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Corticopontine projection in the rat: the distribution of labelled cortical cells after large injections of horseradish peroxidase in the pontine nuclei.

The distribution of cortical cells projecting to the pontine nuclei in rats was studied by making large injections of horseradish peroxidase that filled the basilar pons and measuring the density of labelled cells in each cortical area. All retrogradely labelled cells were layer V pyramidal cells. The highest densities of labelled cells were observed in the motor areas. The lowest densities were in temporal association cortex and perirhinal cortex. Visual cortical areas, including the primary visual cortex, provided a major source of pontine projections. The distribution of corticopontine cells within the primary visual cortex was studied in more detail. In all cases the highest density of labelled cells was observed in the region of cortex that represents the nasal visual field. Control injections into brainstem regions adjacent to the pontine nuclei produced a much lower absolute density of retrogradely labelled cortical cells and the distribution of those cells was different from that observed following pontine injections. We conclude that every area of the rat's cerebral cortex projects to the pontine nuclei and that there are consistent variations in the density of the projections both between and within areas.

Animals↗

Prosencephalic connections of striate and extrastriate areas of rat visual cortex.

Afferent connections of rat primary visual cortex (area 17 or V1 area) and the rostral and caudal parts of areas 18a and 18b were studied, by placing in each of the areas, small electrophoretic injections of enzyme horseradish peroxidase (HRP) or wheat germ agglutinated-HRP. The results indicate that: 1) each of the areas has a distinct pattern of distribution of afferent neurons in the ipsilateral visual thalamus - area 17 receives its principal thalamic input from the dorsal lateral geniculate nucleus, the caudal parts of areas 18a and 18b receive a major thalamic input from the lateral posterior nucleus and a minor input from the posterior nucleus, while the rostral parts of areas 18a and 18b receive a major input from the posterior nucleus, and a minor projection from the lateral posterior nucleus; 2) the rostral and caudal parts of areas 18a and 18b each receive an associational input from area 17; 3) the rostral parts of areas 18a and 18b each receive associational input from three different extrastriate regions, the caudal part of the same extrastriate area, and the rostral and caudal parts of the other extrastriate area, whereas the caudal parts of areas 18a and 18b receive associational inputs only from one or two extrastriate regions; 4) area 17, area 18b and rostral area 18a each receive a substantial associational input from lamina V of the caudal part of the frontal eye field (FEF) in the motor cortex; however the input from the FEF to caudal area 18a (if present) is very small; 5) The extrastriate areas studied receive associational input from the restrosplenial cingulate area 29d; however, the input from area 29d to area 17 appears to be very small. The distinct patterns of distribution of prosencephalic afferents suggest to us that multiple retinotopically organized areas described previously in the rat cortex (cf Montero 1981; Espinoza and Thomas 1983) represent functionally distinct areas.

Afferent Pathways↗

Gene expression analysis of the critical period in the visual cortex.

The development of the primary visual cortex in animals possessing binocular vision is a classical paradigm for the study of activity-dependent neuronal plasticity. To elucidate the genetic determinants of this period of substantial plasticity, we conducted an unbiased and comprehensive transcript profiling analysis with differential display and DNA array techniques. We characterized the transcripts that change significantly between the critical and postcritical periods in the rat binocular visual cortex. We determined if these changes are specific for the visual cortex by simultaneously profiling the hippocampus and examined the impact of sensory experience on the accumulation of the identified transcripts. Our results uncover visual cortex-specific and unspecific transcription programs. Transcripts for protein kinases and phosphatases are particularly regulated. The identified transcripts support the notion that the critical period provides a permissive state for plasticity.

Animals↗

Neuropeptide Y-containing neurons are situated predominantly outside cytochrome oxidase puffs in macaque visual cortex.

