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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↗

LTP and LTD vary with layer in rodent visual cortex.

Mechanisms of plasticity in the visual cortex have been studied with long-term potentiation (LTP), long-term depression (LTD) and ocular dominance plasticity (ODP). It is now possible to compare results from these three forms of plasticity using knockout mice, and also by pharmacological manipulations. A review of the literature shows that if both LTP and LTD are completely abolished, then ODP will also be abolished. In other situations, there is little correlation. We hypothesize that this lack of correlation occurs because the mechanisms for LTP and LTD vary with layer in the visual cortex, and results show that they do.

Animals↗

[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↗

NMDA receptor-dependent ocular dominance plasticity in adult visual cortex.

The binocular region of mouse visual cortex is strongly dominated by inputs from the contralateral eye. Here we show in adult mice that depriving the dominant contralateral eye of vision leads to a persistent, NMDA receptor-dependent enhancement of the weak ipsilateral-eye inputs. These data provide in vivo evidence for metaplasticity as a mechanism for binocular competition and demonstrate that an ocular dominance shift can occur solely by the mechanisms of response enhancement. They also show that adult mouse visual cortex has a far greater potential for experience-dependent plasticity than previously appreciated. These insights may force a revision in how data on ocular dominance plasticity in mutant mice have been interpreted.

Aging↗

Language-specific tuning of visual cortex? Functional properties of the Visual Word Form Area.

The first steps in the process of reading a printed word belong to the domain of visual object perception. They culminate in a representation of letter strings as an ordered set of abstract letter identities, a representation known as the Visual Word Form (VWF). Brain lesions in patients with pure alexia and functional imaging data suggest that the VWF is subtended by a restricted patch of left-hemispheric fusiform cortex, which is reproducibly activated during reading. In order to determine whether the operation of this Visual Word Form Area (VWFA) depends exclusively on the visual features of stimuli, or is influenced by language-dependent parameters, brain activations induced by words, consonant strings and chequerboards were compared in normal subjects using functional MRI (fMRI). Stimuli were presented in the left or right visual hemifield. The VWFA was identified in both a blocked-design experiment and an event-related experiment as a left-hemispheric inferotemporal area showing a stronger activation to alphabetic strings than to chequerboards, and invariant for the spatial location of stimuli. In both experiments, stronger activations of the VWFA to words than to strings of consonants were observed. Considering that the VWFA is equally activated by real words and by readable pseudowords, this result demonstrates that the VWFA is initially plastic and becomes attuned to the orthographic regularities that constrain letter combination during the acquisition of literacy. Additionally, the use of split-field stimulation shed some light on the cerebral bases of the classical right visual field (RVF) advantage in reading. A left occipital extrastriate area was found to be activated by RVF letter strings more than by chequerboards, while no symmetrical region was observed in the right hemisphere. Moreover, activations in the precuneus and the left thalamus were observed when subjects were reading RVF versus left visual field (LVF) words, and are likely to reflect the attentional component of the RVF advantage.

Adult↗

[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↗

State-dependent effects of light-dark cycle on somatosensory and visual cortex EEG in rats.

Somatosensory (SSctx) and visual cortex (Vctx) EEG were evaluated in rats under a 12:12-h light-dark (LD) cycle and under constant light (LL) or constant dark (DD) in each sleep or wake state. Under LD conditions during light period, relative Vctx EEG slow-wave activity (SWA) was higher than that of the SSctx, whereas during dark period, relative Vctx EEG SWA was lower than in the SSctx. These effects were state specific, occurring only during non-rapid eye movement sleep (NREMS). Under LL conditions, the duration of REMS and NREMS during the period that would have been dark if the LD cycle had continued (subjective dark period) was greater than under LD conditions. DD conditions had little effect on the duration of NREMS and REMS. SSctx and Vctx EEG SWA were suppressed by LL during the subjective dark period; however, the degree of Vctx SWA suppression was smaller than that of the SSctx. DD conditions during the subjective light period enhanced SSctx SWA, whereas Vctx SWA was suppressed. Under LL conditions during the subjective dark period, Vctx EEG power was higher than that of the SSctx across a broad frequency range during NREMS, REMS, and wakefulness. During DD, SSctx EEG power during NREMS was higher than that of the Vctx in the delta wave band, whereas SSctx power during REMS and wakefulness was higher than that of the Vctx in frequencies higher than 8 Hz. We concluded that the SSctx and Vctx EEGs are differentially affected by light during subsequent sleep. Results provide support for the notion that regional sleep intensity is dependent on prior regional afferent input.

