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Stimulus-specific neuronal oscillations in orientation columns of cat visual cortex.

In areas 17 and 18 of the cat visual cortex the firing probability of neurons, in response to the presentation of optimally aligned light bars within their receptive field, oscillates with a peak frequency near 40 Hz. The neuronal firing pattern is tightly correlated with the phase and amplitude of an oscillatory local field potential recorded through the same electrode. The amplitude of the local field-potential oscillations are maximal in response to stimuli that match the orientation and direction preference of the local cluster of neurons. Single and multiunit recordings from the dorsal lateral geniculate nucleus of the thalamus showed no evidence of oscillations of the neuronal firing probability in the range of 20-70 Hz. The results demonstrate that local neuronal populations in the visual cortex engage in stimulus-specific synchronous oscillations resulting from an intracortical mechanism. The oscillatory responses may provide a general mechanism by which activity patterns in spatially separate regions of the cortex are temporally coordinated.

Animals↗

NADH augments blood flow in physiologically activated retina and visual cortex.

The mechanism(s) that increase retinal and visual cortex blood flows in response to visual stimulation are poorly understood. We tested the hypothesis that increased transfer of electrons and protons from glucose to cytosolic free NAD(+), reducing it to NADH, evoked by increased energy metabolism, fuels redox-signaling pathways that augment flow. The near-equilibrium between free cytosolic NADH/NAD(+) and lactate/pyruvate ratios established by lactate dehydrogenase predicts that transfer of additional electrons and protons from injected lactate to NAD(+) will augment the elevated blood flows in stimulated retina and cortex, whereas transfer of electrons and protons from NADH to injected pyruvate will attenuate the elevated flows. These predictions were tested and confirmed in rats. Increased flows evoked by stimulation also were prevented by inhibition of nitric oxide synthase. These findings support an important role for cytosolic free NADH in fueling a signaling cascade that increases *NO production, which augments blood flow in photostimulated retina and visual cortex.

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↗

Postnatal expression profile of OBCAM implies its involvement in visual cortex development and plasticity.

This study examined the expression of a neuron-specific cell adhesion molecule, OBCAM (opioid-binding cell adhesion molecule), at both the mRNA and protein levels in the cat primary visual cortex at various postnatal ages, using cDNA array analysis and immunocytochemistry. Results obtained using both methods showed that the expression level of OBCAM was high in young and low in older and adult visual cortex. OBCAM-immunoreactivities were associated predominantly with perikarya and dendrites of pyramidal neurons, and OBCAM-immunopositive neurons were present in all cortical layers. Immunostaining of OBCAM in adult visual cortex showed a reduced number of immunopositive neurons and neurites and relatively lower staining intensities as compared with younger animals. In addition, the number of OBCAM-immunopositive neurons was significantly higher in the visual cortex of 4-month-old animals dark-reared from birth than those in age-matched normally reared animals. These results suggest that OBCAM may play an important role in visual cortex development and plasticity.

Aging↗

Extrageniculate projections to the visual cortex in the macaque monkey: an HRP study.

Extrageniculate projections to the visual cortex were examined in the macaque monkeys by the horseradish peroxidase (HRP) method. Extrageniculate neurons sending fibers to the visual cortex were found in the lateral and inferior pulvinar nuclei, paracentral thalamic nucleus, claustrum, basal nucleus of Meynert, lateral part of the basal amygdaloid nucleus, lateral hypothalamus, locus coeruleus, and dorsomedial and midline regions of the pontine tegmentum.

Animals↗

Small lesions in the primary visual cortex of rats cause a specific reorganization of associational connections.

Neuroplastic changes in associational connections were investigated 3 weeks after the intrinsic organization of the visual cortex of rats had been partially damaged by small cylindrical lesions (type I). These lesions caused the degeneration of short intracortical connections and associational connections that form patches in the primary and secondary visual areas. The resulting terminal degeneration disappeared within 20 days p.o. after which only some fiber degeneration was evident in the infragranular layers. Patches of terminal degeneration reappeared in the vicinity of the stab wounds, when the associational connections between the retrosplenial and the primary visual cortex had been secondarily interrupted by elongated lesions (type II), which penetrated the paramedian cortex and subcortical white matter. When type-II lesions were made in the intact cortex, patches of degeneration were absent, although in both cases some terminal degeneration was diffusely distributed in the primary visual cortex. Horseradish peroxidase (HRP) was applied to sites similar to those where type-I lesions were applied. In the intact cortex, HRP caused a granular labeling of numerous neurons in various positions including the retrosplenial cortex and patches of the postero-median visual cortex. HRP was also applied to type-I lesions that had been made 3 weeks earlier. In these cases, apparently HRP labeled the same subpopulations of neurons as it did in the intact cortex. However, a fraction of the labeled neurons showed a Golgi-like staining (e.g., 27% of the labeled neurons in the retrosplenial cortex) only when HRP was applied to stab wounds.(ABSTRACT TRUNCATED AT 250 WORDS)

Afferent Pathways↗

The processing of stereoscopic information in human visual cortex: psychophysical and electrophysiological evidence.