Layers II/III of the primary visual cortex contain a regular pattern of histochemically detectable cytochrome oxidase (CO)-rich "puffs," which differ from the interpuff regions in their thalamo-cortical and cortico-cortical connectivity, receptive-field properties, and the density of inhibitory GABA-containing synaptic terminals. We used an immunocytochemical method, in combination with cytochrome oxidase histochemistry, to analyze the spatial relationship between neurons that contain neuropeptide Y (NPY) and the CO puffs. Of a total of 606 neurons, only 2.6% of the NPY-containing cells are located in the puffs, whereas the rest are situated in the interpuffs, or at the interface between puffs and interpuffs. The number of NPY-containing neurons in the puffs is substantially less than that expected in an equal volume of the interpuffs (X2 = 13.86; df = 1; P less than 0.001). These observations indicate that columns containing the puffs may differ also from those in the interpuff regions in that they contain a unique array of chemically and morphologically distinct local circuit neurons.

Animals↗

Interocular control of neuronal responsiveness in cat visual cortex.

Neurons in the cat primary visual cortex are selective for particular contour orientations but their responsiveness can vary under certain conditions. After prolonged stimulation (adaptation), the contrast sensitivity of cortical cells is reduced and the 'gain' (the strength of response as a function of contrast) falls. The response to an optimal contour is also reduced when a different stimulus is superimposed on the receptive field in the same eye. Here we report that the sudden appearance of an inappropriate stimulus in one eye can interocularly suppress the activity of cortical neurons if they are already responding to an optimally oriented stimulus in the other eye. In strabismic cats, whose cortical neurons lack binocular facilitation, even contours of similar orientation shown to the two eyes trigger such suppression. This interocular control of cortical responsiveness could serve to veto signals from one eye under conditions that would otherwise cause double vision and perceptual confusion.

Animals↗

Motion and shape perception in cerebral akinetopsia.

Motion cues serve many purposes in primate vision. Consequently, akinetopsia, a defect of movement perception due to cerebral lesions, would be expected to comprise a range of motion-related defects. To address this issue we explored further the perceptual profiles in akinetopsic subject L.M. who has motion perception deficits due to extensive bilateral lesions of the dorsolateral visual association cortex that spare primary visual cortex, area V1. We used several different experimental procedures. Using random-dot motion stimuli, we showed that L.M. can still perceive global coherent motion and discriminate motion direction, yet these abilities fail even at moderate levels of background noise. L.M. also viewed a two-frame apparent (phi) motion stimulus known as a Ternus display. Her performance on this test suggests defective 'long-range' apparent motion mechanisms. These results are clearly in agreement with previous reports. Additional experiments showed that L.M. can still perceive 2-D shape and 3-D structure-from-motion (SFM). However, like motion direction discrimination, these abilities broke down at moderate levels of moving and stationary noise. Surprisingly, L.M. also had trouble perceiving 2-D shapes defined by non-motion signals including 'on' and 'off' transients, dynamic and static binocular disparity, and static texture cues. Our findings highlight the role of the visual association cortex in extracting salient information from noise.

Adult↗

The Role of Horizontal Connections in Generating Long Receptive Fields in the Cat Visual Cortex.

The cells in the primary visual cortex possess numerous functional properties that are more complex and varied than those seen in the cortical input. These properties result from the network of intrinsic cortical connections running across the cortical layers and between cortical columns. In the current study we relate the long receptive fields that are characteristic of layer 6 cells to the input that these cells receive from layer 5. The axons of layer 5 pyramidal cells project over long distances within layer 6, enabling layer 6 cells to collect input from regions of cortex representing large parts of the visual field. When layer 5 was locally inactivated by injection of the inhibitory transmitter GABA, layer 6 cells lost sensitivity over the portion of their receptive fields corresponding to the inactivated region of layer 5. This suggests that the extensive convergence in the projection from layer 5 to layer 6 is responsible for generating the long receptive fields characteristic of the layer 6 cells.

Journal Article↗

Acupuncture stimulation of the vision-related acupoint (Bl-67) increases c-Fos expression in the visual cortex of binocularly deprived rat pups.

Our previous study with functional magnetic resonance imaging (MRI) demonstrated that acupuncture stimulation of the vision-related acupoint, B1-67, activates the visual cortex of the human brain. As a further study on the effect of B1-67 acupuncture stimulation on the visual cortex, we examined c-Fos expression in binocularly deprived rat pups. Binocular deprivation significantly reduced the number of c-Fos-positive cells in the primary visual cortex, compared with that of normal control rat pups. Interestingly, acupuncture stimulation of B1-67 resulted in a significant increase in the number of c-Fos-positive cells in the primary visual cortex, while acupuncture stimulation of other acupoints less important for visual function had no significant effect on c-Fos expression in the primary visual cortex. The results suggest the possibility of vision-related acupoint (B1-67) having an influence over the activity of the primary visual cortex.