Adaptation, Physiological↗

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↗

Binocular impulse blockade prevents the formation of ocular dominance columns in cat visual cortex.

Ocular dominance columns in the cat's visual cortex appear to develop out of an initially overlapping projection by a progressive segregation of the geniculocortical afferents serving the 2 eyes (reviewed in LeVay and Stryker, 1979). To determine whether electrical activity in the visual afferent pathway is involved in this normal, developmental rearrangement of synaptic connections, we blocked the discharge of retinal ganglion cells in both eyes by making repeated intravitreal injections of tetrodotoxin (TTX) during the period in which geniculocortical afferent segregation would normally be taking place. Control experiments for the side effects of the injection procedure, the systemic effects of TTX, and the effects of visual deprivation were carried out, and a series of normal animals of appropriate ages was also studied. We then examined the effects of retinal blockade and the various control procedures on the formation of ocular dominance columns using an anatomical assay, the autoradiographic labeling of geniculocortical afferent terminals in layer IV of the visual cortex by the transneuronal transport of tritiated proline injected into 1 eye, and a physiological assay, the ocular dominance of single cortical cells recorded extracellularly. After retinal TTX blockade, layer IV was labeled uniformly without periodic fluctuation in grain density, and nearly all cortical cells were driven well through both eyes. These assays thus indicated that retinal blockade completely blocked the formation of ocular dominance columns, unlike any of the control procedures, suggesting that the spontaneous maintained discharge of retinal ganglion cells may have an important role in the normal development of binocular connections in the visual cortex.

Animals↗

Distribution of glial fibrillary acidic protein and vimentin immunoreactivity during rat visual cortex development.

The postnatal maturation of astrocytes in the rat visual cortex was analysed by immunostaining the astroglial proteins vimentin and glial fibrillary acidic protein with poly- and monoclonal antibodies. Vimentin immunoreactivity was present in the visual cortex up to the third postnatal week, whereas immunolabelling first disappeared in the cortical layers and then in the white matter. In the early postnatal period, vimentin antibodies labelled radial glial fibres. After the first postnatal week staining of radial glial fibres gradually disappeared and vimentin immunoreactivity was localized in a few protoplasmic astrocytes in the grey matter and fibrous astrocytes in the white matter. The development of glial fibrillary acidic protein-positive astrocytes was not fully complete until postnatal day 50. Glial fibrillary acidic protein-positive radial glial fibres were present after birth and disappeared towards the end of the third postnatal week. Staining of astrocytes in the white matter and in cortical layers I and VI reached an adult density at postnatal days 8 and 20, respectively. A progressively later development of glial fibrillary acidic protein-positive astrocytes was observed in cortical layers II-V which was completed between postnatal days 47 and 50. In the adult rat visual cortex glial fibrillary acidic protein-positive astrocytes were especially dense in layers I and VI, moderate in layers II/III and V and nearly absent in layer IV and lower layer III. The time course of the loss of vimentin and the gradual appearance of glial fibrillary acidic protein immunoreactivity in the visual cortex is considered as an index of astrocytic maturation and the spatiotemporal sequence of this maturation pattern is discussed in terms of reciprocal neuron-astrocyte interactions during brain development.

Animals↗

Projections from V1 to lateral suprasylvian cortex: an efferent pathway in the cat's visual cortex that originates preferentially from CO blob columns.

The patchy pattern of retrograde labeling produced by injections of anatomical tracers into the lateral suprasylvian (LS) visual area was compared to the cytochrome oxidase (CO) blobs in cat visual cortex. Following large injections of anatomical tracers in LS, retrograde labeling formed an irregular lattice of patches with a spacing of slightly less than 1 mm in area 17, and slightly greater than 1 mm in area 18. By comparing labeling in alternate serial sections, patches of LS-projecting cells in both areas were found to align with CO blobs. The conclusion of alignment between CO blob columns and patches of LS-projecting cells was confirmed by a quantitative analysis which showed a significant correlation between the local density of LS-projecting cells in reconstructions of charted cells and the intensity of CO staining in the CO-reacted sections. As for areas 17 and 18, labeling in other afferent areas of LS was also patchy with a spacing on the order of 1 mm except for area 19 where we found patches of LS-projecting cells with a larger spacing, roughly 2 mm. No matching fluctuations in CO density could be discerned in area 19, however. In conjunction with recent evidence that CO blob columns in cats receive strong input from Y-cells of the lateral geniculate nucleus (Boyd & Matsubara, 1996; Shoham, et al., 1996), these data support the hypothesis (Shipp & Grant, 1991) that the patches of LS-projecting cells correspond to Y-cell input columns. As a relationship between the CO architecture and certain classes of efferent cells has previously been shown in primates, these findings show new similarities between CO blobs in different mammalian species.

