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Noradrenergic sprouting and beta-adrenergic receptor binding in the lateral geniculate nucleus of rats after unilateral visual cortex ablation.

One and two weeks after unilateral visual cortex (VC) ablation beta-adrenergic receptor binding is increased in the lateral geniculate nucleus (LGN) of both sides. 6 month later beta-receptor binding in the LGN is decreased again and no differences to untreated control animals are detectable. Using the glyoxylic acid fluorescence method for the visualization of amines a transient increase in the density of noradrenergic fibers in the dorsal part of the ipsilateral LGN can be demonstrated with a maximum 2 weeks after VC ablation. With longer survival time noradrenergic fibre density in the ipsilateral dorsal LGN (LGNd) decreases again and one year after the operation only few fluorescent fibres can be observed in the lateral part of the LGN compared to untreated control animals. Histologically an increased gliosis in the ipsilateral LGNd develops following removal of the visual cortex. In addition, degenerative changes in nerve fibers and terminals as well as neuronal degenerative changes are present and are most pronounced in the medial part of the ipsilateral dorsal LGN. Electron microscopically degenerating terminals in the dorsal part of the ipsilateral LGN can be identified as cortical afferents. Using potassium permanganate fixation typical noradrenergic axons with small dense core vesicles can be demonstrated in the LGN. In the denervated LGNd (i.e. the ipsilateral LGNd after visual cortex ablation) axon terminals with dense core vesicles appear exhibiting the structural peculiarities of growth cones seen during ontogenesis. They could be regarded as ultrastructural equivalent of the newly formed noradrenergic sprouts.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals

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.

Animals

Arrangement of ocular dominance columns in human visual cortex.

The arrangement of the ocular dominance columns in the human primary visual cortex was studied by examining cytochrome oxidase activity in autopsy specimens of occipital lobes obtained from two patients who became blind in one eye before death. By artificially flattening the cortex before processing, it was possible to reconstruct the pattern formed by the ocular dominance columns throughout most of the primary visual cortex. The columns form a mosaic of irregular parallel stripes about 500 microns to 1000 microns wide (right eye column plus left eye column measures 1 to 2 mm), oriented at right angles to the boundary of the primary visual cortex. The columns are wider near the boundary of the primary visual cortex and within the representation of the peripheral visual field, the ocular dominance columns of the ipsilateral eye become fragmented until they disappear altogether at the border of the monocular crescent representation. The arrangement of ocular dominance columns in the human visual cortex is very similar to the pattern reported in the macaque monkey, although the columns in humans are wider.

Aged

Visual cortex: a cat's-eye view of the visual system.

Optical imaging data show that the cat's visual cortex contains patches of cells that respond to low spatial and high temporal frequencies; outside the patches, cells respond to high spatial and low temporal frequencies. The results suggest a possible anatomical substrate for psychophysically defined spatial frequency channels.

Animals

[Image processing in the primary visual cortex].

INTRODUCTION: Area 17 or the primary visual area forms the first link in the chain of cerebral analysis of a visual image. The neurones forming the primary visual cortex are characterized by the extreme precision of their connections, functional specialization and hierarchic organization. The spatial precision of the connections within the system for vision permit retinotopic representation in the visual cortex, so that each point of the retina is projected into a specific area of the cortex. The cortical neurones which analyze the characteristics of the image situated in a precise zone of the visual field are themselves organized into a basic functional unit known as a hypercolumn. Within each hypercolumn there are various columnar cell systems with receptive fields having similar characteristics. Thus, each hypercolumn is made up of multiple orientation columns, two ocular dominance columns and 'blob' regions. All these systems permit the analysis of different aspects of the image. The neurones belonging to the orientation columns are sensitive to the orientation, spatial frequency and movement of a visual stimulus; those of the 'blob' regions to colour, and the binocular neurones of the ocular dominance columns to depth. Within each column, the hierarchical pattern of neurone interconnections determines the successive appearance of cells with receptive fields having new properties.