Three-dimensional depth perception relies in part on the binocular fusion of horizontally disparate stimuli presented to the left and right eye. The mammalian visual system offers a unique possibility to study electrophysiologically cortical neuronal mechanisms: since the input of the two eyes remains separated up to the level of the visual cortex, evoked potential components that are generated exclusively by cortical structures may be explored when dynamic random-dot stereograms (dRDS) are presented. In a series of independent studies, we determined the scalp topography of dRDS evoked brain activity in different groups of healthy subjects, and we found consistent results. Major differences between stereoscopic and contrast evoked brain activity are seen in the strength of the potential fields as well as in their topography. Our findings suggest that there are fewer neurons in the human visual cortex that are responsive to horizontal disparity, and that higher visual areas like V2 are more engaged with stereoscopic processing than the primary visual cortex. On the other hand, component latencies of evoked brain activity show no effect signifying that the binocular information flow to the visual cortex has a similar time course for both the processing of contrast information and of dRDS stimuli. We could also verify that healthy subjects can learn to perceive 3D structure contained in dRDS. Changes in perceptual ability as measured with psychophysical tests are paralleled by systematic alterations in the topography of stereoscopically evoked potential fields. Stereoscopic VEP recordings may also be of clinical use: in patients with selectively disturbed depth perception but normal visual acuity there is a high correlation between clinical symptoms, perceptual deficiency, and altered VEP amplitudes and latencies.

Brain Mapping↗

Experience-dependent plasticity in adult visual cortex.

Experience-dependent plasticity is a prominent feature of the mammalian visual cortex. Although such neural changes are most evident during development, adult cortical circuits can be modified by a variety of manipulations, such as perceptual learning and visual deprivation. Elucidating the underlying mechanisms at the cellular and synaptic levels is an essential step in understanding neural plasticity in the mature animal. Although developmental and adult plasticity share many common features, notable differences may be attributed to developmental cortical changes at multiple levels. These range from shifts in the molecular profiles of cortical neurons to changes in the spatiotemporal dynamics of network activity. In this review, we will discuss recent progress and remaining challenges in understanding adult visual plasticity, focusing on the primary visual cortex.

Animals↗

Neuronal activity in the primary visual cortex of the cat freely viewing natural images.

Many studies have now demonstrated that neurons in the visual cortex of cats and monkeys change their activity when stimuli are presented beyond their classical receptive field, and that these responses are not readily apparent from their receptive field properties. However few studies have been conducted to investigate the discharge properties of neurons in the visual cortex of animals when they are allow to freely view natural images. We employ tetrodes, which enable simultaneous and separable recordings of small numbers of neighboring neurons, to record 102 single units from 59 sites from areas 17 and 18 of two alert cats. While the animals viewed either natural images or black screens, they made frequent saccadic eye movements and gaze fixations. Fixations onto an image's location increased neuronal firing peaking at 80-100 ms after the fixation onset, to then decrease steadily with time despite continuous fixation. Saccades trigger a fast decrease in firing rate for both images and darkness. When we examined the incidence of correlated firing, we observed significant synchrony during the initial phases of visual fixations when the animals viewed natural scenes. Such synchrony was absent during saccadic eye movements and during eye movements in darkness. Our data revealed that scanning of natural scenes is associated with a rapid succession of distinct fixation-related activation patterns that included transient rate changes and excess coincident firing. The transient nature of these synchronization phenomena suggests a fast acting mechanism, which is in good agreement with the evidence that basic operations of scene analysis must be accomplished within a few tens of milliseconds in primary visual cortex.

Action Potentials↗

The effects of panretinal photocoagulation on the primary visual cortex of the adult monkey.

