myelinated fibres, dendrites and axonal presynaptic terminals. Vacuoles of myelinated fibres were observed within myelin and possibly also in the inner tongue of oligodendroglial cytoplasm. Whorled membrane configurations were also seen. Tubulovesicular particles, 40 nm in diameter, were recognised in two scrapie-infected mice. It is suggested that some scrapie vacuoles arise as a result of incorporation of abnormal membrane into organelles, possibly mitochondria, in neuronal perikarya and neurites and probably also within oligodendroglial cytoplasm and myelin.

Animals

[Facial motor responses to microstimulation of the superior colliculi in the white mouse].

Facial motor responses to microstimulation of different zones of the superior colliculi have been investigated in the albino mice craniotomized under thiopental anaesthesia. Local responses of the mystacial vibrissae, upper lip and eyelids were initiated by microstimulation of the rostral parts of the inner layers of the colliculus superior (high-frequency volleys of 5-7 pulses with a current limit of 35 microA). Sequential changes in the pattern of facial responses were observed within microelectrode traces indicating vertical orientation of facial motor representations in the superior colliculus. Some differences in the localization and pattern of facial responses in the right and left superior colliculi were revealed: 1) vibrissae and lip representations in the right superior colliculus occupy more extensive zone (vertical distribution from 300 to 2,300 microns) as compared to those in the left one (700-2,000 microns); 2) microstimulations of the right superior colliculus produce both uni- and bilateral vibrissal motor responses, whereas stimulation of the left superior colliculus evokes only unilateral responses. The duration of the latent period of the vibrissal and lip motor responses to stimulation of the right superior colliculus varied from 10 to 26 ms (16.1 +/- 2.4 ms; n = 199), to stimulation of the left one-from 10 to 18 ms (mean 14.9 +/- 1.8 ms; n = 55). It is suggested that polysynaptic motor responses to microstimulation of the superior colliculi are realized via the reticular and other premotor nuclei of the brain stem which have direct inputs from the superior colliculus and direct projections to the facial motor nucleus.

Animals

Simultaneous unitary neuronal activity in both superior colliculi and its relation to eye movements in the cat.

The analysis of simultaneous unitary neuronal activity related to eye movement and recorded in both superior colliculi has shown a mirror-functioning image. Increase of the frequency discharge in a collicular unit, was associated with a decrease of the frequency discharge in the contralateral superior colliculus unit. This unitary neuronal reciprocal behaviour was observed each time a horizontal or oblique eye movement was produced. It is possible that this reciprocal functioning between neurones in both superior colliculi could exert an important influence on oculomotor brainstem structures. These results give a better idea of the role played by both superior colliculi in the control of conjugate eye movements.

Animals

Effects of damage to superior colliculi and pre-tectum on movement discrimination in rhesus monkeys.

Eight rhesus monkeys (Macaca mulatta) were trained to detect an instantaneous lateral displacement of a small spot of light. The smallest movement for 79% correct performance was then determined. Severing the splenium of the corpus callosum, which has to be done to reveal the superior colliculi, had no effect on the movement threshold. But when the superior colliculi were damaged in addition, there was an initial substantial impairment in the detection of movement and a smaller but permanent elevation in the threshold. Only if the rostral superior colliculi and pretectum were spared, was there no change in threshold, and there was evidence that the pretectal rather than collicular damage may be more important in relation to movement discrimination. Tests with human observers performing the same task showed that the threshold can be elevated by imprecise fixation and accomodation, suggesting that the consequences of mid-brain damage on movement detection may be wholly or in large part attributed to oculomotor disorders.

Accommodation, Ocular

Retinal ganglion cell survival and neurite regeneration requirements: the change from Müller cell dependence to superior colliculi dependence during development.

A study has been made of the effects of Müller conditioned media or neonatal superior collicular extracts on the survival in dissociate culture of retinal ganglion cells (RGC) from differently aged rats. Embryonic and neonatal RGC were identified either by retrograde horseradish peroxidase (HRP) or Thy-1 antibody labelling techniques. Müller conditioned media supported the survival, over 24 h in culture, of 85% of the RGC plated from 17-day embryos (E17); in contrast, superior collicular extract only maintained 45% of these RGC. With further development there was a decline in the survival enhancing effects of the Müller conditioned media and an increase in the survival due to superior collicular extracts: by postnatal day 12 (P12), the survival of RGC had declined to 20% in the Müller media, but had increased to over 90% in the colliculus extract. This transition in the dependence of RGC from Müller cells to superior colliculi was examined with homogeneous cultures of RGC, obtained using cell sorting techniques. RGC in these cultures showed the same transition in dependence from Müller cells to superior colliculi, indicating that the survival-enhancing effects were not mediated by the other intrinsic cells of the retina. The results suggest that the survival of RGC is at first dependent on the intrinsic glia of the retina and only later, as development proceeds, on the targets of the RGC in the superior colliculus. The normal RGC death period in the rat extends from near birth to about 6 days postnatal. The question then arises as to the timing of trophic support from Müller glia and from neurones in the tectum in relation to the duration of the normal RGC death period. This has been examined in the present work.