Geniculate Bodies

Spatio-temporal frequency domains and their relation to cytochrome oxidase staining in cat visual cortex.

Spatial and temporal frequencies are important attributes of the visual scene. It is a long-standing question whether these attributes are represented in a spatially organized way in cat primary visual cortex. Using optical imaging of intrinsic signals, we show here that grating stimuli of different spatial frequencies drifting at various speeds produce distinct activity patterns. Rather than observing a map of continuously changing spatial frequency preference across the cortical surface, we found only two distinct sets of domains, one preferring low spatial frequency and high speed, and the other high spatial frequency and low speed. We compared the arrangement of these spatio-temporal frequency domains with the cytochrome oxidase staining pattern, which, based on work in primate striate cortex, is thought to reflect the partition of the visual cortex into different processing streams. We found that the cytochrome oxidase blobs in cat striate cortex coincide with domains engaged in the processing of low spatial and high temporal frequency contents of the visual scene. Together with other recent results, our data suggest that spatiotemporal frequency domains are a manifestation of parallel streams in cat visual cortex, with distinct patterns of thalamic inputs and extrastriate projections.

Animals

The development of basal forebrain projections to the rat visual cortex.

The development of the basal forebrain projections to the visual cortex of the rat were studied using retrograde tracer techniques. Injections of wheat germ agglutinin-horseradish peroxidase placed in the visual cortex of newborn animals resulted in labelling of neurons throughout the basal forebrain nuclei. Although at this time the overall distribution of retrogradely labelled cells within the basal forebrain appeared similar to that seen in the adult, cells were smaller and weakly stained. It was only at the end of the second postnatal week that the somata of stained neurons showed sizes and staining intensity comparable to the adult. This precedes or coincides with the reported significant increases in cortical and basal forebrain ChAT activity and the first detection of ChAT-labelled fibres in this cortical area. These data suggest an important developmental point around the end of the second postnatal week that may correspond to the time when a significant number of cholinergic axons first appear within the visual cortex. They also suggest that the cholinergic projections to the visual cortex develop late in comparison with the thalamic and other subcortical afferents in this cortical area.

Aging

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.

Animals

Isochronic transplantation of neonatal grafts in the visual cortex of cats: responsiveness, ocular dominance and specificity of cortical cells to visual stimulation.

The visual cortex of adult cats was studied physiologically following neonatal isochronic transplantation of grafts from areas 17,18, which were placed homotopically, in order to reveal their functional integration and thus possible repairing of damaged cortical neuronal circuits. Three homograft cats, in which transplantation was carried out between siblings (228 cortical cells) were compared to 4 animals receiving reimplanted autografts of the equivalent size (131 cells) as well as 3 animals with analogous sectioning of the visual cortex (162 cells) (pseudograft controls). The location of the boundaries between the transplant region and the host were determined using the Nissl's method for staining histological cross sections. Extracellular unit recording revealed typical waveform of the action potentials in the transplanted region and in the surrounding host tissue of all groups of cats. Visual responsiveness in the homograft cats was 17.5% in the transplanted region and 80.4% in the unoperated hemisphere; the corresponding results were 40.3% for the transplanted region and 82.2% for the unoperated hemisphere in the autografts and 23.1% and 73.4% in the pseudografts. The specificity of the cells to visual stimulation as expressed by their orientation and direction specificity, indicated preservation of these properties in the transplanted cats. While all responsive cells in the transplanted region of the homografts were orientation specific, their proportion was 60% in the autografts and 55.5% in the analogous region in the pseudograft controls. As to the direction specific cells, their performance in the grafted region of the grafted cats was even much higher than that of the pseudograft controls. The ocular dominance distribution of the cells showed preservation of binocularity in the transplanted region (90.0% binocular cells) of the homografts; it was however smaller in the equivalent region of the autografts (65.0%) and remarkably reduced (20.0%) in the pseudografts. It was concluded that despite the deafferentation induced during the transplantation procedure, a remarkable visual responsiveness was found in the transplanted region, indicating postoperative recovery. However, the cells there were mainly affected in their activity and less in their specificity to visual stimulation.

