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Fibre divergence in the distal optic radiation: possible basis of functional plasticity in adult primate visual cortex.

The precision of retinotopy in primate visual cortex is commonly thought to result from highly ordered arrangement of fibres in the visual pathways. However, rigid point-to-point representation is hardly compatible with findings of a substantial reorganization of visual cortical maps after peripheral and central lesions. Such observations could be accounted for by divergence in the optic radiation. To explore the hypothesis of fibre divergence, we made small knife cuts in the distal optic radiation of macaca fascicularis. After subsequent axonal tracing by injecting WGA-HRP into lateral geniculate nucleus, we studied the course of distal fibres in white matter. The amount of divergence was assessed by measuring, relative to the prevailing fibre course, length and orientation of labelled fibres between lesion and entry into cortex. Lesion sizes between 1 mm to 3 mm did not result in any detectable diminution of terminal labelling in layer IVC of striate cortex. Individual labelled fibres were found to diverge symmetrically from both sides into the gap distal to the lesion. Divergence starts at a distance of about 3 mm before cortex. At the white matter boundary, less than 10% of all fibres still retain the original direction, with the remaining fibres taking any other orientation without preference. We estimate that this corresponds to a divergence of visual afferents encompassing about 6-10 mm of cortical distance, if intracortical arborization of terminal fibres is taken into account. Possible consequences for functional plasticity in the adult primate visual cortex are discussed.

Animals↗

[Functional magnetic resonance tomography of the visual cortex].

Functional magnetic resonance imaging (MRI) allows to directly visualize regional activity of the visual cortex during stimulation. The value of the method to evaluate physiologic and pathologic conditions is elaborated on in comparison with positron emission tomography (PET). Aspects as sequence selection and stimulus paradigms are discussed. Subtle activity in the primary cortex (V1), processing areas as MT/V5 and subcortical areas are visualized by MRI. Cooperation of the subjects is a essential factor. Despite promising initial results in clinical trials the focus of research in clinical trials the focus of research in the visual cortex yet remains in the preclinical field.

Animals↗

Depth is encoded in the visual cortex by a specialized receptive field structure.

Binocular neurons in the visual cortex are thought to perform the first stage of processing for the fine stereoscopic depth discrimination exhibited by animals with frontally located eyes. Because lateral separation of the eyes gives a slightly different view to each eye, there are small variations in position (disparities), mainly along the horizontal dimension, between corresponding features in the two retinal images. The visual system uses these disparities to gauge depth. We studied neurons in the cat's visual cortex to determine whether the visual system uses the anisotropy in the range of horizontal and vertical disparities. We report here that there is a corresponding anisotropy in the cortical representation of binocular information: receptive-field profiles for left and right eyes are matched for cells that are tuned to horizontal orientations of image contours. For neurons tuned to vertical orientations, left and right receptive fields are predominantly dissimilar. Therefore, a major modification is required of the conventional notion of disparity processing. The modified scheme allows a unified encoding of monocular form and binocular disparity information.

Action Potentials↗

Transient patterns of serotonergic innervation in the rat visual cortex: normal development and effects of neonatal enucleation.