Animals

[Function of the superior colliculi of the corpora quadrigemina during creation of a local focus of increased excitability in the mesencephalic reticular formation and sensomotor cortex].

In chronic experiments on waking rabbits, the foci of heightened excitability in the sensorimotor cortex and mesencephalic reticular formation affected in a similar way the background neuronal activity in the superior colliculi and that evoked by light stimuli. The effect was manifested in elimination of inhibitory pauses in the neuronal response to light stimulus and in a general increase of discharge frequency. Similarity of the cortical and reticular influences is due to their possible mediation by the same collicular interneurones participating in inhibitory pauses formation in the process of backward inhibition. Increased neuronal activity in the superior colliculi under the action of local foci in the sensorimotor cortex and mesencephalic reticular formation correlated with appearance of forelimb motor reaction to isolated light stimulus testifying to a formation of a functional connection between the visual and motor analyzers. Possible role of the superior colliculi in this process and their participation in the formation of a visually controlled reaction is discussed.

Animals

The role of frontal eye-fields and superior colliculi in visual search and non-visual search in rhesus monkeys.

Rhesus monkeys were tested on a visual search task in which they had to find and retrieve a peanut from a display of visually similar but inedible objects. The speed with which they did so was measured. Animals in which the superior colliculi or frontal eye-fields had been removed took longer to find the peanut than two operated control groups. Animals with collicular lesions had longer latencies than those with frontal eye-fields removed. These two groups were also tested on a second task, non-visual search, in which a peanut was concealed in each of 25 identical holes. The animals' task was to retrieve all 25 peanuts as quickly as possible. The group with frontal eye-fields removed made significantly more return errors, i.e. returning to a hole already sampled, than the control group but, in contrast to the first task, the animals with collicular lesions were not impaired. The results are related to the physiological properties of frontal eye-fields and superior colliculi and to the effects of frontal cortical brain damage in man. It is suggested that the frontal eye-fields are concerned with internally organized, i.e. voluntary, eye scanning whereas the superior colliculi are concerned with the detection and location of targets which are then fixated involuntarily.

Animals

Inhibition of labyrinthine nystagmus by visual fixation: effects of ablation of visual cortex and superior colliculi.

A study was conducted to destroy two specific areas of the cat's visual system in order to determine if these lesions would affect the visual inhibition of calorically-induced vestibular nystagmus. The occipital visual cortex was removed in eight cats and the superior colliculi were removed bilaterally in nine cats. Postoperative vestibular testing revealed no significant change in the electronystagmography tracings and response to visual fixation. These findings suggest that, in cats, the visual inhibition of labyrinthine nystagmus is not dependent upon the integrity of the visual cortex or superior colliculi. The hypothesis is brought forward that the visual inhibition of the vestibular nystagmus is merely a reflex of the brain stem to light stimulus, mediated via the cerebellum.

Animals

Abnormalities in grooming behavior and tryptophan hydroxylase activity in the superior colliculi in cats with pontile and frontal neocortical lesions.

Cats with pontile or frontal neocortical lesions display a dissociation of the appetitive and consummatory components of grooming behavior when their body surface is tactually stimulated, an abnormal behavior that waxes and wanes with the seasons of the year. Tryptophan hydroxylase activity is significantly decreased in the superior colliculi of cats with pontile lesions and of cats with frontal neocortical lesions. The results suggest that the change in tryptophan hydroxylase activity is mediated neuronally and is a transneuronal effect on the serotonergic input to the superior colliculi. Pharmacological manipulations of the serotonergic system in normal cats failed to induce the abnormal behavior, indicating that other factors are involved in the genesis of the abnormal behavior.

Animals

The effect of ablation of frontal eye-fields and superior colliculi on visual stability and movement discrimination in rhesus monkeys.

The neurological basis of the maintenance of a stable visual scene by means of a corollary discharge mechanism was investigated. Monkeys were trained to detect and respond to sudden rapid movement of a small spot of light in an otherwise totally dark environment. There was no evidence that after removal of the frontal eye-fields, superior colliculi, or caudal superior temporal sulcus the animals confused real movement of the target with retinal image movement caused by changing the position of head and eyes. The result was confirmed by an examination of the ipsiversive turning that follows unilateral frontal eye-field or collicular ablation. If the turning is a compensation for apparent movement of the visual world when the eyes are moved it should not be present in total darkness. It was still present. The thresholds for the smallest detectable instantaneous displacement of the target were also measured. The threshold was impaired by bilateral superior colliculus lesions but not by removal of the frontal eye-fields or cortex of the caudal part of the superior temporal sulcus.

Animals

[Directional selectivity of neurons of the superior colliculi of the lamina quadrigemina in visually deprived rabbits].