Animals↗

Perceptual and neuronal correspondence in primary visual cortex.

Recent findings from the study of primary visual cortex in humans and animals blur the distinction between early and late visual processing. Under some conditions, the activity of neurons in primary visual cortex appears as close or closer to perception than activity in 'higher' visual areas.

Animals↗

Plasticity of 5-hydroxytryptamine(1B) receptors during postnatal development in the rat visual cortex.

The distribution of 5-hydroxytryptamine1A and 5-hydroxytryptamine1B receptors in the visual cortex was studied by quantitative autoradiography during postnatal development. Overall, receptor densities increased throughout development, but exhibited regional rearrangements, particularly in the case of 5-hydroxytryptamine1B receptors. Neonatal treatment with 5,7-dihydroxytryptamine, which causes selective degeneration of serotoninergic neurons, had no effect on the density of 5-hydroxytryptamine1A receptors in the visual cortex. However, a transient increase in 5-hydroxytryptamine1B at postnatal days 10-12 was observed after this treatment, suggesting a regulation of postsynaptic receptors. Neonatal enucleation resulted in a marked increase in 5-hydroxytryptamine1B binding sites in all layers of the visual cortex by P16, whereas it had no effect upon 5-hydroxytryptamine1A binding sites. These results show that both receptor subtypes do not exhibit striking transient features in the visual cortex during postnatal development, but rather undergo discrete reorganizations. 5-Hydroxytryptamine1B receptors show changes in density after either neonatal degeneration of serotoninergic neurons or enucleation, indicating that the serotoninergic system involving this receptor subtype can exhibit some postnatal plasticity in the visual cortex.

5,7-Dihydroxytryptamine↗

Organization of corticocortical connections in human visual cortex.

Clinical and psychophysical observations indicate that the visual cortex is critical for the perception of color, form, depth, and movement. Little, however, is known about the cortical circuitry that underlies these functions in humans. In an attempt to learn more about these connections, we have traced projections of primary (V1) and secondary (V2) visual cortex in the postmortem, fixed human brain, using the fluorescent dye 1,1'-dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate as an axonal marker. The results show that V1 makes a forward projection to layers 3 and 4 of V2, and V2 projects back to layers 1, 2, 3, 5, and 6 of V1. Some V2 injections also show an input to layer 4B of V1. Projections to 4B probably originate from cytochrome oxidase (CO)-reactive stripes that we have identified in V2. Differential connections between CO-rich (blobs) and CO-poor regions (interblobs) also exist within V1; blobs are connected to blobs and interblobs are connected to interblobs. The results show that the connections in human visual cortex are similar to those of nonhuman primates and that their organization is consistent with the concept of multiple processing streams in the visual system.

Adult↗

fMR-adaptation reveals a distributed representation of inanimate objects and places in human visual cortex.

The way information about objects is represented in visual cortex remains controversial. It is unclear, for example, whether information is processed in modules, specialized for different categories of objects or whether information is represented in a distributed fashion across a large network of overlapping visual areas. In this study, we used fMR-adaptation to investigate the extent to which 'specialized' regions of visual cortex are involved in representing information about inanimate objects and places. We found adaptation in the object-selective lateral occipital complex (LOC) following repeated presentations of the same inanimate object. However, we also found fMR-adaptation to inanimate objects in fusiform face area (FFA) and the parahippocampal place area (PPA). Furthermore, this adaptation was not affected by changes in the size of the stimulus. In the second part of the experiment, we found adaptation to repeated images of places in the place-selective PPA, which was both size- and viewpoint-invariant. fMR-adaptation to repeated images of places was also observed in the LOC, but not in the FFA. These results suggest that the representation of inanimate objects and places is not restricted to those regions showing maximal responses to these particular categories of objects, but is distributed across human visual cortex and can include 'face-selective' regions such as the FFA.

Adaptation, Psychological↗