The transient aggregation of serotonin (5-HT)-containing fibers in the early development of rat visual cortex was examined immunohistochemically. The aggregation of 5-HT immunoreactive (IR) fibers consisted of three stages which were classified according to the course of time and degree of space occupied. The primary aggregation appeared in the subplate and moved upward along the development of the cortex. The aggregation proceeded to the secondary stage in presumptive layer IV. The fibers extended in a column-like structure following the secondary aggregation and formed the tertiary aggregation. The upper edge of the tertiary aggregation formed a lattice-like pattern in layer I and its structure was recognized to be similar to the structure of a 'blob' which characterizes the primary visual cortex in monkey. This transient aggregation of 5-HT-IR fibers began in the subplate of the anterior visual cortex on postnatal day 2 (PND 2) and progressed towards the posterior. On PND 11, the secondary and tertiary aggregations were completed in the entire region. No further aggregation of 5-HT-IR fibers was observed on PND 15. The anterior-to-posterior axis in the aggregation process corresponds to the direction of differentiation in the layer structure of cortex. In order to investigate the relationship between the transient aggregation of 5-HT-IR fibers and the development of the visual pathway, the secondary and tertiary aggregation on PND 11 were observed after postnatal monocular or binocular enucleation. Enucleation of eye balls did not affect either the area occupied by the 5-HT-IR fibers in the secondary aggregation or the number of column structures in the tertiary aggregation. However, the contralateral and ipsilateral cortices of monocularly enucleated cases were irregularly shaped in the secondary aggregation. The distribution of 5-HT-IR fiber terminals in the binocular area (Oc1B) increased in density on the contralateral side in the monocular enucleation, while that of both sides in the binocular enucleation was of non-homogeneous density and were shaped irregularly. The above results suggest that the transient aggregation of 5-HT-IR fibers observed in the early stage of development of visual cortex is regulated primarily by the intrinsic factors, and that extrinsic factors, such as visual pathway input, affect the aggregation within the boundary of such intrinsic factors. That is, the visual pathway input and the input balance from both eyes affect the distribution density of 5-HT-IR fibers and the shape of the visual cortex, respectively.

Animals↗

Primary culture of identified neurons from the visual cortex of postnatal rats.

We have examined the properties of neurons from the visual cortex of postnatal Long Evans rats in dissociated cell culture. Visual cortex from rat pups 1-15 d old was subjected to enzymatic and mechanical dissociation to yield a suspension of single cells. Neurons plated onto collagen or a feeder layer of astrocytes rapidly extended processes and survived for 4-10 weeks. Antisera to glutamic acid decarboxylase, choline acetyltransferase, and vasoactive intestinal polypeptide stained 22 +/- 2, 2.3 +/- 0.3, and 2.4 +/- 0.2% of all neurons, respectively, suggesting that different neuronal classes survived roughly in proportion to their number in vivo. In order to study a particular identified class of cortical neurons, we prelabeled cells in vivo by retrograde transport of a fluorescent tracer. Neurons in layer V of visual cortex that project to the superior colliculus were labeled after injecting fluorescent latex microspheres into the colliculus. Retrogradely labeled neurons were readily identified immediately after dissociation and throughout the period in vitro. After 2 weeks in culture, labeled cells exhibited many ultrastructural features characteristic of pyramidal neurons in vivo. Intracellular recording techniques were used to evaluate the response properties of labeled layer V neurons, as well as other, unlabeled neurons, to excitatory amino acid agonists and antagonists. Glutamate and aspartate--as well as the synthetic agonists N-methyl-D-aspartate (NMDA), kainate, and quisqualate--excited every cortical neuron tested. The antagonist 2-amino-5-phosphonovaleric acid had no effect on responses to quisqualate and kainate but completely blocked depolarizations due to NMDA and aspartate and reduced depolarizations elicited by low concentrations of glutamate. Kynurenic acid, piperidine dicarboxylic acid, and gamma-D-glutamylglycine antagonized responses to all 5 of the agonists. These results provide evidence that corticocollicular neurons in culture express both NMDA-type and non-NMDA receptors for excitatory amino acids.

Animals↗

Activity-dependent regulation of NMDAR1 immunoreactivity in the developing visual cortex.