Animals

Effects of neonatal monocular and binocular enucleation on transient acetylcholinesterase activity in developing rat visual cortex.

Geniculo-recipient layers of visual cortical area 17 in the laboratory rat display a transient pattern of acetylcholinesterase (AChE) activity during the second and third postnatal weeks of life. The appearance of the AChE histochemical reaction product and its distribution in thalamic recipient layers of cortical area 17 suggest that this transient AChE serves as a marker for the region of geniculocortical axon terminals. In the present study, infant rats were enucleated monocularly or binocularly on the day of birth. Animals were sacrificed at postnatal days 5-21. Frozen sections cut in either the transverse plane or parallel to the pial surface were processed for AChE histochemistry. Neonatal monocular enucleation resulted in a marked reduction of transient AChE activity in thalamic recipient layers of the medial part of area 17 contralateral to the enucleated orbit, i.e., the monocular segment of area 17. No loss of AChE was observed in area 17 ipsilateral to the enucleation. Pigmented and albino strains of rats did not differ significantly in the extent to which monocular enucleation reduced the transient AChE in contralateral visual cortex. Neonatal binocular enucleation resulted in an almost complete loss of AChE histochemical staining in thalamic recipient layers throughout cortical area 17, without loss of AChE in other cortical regions. These data support the hypothesis that transient AChE serves as a marker for the region of geniculocortical axon terminals, and also demonstrate that the transient expression of AChE in visual cortex depends upon normal innervation or activity of the geniculocortical neurons.

Acetylcholinesterase

The monocular and binocular subfields of the rat's primary visual cortex: a quantitative morphological approach.

Primary visual cortex in the rat was studied by a variety of methods: transsynaptic transport of labelled amino acids, 2-deoxyglucose, and staining for perikarya, myelin, and acetylcholinesterase. The analysis was aided by a computer-controlled television image analyzer. The results obtained with different methods agree with one another in describing the position and extent of the entire primary visual cortex as well as its monocular (medial) and binocular (lateral) subareas.

Acetylcholinesterase

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

[Effect of stimulation of interanalyzer association areas of rabbit cortex on evoked potentials and neuronal responses of the visual cortex].

The study was concerned with functional influences of associative cortical areas on EPs and neuronal responses to flashes in the visual projection zone of a rabbit. Stimulation of the anterior associative cortical parts leads to a prolongation of the inhibitory pause and to a decrease of late discharges in responses of visual cortical neurones which is manifested in a significant reduction of the secondary negative-positive EP complex. Stimulation of the posterior associative area causes a prolongation and a deepening of the inhibitory pause in the neuronal activity and an increase of late discharges in the neuronal response in the visual cortex with a corresponding significant enhancement of the secondary negative-positive EP complex. Different mechanisms of functional effects of the anterior and posterior interanalyser brain parts on the visual projection area are suggested.

Animals

Segregation of pathways leading from area V2 to areas V4 and V5 of macaque monkey visual cortex.

V5 and V4 are areas of macaque monkey prestriate visual cortex that are specialized for involvement in different aspects of visual perception, namely motion for V5 (refs 1-4) and colour vision, with other possible functions, for V4 (refs 2, 5-9). Thus, it is unlikely that they should be fed the same information for further processing, yet both receive a strong input from patches of the upper layers of V2 (refs 10, 11), the area immediately adjoining the primary visual cortex, V1. V2, however, seems to comprise functionally distinct subregions, which can be revealed by staining the tissue for the mitochondrial enzyme cytochrome oxidase. Here we report that V4 and V5 are connected with separate cytochrome oxidase-defined subregions of V2, suggesting that cortical pathways dealing with motion and colour perception are segregated in their passage through V2, and reinforcing evidence for functional specialization in the visual cortex.

Animals

Vasoactive intestinal polypeptide immunoreactive neurons in the primary visual cortex of the cat.