The reactivity of neurones in the superior colliculi to the movement of the optic stimulus was studied in monocularly and binocularly deprived rabbits (MD and BD). Deprivation was effected by suturing the eyelids before the eyes opened. The neurones were classified into three groups by the degree of their directional selectivity (absolute, relative and zero). The effect of a prolonged (ten weeks or more) early MD consisted in the absence of collicular units with absolute selectivity. Proceeding from the existence of such units in the MD type rabbits in the third week of their life, it is suggested that prolongation of the MD time leads to their functional reduction. As the recorded characteristics of unit responses do not change in BD rabbits, it is assumed that the MD effect is not linked to limiting the function of vision and is just a consequence of a prolonged asymmetry of optic stimulation. The deprivation effect at the collicular level in different vertebrates is discussed.

Animals

Evidence that superior colliculi are involved in the control of amygdala-kindled seizures.

The effects of bilateral high radiofrequency lesions of superior colliculus (SC) were studied on amygdala kindling. The results demonstrate that a selective destruction of the SC only slightly facilitated the development of kindling. However, the most remarkable effect was the increase of afterdischarge and motor seizure duration observed when SC-lesioned animals reached the generalized seizures. These data confirm that the superior colliculi participate in controlling the generalization of kindled seizures.

Amygdala

Single-cell responses associated with rhythmic slow-wave potentials in rat superior colliculi.

The present study was designed to investigate the features of rhythmic slow-wave potentials in the superior colliculi of rats and to study the relationship between these potentials and the activity of single colliculi neurons. In contrast to studies in other portions of the visual system, rhythmic slow-wave potentials in the colliculi fell within a single limited range of frequencies from 9 to 26 cycl/sec. Single-cell recordings revealed that numerous collicular neurons were also responding rhythmically, with discharge frequencies generally coinciding with those established for collicular slow-wave potentials. In some animals slow-wave and single-cell potentials were recorded simultaneously through adjacent electrodes, and in all instances the frequency of discharge of single neurons coincided with the frequency of oscillation of the slow-wave potentials. These data suggest that rhythmic postsynaptic potentials in individual collicular neurons may be the mechanism by which rhythmic slow-wave potentials are generated.

Animals

[Interaction of recovery cycles of evoked potentials in the cerebral cortex of the cat in response to stimulation of the superior colliculi and pulvinar].

Similar character of recovery cycles of evoked potentials in the visual cortex to electric stimulation of the superior colliculi (SC) and pulvinar was found in chronic experiments on alert cats irrespective of stimuli presentation order. In the association cortex preceding SC stimulation facilitated the response to test stimulation of pulvinar almost at all delays between the stimuli. If the pulvinar stimulation was applied as a conditioned stimulus, then the response to SC stimulation under intervals of 20-200 ms was depressed. The obtained data point to equivalence of the inputs from SC and pulvinar to the visual cortex, to different informational value of inputs from SC to the association and visual cortex, and to mutual function dependence of the inputs from SC and pulvinar to the association cortex.

Animals

Single retinal ganglion cells sending axon collaterals to the bilateral superior colliculi: a fluorescent retrograde double-labeling study in the Japanese monkey (Macaca fuscata).

Single retinal ganglion cells projecting bilaterally to the superior colliculi (SC) by way of axon collaterals were revealed in the Japanese monkey (Macaca fuscata). After injecting Fast blue into the SC on one side and Diamidino yellow into the SC on the opposite side, some retinal ganglion cells were double-labeled with both tracers. Most of them were large cells (more than 25 microns in diameter), and were localized in a narrow strip around the vertical meridian of the retina on each side. This retinal area roughly corresponds to the reported strip of nasotemporal overlap, where both crossed and uncrossed retinofugal projections arise.

Amidines

Increase by visual stimuli in turnover of 5-hydroxytryptamine in the superior colliculi of the rabbit.

1. Irregular light falshes were played on to one eye of dark adapted rabbits for periods of 20-80 min. The concentration of 5-hydroxyindol-3-ylacetic acid (5-HIAA) and of 5-hydroxytryptamine (5-HT) were estimated in left and right superior colliculi, thalami and hippocampi. 2. In rabbits exposed to such visual stimuli for 30-60 min, there was an increase in the 5-HIAA content of the colliculus contralateral to the stimulated retina which aberaged 17% (P = 0-02), but no rise was seen if the exposure was shortened to 20 or prolonged to 80 min. At no time was there a difference in 5-HIAA content between right and left thalamus or right and left hippocampus. 3. Stationary or strictly repetitive visual stimuli produced no difference between the 5-HIAA content of left and right superior colliculus. 4. No difference in 5-HT concentration between the two colliculi was found after any form of visual stimulation, nor did any changes occur in the other parts of the brain which were examined. 5. Irregular, prolonged visual stimualtion thus appears to activate tryptaminergic neurones terminating in the colliculi. The possibility is discussed that the 5-HT released at this site might act as a brake to neuronal activity under conditions when habituation to the stimuli is not yet complete.

Animals

[Oscillating potentials of the retina and superior colliculi and correlations with neuronal activity].

The oscillatory potentials of retina and superior colliculus induced with photostimuli were studied in unanesthetized rabbits. Increasing intensity of light flashes induces a better correlation of the responses in these structures. A common genesis of the oscillatory processes and neuronal activity was revealed. The peculiarities of formation of the short-latency responses in the superior colliculus and the specificity of generation of the responses of photostimuli of extreme brightness, are discussed.

Animals