NMDA receptors have been implicated in activity-dependent synaptic plasticity in the developing visual cortex. We examined the distribution of immunocytochemically detectable NMDAR1 in visual cortex of cats and ferrets from late embryonic ages to adulthood. Cortical neurons are initially highly immunostained. This level declines gradually over development, with the notable exception of cortical layers 2/3, where levels of NMDAR1 immunostaining remain high into adulthood. Within layer 4, the decline in NMDAR1 immunostaining to adult levels coincides with the completion of ocular dominance column formation and the end of the critical period for layer 4. To determine whether NMDAR1 immunoreactivity is regulated by retinal activity, animals were dark-reared or retinal activity was completely blocked in one eye with tetrodotoxin (TTX). Dark-rearing does not cause detectable changes in NMDAR1 immunoreactivity. However, 2 weeks of monocular TTX administration decreases NMDAR1 immunoreactivity in layer 4 of the columns of the blocked eye. Thus, high levels of NMDAR1 immunostaining within the visual cortex are temporally correlated with ocular dominance column formation and developmental plasticity; the persistence of staining in layers 2/3 also correlates with the physiological plasticity present in these layers in the adult. In addition, visual experience is not required for the developmental changes in the laminar pattern of NMDAR1 levels, but the presence of high levels of NMDAR1 in layer 4 during the critical period does require retinal activity. These observations are consistent with a central role for NMDA receptors in promoting and ultimately limiting synaptic rearrangements in the developing neocortex.

Age Factors↗

Parvalbumin immunoreactivity: a reliable marker for the effects of monocular deprivation in the rat visual cortex.

In mammals, monocular deprivation performed during the early stages of postnatal development (critical period) dramatically affects the functional organization of the visual cortex. Since the early work of Hubel and Wiesel, the effects of monocular deprivation are accounted for by the fibers driven by the two eyes competing for the control of cortical territories. In cat and monkey striking structural changes accompany the functional effects of monocular deprivation. Also, in the rat, monocular deprivation causes functional alteration at the level of visual cortex; no structural correlates of these effects, however, have so far been described. Parvalbumin is a calcium binding protein that in the neocortex colocalizes with a subpopulation of GABAergic neurons. Here we report that in the rat monocular deprivation results in a dramatic reduction of parvalbumin-like immunoreactivity in the visual cortex contralateral to the deprived eye. This effect is due to competitive phenomena and not to visual deprivation itself, it is restricted to the binocular portion of the visual cortex and neither binocular deprivation, nor dark rearing can induce it. We conclude that parvalbumin-like immunoreactivity is a useful immunohistochemical marker for the effects of monocular deprivation in the rat visual cortex.

Animals↗

Glutamate-like immunoreactivity in the cat superior colliculus and visual cortex: further evidence that glutamate is the neurotransmitter of the corticocollicular pathway.

Biochemical studies provide evidence that the pathway from visual cortex to the superior colliculus (SC) utilizes glutamate as a neurotransmitter. In the present study, we have used immunocytochemistry, visual cortex lesions, and retrograde tracing to show directly by anatomical methods that glutamate or a closely related analog is contained in corticocollicular neurons and terminals. A monoclonal antibody directed against gamma-L-glutamyl-L-glutamate (gamma glu glu) was used to localize glutamate-like immunoreactivity in both the superior colliculus (SC) and visual cortex (VC). Unilateral lesions of areas 17-18 were made in four cats to determine if gamma glu glu labeling was reduced in SC by this lesion. WGA-HRP was injected into the SC of 10 additional cats in order to determine if corticocollicular neurons were also labeled by the gamma glu glu antibody. A distinctive dense band of gamma glu glu immunoreactivity was found within the deep superficial gray and upper optic layers of SC where many corticotectal axons are known to terminate. Both fibers and cells were labeled within the band. Immunoreactivity was also found in cells and fibers throughout the deep layers of SC. Measures of total immunoreactivity (i.e. optical density) in the dense band were made in sections from the SC both ipsilateral to and contralateral to the lesions of areas 17-18. A consistent reduction in optical density was found in both the neuropil and in cells within the dense band of the SC ipsilateral to the lesion. A large percentage of all corticocollicular neurons that were retrogradely labeled by WGA-HRP also contained gamma glu glu. These results provide further evidence that the corticocollicular pathway in mammals is glutamatergic. The results also suggest that visual cortex ablation alters synthesis or storage of glutamate within postsynaptic SC neurons, presumably as a result of partial deafferentation.