Acupuncture Points↗

The role of spared calcarine cortex and lateral occipital cortex in the responses of human hemianopes to visual motion.

Some patients, who are rendered perimetrically blind in one hemifield by cortical lesions, nevertheless exhibit residual visual capacities within their field defects. The neural mechanism that mediates the residual visual responses has remained the topic of considerable debate. One explanation posits the subcortical visual pathways that bypass the primary visual cortex and innervate the extrastriate visual areas as the substrate that underlies the residual vision. The other explanation is that small islands of the primary visual cortex remain intact and provide the signals for residual vision. We have performed behavioral and functional magnetic resonance imaging experiments to investigate the validity of the two explanations of residual vision. Our behavioral experiments indicated that of the seven hemianopes tested, two had the ability to discriminate the direction of a drifting grating. This residual visual response was shown with fMRI to be the result of spared islands of calcarine cortical activity in one of the hemianopes, whereas only lateral occipital activity was documented in the other patient. These results indicate that the underlying neural correlates of residual vision can vary between patients. Moreover, our study emphasizes the necessity of ruling out the presence of islands of preserved function and primary visual cortex before assigning residual visual capacities to the properties of visual pathways that bypass the primary visual cortex.

Adult↗

Development of blobs in the visual cortex of macaques.

We have examined the area of the primary visual cortex and the number and size of blobs within it in 10 neonatal and 11 adult rhesus monkeys. The average area of the primary visual cortex (V1) increases from 919 mm2 in newborns to 1,069 mm2 in adult animals (16%). The number of blobs decreases per unit area from an average of 5.2/mm2 at birth to 4.3/mm2 in maturity (18%). As a consequence, the number of blobs remains approximately the same during maturation, at about 4,800/hemisphere. These observations correct a preliminary report on a subset of the animals studied here (Purves and LaMantia: Proc Natl Acad Sci 87:5765, '90), in which it appeared that blob number might increase between birth and maturity. As in other regions of the developing postnatal brain, we found no net loss of modular circuitry.

Animals↗

Gaze direction controls response gain in primary visual-cortex neurons.

To localize objects in space, the brain needs to combine information about the position of the stimulus on the retinae with information about the location of the eyes in their orbits. Interaction between these two types of information occurs in several cortical areas, but the role of the primary visual cortex (area V1) in this process has remained unclear. Here we show that, for half the cells recorded in area V1 of behaving monkeys, the classically described visual responses are strongly modulated by gaze direction. Specifically, we find that selectivity for horizontal retinal disparity-the difference in the position of a stimulus on each retina which relates to relative object distance-and for stimulus orientation may be present at a given gaze direction, but be absent or poorly expressed at another direction. Shifts in preferred disparity also occurred in several neurons. These neural changes were most often present at the beginning of the visual response, suggesting a feedforward gain control by eye position signals. Cortical neural processes for encoding information about the three-dimensional position of a stimulus in space therefore start as early as area V1.

Animals↗

Thinning of the cerebral cortex in aging.

The thickness of the cerebral cortex was measured in 106 non-demented participants ranging in age from 18 to 93 years. For each participant, multiple acquisitions of structural T1-weighted magnetic resonance imaging (MRI) scans were averaged to yield high-resolution, high-contrast data sets. Cortical thickness was estimated as the distance between the gray/white boundary and the outer cortical surface, resulting in a continuous estimate across the cortical mantle. Global thinning was apparent by middle age. Men and women showed a similar degree of global thinning, and did not differ in mean thickness in the younger or older groups. Age-associated differences were widespread but demonstrated a patchwork of regional atrophy and sparing. Examination of subsets of the data from independent samples produced highly similar age-associated patterns of atrophy, suggesting that the specific anatomic patterns within the maps were reliable. Certain results, including prominent atrophy of prefrontal cortex and relative sparing of temporal and parahippocampal cortex, converged with previous findings. Other results were unexpected, such as the finding of prominent atrophy in frontal cortex near primary motor cortex and calcarine cortex near primary visual cortex. These findings demonstrate that cortical thinning occurs by middle age and spans widespread cortical regions that include primary as well as association cortex.