When cat visual cortex (area 17) is reacted with an antibody to vasoactive intestinal polypeptide (VIP) a variety of neuronal types is labelled. Many of the labelled neurons are bipolar in form and are most common in layers II and III, although significant numbers of bipolar neurons are also encountered in layer V. Multipolar cells are also labelled. These are most frequent in layer IV and have a variety of shapes. In layer I, the labelled cells are of three varieties, i.e. horizontal bipolar cells, horizontal bitufted cells and multipolar neurons, while in layer VI the few VIP-positive neurons are horizontal bipolar cells. This suggests that all of the VIP-labelled neurons in cat area 17 are non-pyramidal in form, and this has been confirmed by electron microscopy. In these preparations, axon terminals are also labelled and under the light microscope it can be seen that these terminals occur both within the neuropil and around the cell bodies of some neurons, particularly neurons in layers II and III. Electron microscopy has shown that all of the labelled axon terminals form symmetric synapses and that those in the neuropil synapse with the shafts of smooth dendrites. These axodendritic synapses account for about 90% of the synapses formed by the labelled axon terminals. The remainder of the labelled axon terminals synapse with the cell bodies of pyramidal neurons. Parallels are drawn between these results and those previously obtained by examining those neuronal elements labelled with VIP antibodies in rat visual cortex.

Animals

[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.

Acetylcholinesterase

Development of the visual cortex in a wallaby--phylogenetic implications.

The visual cortex of one of the smallest macropod marsupials, a wallaby, the quokka Setonix brachyurus, was examined at maturity and during development from postnatal day 1 to 150 in Nissl-stained or Golgi-stained sections. Injections of horseradish peroxidase into the primary visual centres in adults identified cortical neurons projecting to these regions. The pattern of cell generation was determined by tritiated thymidine/autoradiography. The adult visual cortex was composed of the usual six layers of cells with dendritic morphologies similar to those seen in eutherian (placental) mammals. The margin of Layer 6 overlapped slightly with the white matter, and pyramidal cells projecting to subcortical regions were located at all depths of Layer 6. Cortical development was similar to that of eutherian mammals, with the formation of transient cellular layers below the developing cortical plate. Autoradiographic studies confirmed that the cortical plate was composed of two layers: a compact zone of densely packed cells located pial to a striated layer containing more loosely packed cells. Postnatal cell generation of Layers 6-2 took place from postnatal day 3 to day 85, with Layer 4 neurons, the main target of incoming visual axons, being generated around postnatal day 40. Cells located at maturity in the white matter, mostly of glial morphology, were generated from postnatal day 40. A subplate zone containing early differentiating cells was not evident, and postnatal 3H-thymidine labelling did not identify a population of early generated neurons below the cortical plate. In the tammar wallaby Macropus eugenii, the majority of target neurons for incoming geniculate axons are generated about 20 days after the first axons enter directly into the cortical plate without a waiting period in a subplate. Geniculate axons in the wallabies have a relatively longer 'wait' for their target neurons than do those in cats or monkeys. Therefore, if geniculate axons in marsupials make temporary synapses, while they wait for their target neurons to be generated, their temporary connections must, unlike those in cats or monkeys, be with cells already in the cortical plate.

Aging

Morphology of corticotectal cells in the primary visual cortex of hooded rats.

In primary visual cortex of hooded rats, pyramidal cells in layer V may be classified as long, medium, or short, on the basis of the layer in which the apical dendrite terminates. The present study determines which of these types of pyramidal cells project to the superior colliculus. Two different strategies were used to label corticotectal cells with horseradish peroxidase (HRP). In the first set of experiments, a large number of corticotectal cells were labeled by retrograde transport following injection of HRP into the superior colliculus. In the second set of experiments, single unit recording was used to identify corticotectal cells physiologically by antidromic activation from the superior colliculus. These cells were then impaled and labeled by intracellular iontophoresis of HRP. The results from both techniques suggest that only long pyramidal cells send an axon to the superior colliculus. These cells are distinguished by an apical dendrite that extends into layer I. We conclude that in hooded rats corticotectal cells in primary visual cortex are the long pyramids in layer V.

Action Potentials