Animals↗

Neural discharge coupled to saccade offset in the cat visual cortex.

The increase in neural activity in cat visual cortex associated with eye movements has been thought to reflect a replica of the motor command signal. We examined the timing of the saccade-related increase in neural activity in cat areas 17 and 18 in relation to saccade onset and offset. The increase in activity was temporally coupled to saccade offset rather than onset both for visually guided saccades and for spontaneous saccades in the dark. Overall, it occurred 63 ms after saccade offset, and the peak was higher and sharper for data aligned at saccade offset than for onset. These results are inconsistent with the idea that saccade-related activity in cat visual cortex reflects a copy of the motor command signal.

Animals↗

Interactions between callosal, thalamic and associational projections to the visual cortex of the developing rat.

The patterns of callosal interconnections between the visual cortices of rats display considerable plasticity in response to various neonatal manipulations. In the present study, many neurones in the principal visual thalamic relay nuclei, the dorsal lateral geniculate nucleus (DLG) and to a lesser extent those in the lateral posterior nucleus (LP) were destroyed by injections of the neurotoxin - kainic acid - on the first day of postnatal life. Four weeks later, as demonstrated with the anterograde and retrograde transport of the enzyme horseradish peroxidase (HRP) injected into the occipital lobe of one hemisphere, callosally projecting neurones and terminals were distributed more widely in the retinotopically organized areas 17, 18a and 18b of the visual cortex ipsilateral to the lesioned visual thalamus than in unoperated control animals of the same age. By contrast, in the visual cortex contralateral to the lesioned visual thalamus the areal distribution of callosally projecting neurones and terminals was similar to that of the controls, that is, largely but not exclusively restricted to the common border of areas 17 and 18a. Both in unoperated and operated animals, cells in lamina V of several cytoarchitectonically defined areas that are not retinotopically organized (area 8 in the frontal lobe, area 29d in the retrosplenial limbic cortex and perirhinal areas 35/13 in the temporal lobe) also project to contralateral visual cortices. In areas 8 and 29d, the total numbers, laminar distributions and densities of labelled callosal cells both ipsilateral and contralateral to the kainate-injected visual thalamus were similar to those in the controls. However, in the temporal lobe, the areal distribution of the labelled callosal neurones was more extensive than that in the controls and labelled cells in areas 35/13 of the cortex contralateral to the kainate-lesioned visual thalamus merged with those in the neighbouring areas 20 and 36. By contrast, the areal distribution of associational neurones in area 18a and in nonretinotopically organized areas projecting to area 17 were very similar in controls and in operated animals (neonatal kainate lesion of the visual thalamus, neonatal section of the corpus callosum or both procedures combined). However, in operated animals, the labelled associational neurones projecting from the supragranular laminae (II/III) of area 18a to area 17 constituted a higher proportion of all cells than did those in the unoperated control animals. Thus, overall the number of associational neurones projecting from area 18a to area 17 was slightly increased by the experimental manipulations performed.(ABSTRACT TRUNCATED AT 400 WORDS)

Afferent Pathways↗

Texture segregation in the human visual cortex: A functional MRI study.

The segregation of visual scenes based on contour information is a fundamental process of early vision. Contours can be defined by simple cues, such as luminance, as well as by more complex cues, such as texture. Single-cell recording studies in monkeys suggest that the neural processing of complex contours starts as early as primary visual cortex. Additionally, lesion studies in monkeys indicate an important contribution of higher order areas to these processes. Using functional MRI, we have investigated the level at which neural correlates of texture segregation can be found in the human visual cortex. Activity evoked by line textures, with and without texture-defined boundaries, was compared in five healthy subjects. Areas V1, V2/VP, V4, TEO, and V3A were activated by both kinds of line textures as compared with blank presentations. Textures with boundaries forming a checkerboard pattern, relative to uniform textures, evoked significantly more activity in areas V4, TEO, less reliably in V3A, but not in V1 or V2/VP. These results provide evidence that higher order areas with large receptive fields play an important role in the segregation of visual scenes based on texture-defined boundaries.