Adolescent↗

Receptive field expansion in adult visual cortex is linked to dynamic changes in strength of cortical connections.

1. Receptive field (RF) sizes of neurons in adult primary visual cortex are dynamic, expanding and contracting in response to alternate stimulation outside and within the RF over periods ranging from seconds to minutes. The substrate for this dynamic expansion was shown to lie in cortex, as opposed to subcortical parts of the visual pathway. The present study was designed to examine changes in cortical connection strengths that could underlie this observed plasticity by measuring the changes in cross-correlation histograms between pairs of primary visual cortex neurons that are induced to dynamically change their RF sizes. 2. Visually driven neural activity was recorded from single units in the superficial layers of primary visual cortex in adult cats, with two independent electrodes separated by 0.1-5 mm at their tips, and cross-correlated on-line. The neurons were then conditioned by stimulation with an "artificial scotoma," a field of flashing random dots filling the region of visual space around a blank rectangle enclosing the RFs of the recorded neurons. The neuronal RFs were tested for expansion and their visually driven output again cross-correlated. After this, the neurons were stimulated vigorously through their RF centers to induce the field to collapse, and the visually driven output from the collapsed RFs was again cross-correlated. Cross-correlograms obtained before and after conditioning, and after RF collapse, were normalized by their flanks to control for changes in peak size due solely to fluctuations in spike rate. 3. A total of 37 pairs of neurons that showed distinct cross-correlogram peaks, and whose RF borders were clearly discernible both before and after conditioning, were used in the final analysis. Of these neuron pairs, conditioning led to a clear expansion of RF boundaries in 28 pairs, whereas in 9 pairs the RFs did not expand. RFs that did expand showed no significant shifts in their orientation preference, orientation selectivity, or ocularity. 4. When the RFs of a pair of neurons expanded with conditioning, the area of the associated flank-normalized cross-correlogram peaks also increased (by a factor ranging from 0.84 up to 3.5). Correlograms returned to their preconditioning values when RFs collapsed.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

Brain representation of hemifield stimulation in poststroke visual field defects.

BACKGROUND AND PURPOSE: Plasticity in extended, parallel, or reciprocal operating networks is well recognized. Changes in neuronal activity after lesions to distinct localized structures, such as the primary visual cortex, are less well characterized. We investigated the cortical reorganization in patients with poststroke visual field defects using blood oxygen level-dependent functional MRI. METHODS: Brain activation was measured in 7 patients with a single occipital cortical lesion and partially recovered hemianopia and in 7 age-matched control subjects. Differences in activation between rest and visual hemifield stimulation were assessed with statistical parametric mapping (SPM'99). RESULTS: In normal subjects, significant activation was found in the contralateral primary visual cortex and bilaterally in the extrastriate cortex. During hemifield stimulation of the unaffected side of stroke patients, a similar pattern was found compared with that seen in control subjects. During stimulation of the hemianopic side, bilateral activation was seen within the extrastriate cortex, stronger in the ipsilateral hemisphere. The primary visual cortex was not significantly activated in either hemisphere during stimulation of the hemianopic side. CONCLUSIONS: Visual field defects after stroke are associated with bilateral activation of the extrastriate visual cortex. This pattern of activation indicates altered neuronal activity in the visual system. Further investigation is necessary to determine the relationship between functional reorganization and recovery of lost visual function after poststroke hemianopia.

Adult↗

Asymmetric suppression outside the classical receptive field of the visual cortex.

Areas beyond the classical receptive field (CRF) can modulate responses of the majority of cells in the primary visual cortex of the cat (). Although general characteristics of this phenomenon have been reported previously, little is known about the detailed spatial organization of the surrounds. Previous work suggests that the surrounds may be uniform regions that encircle the CRF or may be limited to the "ends" of the CRF. We have examined the spatial organization of surrounds of single-cell receptive fields in the primary visual cortex of anesthetized, paralyzed cats. The CRF was stimulated with an optimal drifting grating, whereas the surround was probed with a second small grating patch placed at discrete locations around the CRF. For most cells that exhibit suppression, the surrounds are spatially asymmetric, such that the suppression originates from a localized region. We find a variety of suppressive zone locations, but there is a slight bias for suppression to occur at the end zones of the CRF. The spatial pattern of suppression is independent of the parameters of the suppressive stimulus used, although the effect is clearest with iso-oriented surround stimuli. A subset of cells exhibit axially symmetric or uniform surround fields. These results demonstrate that the surrounds are more specific than previously realized, and this specialization has implications for the processing of visual information in the primary visual cortex. One possibility is that these localized surrounds may provide a substrate for figure-ground segmentation of visual scenes.

Animals↗

Comparison of the hemodynamic response to different visual stimuli in single-event and block stimulation fMRI experiments.

Experiments with three different types of basic visual stimulation were performed to compare cortical activation in single-event and block trials. Independent of the stimulation paradigm, the single-event presentation leads to highly consistent signal responses regarding both the activated cortical areas and the dynamics of the signal time course. In contrast, signal time courses during block paradigms depend on the stimulus applied and are a complex and nonlinear function of the single-event responses. Additionally, the initial dip during the first 2 seconds after stimulus onset is consistently observed. However, the small amplitude change (-0.1% to -0.3%) requires signal averaging to establish statistical significance of the effect. Furthermore, different patterns of activation were observed within the primary visual cortex. In an anterior part of the primary visual cortex, activation was only observed at the onset and at the cessation of stimulation involving luminance changes.

Adult↗

Functional consequences of modification of callosal connections by perinatal enucleation in rat visual cortex.

The effects of neonatal monocular enucleation (right eye) on the callosal connections in the rat visual cortex were studied by physiological and morphological methods. Evoked activity was recorded in the left hemisphere, i.e. contralaterally to the enucleated eye. After enucleation, trans-callosally evoked responses were recorded in a widened stripe of the lateral visual cortex. Compared with the controls, the responsive area was expanded laterally and medially, i.e. into the lateral part of the primary visual area and within the secondary visual cortex (lateral part). Within about 0.5 mm of the expansion, the responses did not differ from those recorded in areas with "normal" callosal connections. Morphological evidence is presented suggesting that this expansion of evoked responses with high amplitudes and short latencies corresponds to an extension of callosal connections with a high density of axon terminals in layers two and three. Further medially within the primary visual cortex, callosally evoked responses with low amplitudes and longer latencies were recorded. The main types of unit responses and characteristic interactions between visually and callosally evoked responses are shown and discussed. These results suggest that following neonatal enucleation (1) the callosal connections expand and form functional synapses in the lateral part of the visual cortex, (2) these connections can activate cortical neurons either directly or by mediation of associational connections between the lateral secondary and primary visual cortex areas and (3) callosal connections can interact with visually evoked potentials and unit responses.

Animals↗

The role of ipsilateral and contralateral inputs from primary cortex in responses of area 21a neurons in cats.

Neuronal responses in cat visual area 21a were analyzed when the primary visual cortex (areas 17 and 18) was deactivated by cooling. Ipsilateral and contralateral cortices were deactivated separately. Results established that (1) cooling the ipsilateral primary cortex diminished the activity of all area 21a cells and, in 30%, blocked responsiveness altogether, and (2) cooling the contralateral primary cortex initially increased activity in area 21a cells but, with further cooling, reduced it to below the original level although only 9% of cells ceased responding. These findings were then compared to earlier results in which bilateral deactivation of the primary cortex greatly reduced and, in most cases, blocked the activity of area 21a cells (Michalski et al., 1993). Despite the response attenuation following cooling of the primary visual cortex (either ipsilateral or contralateral), neurons of area 21a retained their original orientation specificity and sharpness of tuning (measured as the half-width at half-height of the orientation tuning curve). Direction selectivity also tended to remain unchanged. We concluded that for area 21a cells (1) the ipsilateral primary cortex provides the main excitatory input; (2) the contralateral primary cortex supplies a large inhibitory input; and (3) the nature of orientation specificity, sharpness of orientation tuning, and direction selectivity are largely unaffected by removal of the ipsilateral hemisphere excitatory input or the contralateral hemisphere inhibitory input.

Animals↗