Adult↗

Variations by layers and developmental changes in expression of telencephalin in the visual cortex of cat.

The expression of telencephalin in visual cortex of cat and monkey was studied immunohistochemically. In adult cats and monkeys, immunoreactivity to a polyclonal antibody raised against telencephalin was especially low in layer IV, which receives massive afferent input from the thalamus. In kitten visual cortex, the antibody bound both layer IV and other cortical layers during the most sensitive period for ocular dominance plasticity. Outside the sensitive period, the staining of layer IV was selectively reduced. These findings suggest that the expression of telencephalin is developmentally regulated during the early period and may play a role in regulating plasticity during the sensitive period.

Aging↗

Modulations of primary visual cortex activity representing attentive and conscious scene perception.

In the visual cortex, information is transferred from one area to the next by means of feedforward connections. These connections shape the receptive field properties of neurons in subsequent visual areas. Horizontal and feedback connections modulate this neuronal activity, resulting in the phenomenon of contextual modulation. In area V1, where receptive field properties reflect only low level processing, contextual modulation can be observed that represents fully evaluated perceptual saliency of the features within the receptive field. Here, we discuss to what extent these modulations are related to high level visual processes like perceptual organization, attention and visual awareness. Contextual modulation appears to reflects a process very distinct from receptive field based processing. This process seems to integrate information from distant areas in visual cortex to neurophysiologically 'highlight' those neurons that represent image elements or features of objects that stand out perceptually. Moreover, similar modulations are observed in relation to whether objects are attended to or not. Finally, these modulations are only present when subjects are aware of the visual input.

Animals↗

Silent synapses in the developing rat visual cortex: evidence for postsynaptic expression of synaptic plasticity.

In the developing visual cortex activity-dependent refinement of synaptic connectivity is thought to involve synaptic plasticity processes analogous to long-term potentiation (LTP). The recently described conversion of so-called silent synapses to functional ones might underlie some forms of LTP. Using whole-cell recording and minimal stimulation procedures in immature pyramidal neurons, we demonstrate here the existence of functionally silent synapses, i.e., glutamatergic synapses that show only NMDA receptor-mediated transmission, in the neonatal rat visual cortex. The incidence of silent synapses strongly decreased during early postnatal development. After pairing presynaptic stimulation with postsynaptic depolarization, silent synapses were converted to functional ones in an LTP-like manner, as indicated by the long-lasting induction of AMPA receptor-mediated synaptic transmission. This conversion was dependent on the activation of NMDA receptors during the pairing protocol. The selective activation of NMDA receptors at silent synapses could be explained presynaptically by assuming a lower glutamate concentration compared with functional ones. However, we found no differences in glutamate concentration-dependent properties of NMDA receptor-mediated PSCs, suggesting that synaptic glutamate concentration is similar in silent and functional synapses. Our results thus support a postsynaptic mechanism underlying silent synapses, i.e., that they do not contain functional AMPA receptors. Synaptic plasticity at silent synapses might be expressed postsynaptically by modification of nonfunctional AMPA receptors or rapid membrane insertion of AMPA receptors. This conversion of silent synapses to functional ones might play a major role in activity-dependent synaptic refinement during development of the visual cortex.

Animals↗

Orienting behavior in hamsters with lesions of superior colliculus, pretectum, and visual cortex.

We examined cortical and subcortical mediation of visual locomotor orienting function by comparing the behavior of hamsters with discrete bilateral lesions affecting the pretectum, superior colliculus (SC), or visual cortex (VC). Orienting and approach to stationary targets was evaluated by measuring the accuracy of hamsters' approaches to small black apertures, located at eye level along the wall of a circular white arena. Hamsters with bilateral ablation of the visual cortex were slightly impaired for approaches to central field targets, whereas those with ibotenic acid lesions of the pretectum (which spares fibers of passage and thus leaves tectal afferents intact) were totally unimpaired. Hamsters with transection of the brachium of SC (BSC) at the prectectal-SC (PT-SC) border were severely impaired in their ability to approach stationary targets in central and peripheral fields. Thus, we did not detect any of the central field sparing that has been reported by others for rodents with similar lesions. Several possible reasons for the disparity between our results and those of others are discussed. Overall, our results indicate that in hamsters the SC is essential for normal visually guided approach to dark, stationary targets throughout the visual field. Further, our results and qualitative observations indicate that the approach errors are most likely due to deficits of visuomotor integration rather than to a lack of visual scanning.

Animals↗

Effects of neonatal monocular enucleation on the number of GAD-positive puncta in rat visual cortex.

Rats that had one eye removed on the day of birth were examined at various postnatal ages with immunocytochemical methods to determine the effect on the development of the GABAergic axonal plexus in the visual cortex. The monocular segment of visual cortex contralateral to the enucleated orbit had 20-30% fewer GABAergic axon terminals than the monocular segment of visual cortex contralateral to the normal eye. Other cortical areas did not show any significant changes. These findings suggest that sensory deprivation of the visual cortex interferes with the normal development of GABAergic neurons.

Animals↗

[Effect of nonspecific thalamic nuclei on individual ensemble-forming loci in the visual cortex].

Electric stimulation of guinea pigs' medial thalamic nuclei (MT) creates in the area 17 of the visual cortex functional groups of excited neurones, located within the same structural columns as ensembles of neurones excited by flashes of diffuse light. Together with analogous effects of the MBRF stimulation, such results indicate a convergence of different non-specific influences on separate columns of the visual cortex. MT exert their initial and mainly activating influences on the lower, efferent layers, while MBRF, according to literary data, activates afferent, middle layers of the cortex. This points to the existence in the visual cortex of separate non-specific regulation of the input on the part of the reticular formation and of the output on the part of MT.

Afferent Pathways↗

Effect of the richness of the environment on the cat visual cortex.

In a recent study of the cat visual cortex, it was shown that there are interindividual differences in the numerical density (Nv) of symmetrical synapses associated with flat vesicles (FS synapses) but not of asymmetrical synapses associated with round vesicles (RA synapses). Since many of the environment-sensitive properties of visual cortex neurons are GABA-dependent, it was suggested that the interindividual differences in FS synapses might be due to environmental factors. To verify this possibility we estimated the Nv of both types of synapses in two groups of six cats, paired by litter and by sex, and raised either in isolation or in a colony from the time of weaning to the age of 8 months. We also measured the Nv of neurons and the thickness of the cortex and made some gross anatomical measurements. The brains of animals raised in the enriched environment are 7% heavier, and their total body weight is 10% greater: The brain-to-body-weight ratio remains unchanged. The total length of the brain is not affected, but the length and width of the cerebral hemispheres are each 5% greater in the enriched cats. As in comparable rat studies, the thickness of the cortex is 4% greater, but in the present study this difference is not significant. The numerical density of neurons is diminished by 17% in enriched animals. This is probably due to a wider separation of neuronal cell bodies in a larger cortical volume, rather than to a loss of neurons. There are no significant changes in the numerical density of RA synapses between the two milieux, but there are nearly twice as many FS synapses per mm3 of tissue in the impoverished cortex. The coefficient of variation of FS synapses, which in the previous study was on the order of 30%, has been reduced to 10% and 7% in enriched and impoverished cats, respectively. We conclude that environmental conditions can lead to selective interindividual differences in the Nv of FS synapses, as seen in our previous study of animals whose rearing conditions were not controlled. The average diameter of RA synaptic profiles is not affected by the environment but FS synapses are 25% wider in the enriched animals. Because of the smaller neuronal Nv in enriched animals, there are, in fact, 18% more RA synapses and 34% fewer FS synapses per neuron in the enriched condition.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