PURPOSE: To determine the effects of panretinal photocoagulation (PRP) on the levels of cytochrome oxidase (CO), Zif268, synaptophysin, and growth-associated protein 43 (GAP-43) in the primary visual cortex of adult monkeys. METHODS: Ten adult primates underwent unilateral argon laser PRP with instrument settings at 300 to 500 microns spot diameter, 200 to 500 mW power intensity, and 0.1 to 0.2 second duration, causing moderate to severe burns in the peripheral retina. At 20 hours, 12 days, 6 months, and 13 months after laser treatment, the visual cortex was assessed histologically for CO and immunohistochemically for Zif268, synaptophysin, and GAP-43. RESULTS: PRP resulted in transneuronal changes in the relative distributions of CO, Zif268, synaptophysin, and GAP-43 in the primary visual cortex. CO activity was relatively decreased in the lasered eye's ocular dominance columns at 12 days post-PRP, with recovery by 13 months post-PRP. The level of Zif268 was dramatically decreased in the lasered eye's ocular dominance columns at 20 hours post-PRP, with gradual recovery by 13 months post-PRP. Levels of synaptophysin and GAP-43 immunoreactivity were increased in both the lasered and the nonlasered eyes' ocular dominance columns at 6 months post-PRP. CONCLUSION: PRP treatment results in metabolic activity changes in the visual cortex of the adult monkey. These changes are followed chronologically by spatial redistribution of synaptophysin and GAP-43, neurochemicals known to play a role in cortical plasticity. This study demonstrates, for the first time, that PRP as used in the treatment of diabetic retinopathy results in a redistribution of neurochemicals in the adult monkey visual cortex. Such changes may help explain the anomalous visual functional loss often reported by patients after PRP.

Animals↗

Effects of cholinergic depletion on neuron activities in the cat visual cortex.

1. Unilateral lesions of the nucleus basalis magnocellularis (nBM), a source of cholinergic projection to the cerebral cortex, were produced by injection of kainic acid in the cat. The lesions caused a significant reduction in density of choline acetyltransferase-immunoreactive terminals in the visual cortex ipsilateral to the lesions. 2. In the primary visual cortex ipsilateral to the lesions [acetylcholine (ACh)-depleted cortex], about half of the cells had weak or undetectable visual responses, whereas in the contralateral visual cortex almost all the cells had normal responsivity. The response selectivity, such as orientation and direction selectivities, of cortical cells was not affected by the depletion of ACh. 3. The microionophoretic application of ACh to cells under observation facilitated visual responses in 83% of the cells recorded from the ACh-depleted cortex, whereas it suppressed the responses in only 9%. The application of a muscarinic antagonist, atropine, to cells in the ACh-depleted cortex was ineffective, suggesting no residual ACh activity. 4. The mean current required to induce facilitation in the cortex ipsilateral to the lesion was significantly smaller than that required in the contralateral cortex and the visual cortex of the normal cat, suggesting a supersensitivity of receptors mediating the effect or a reduction in catabolism of exogenous ACh in the ACh-depleted cortex. 5. More than half of the cells that had been unresponsive to visual stimuli became clearly responsive during the ACh application. The response magnitude of cortical cells, as a whole, increased to the same degree as that observed during the ACh application in the normal cat. 6. In addition to the decrease in the average response magnitude, there was a remarkable variability in responses of cells to motion of the slit from sweep to sweep in the ACh-depleted cortex. The application of ACh to cortical cells decreased the variability of responses and consequently made the responses much more consistent. 7. These results suggest that without ACh supplied from the nBM, most of the cortical neurons could not respond briskly and consistently to excitatory inputs and that exogenously applied ACh could reverse such an impairment of cortical neurons through intact or even supersensitive postsynaptic receptors.

Acetylcholine↗

Major glutamatergic projection from subplate into visual cortex during development.

Subplate neurons, the first neurons of the cerebral cortex to differentiate and mature, are thought to be essential for the formation of connections between thalamus and cortex, such as the system of ocular dominance columns within layer 4 of visual cortex. To learn more about the requirement for subplate neurons in the formation of thalamocortical connections, we have sought to identify the neurotransmitters and peptides expressed by the specific class of subplate neurons that sends axonal projections into the overlying visual cortex. To label retrogradely subplate neurons, fluorescent latex microspheres were injected into primary visual cortex of postnatal day 28 ferrets, just prior to the onset of ocular dominance column formation. Subsequently, neurons were immunostained with antibodies against glutamate, glutamic acid decarboxylase (GAD-67), parvalbumin, neuropeptide Y (NPY), somatostatin (SRIF), or nitric oxide synthase (NOS). Retrograde labeling results indicate that the majority of subplate neurons projecting into the cortical plate reside in the upper half of the subplate. Combined immunostaining and microsphere labeling reveal that about half of cortically projecting subplate neurons are glutamatergic; most microsphere-labeled subplate neurons do not stain for GAD-67, parvalbumin, NPY, SRIF, or NOS. These observations suggest that subplate neurons can provide a significant glutamatergic synaptic input to the cortical plate, including the neurons of layer 4. If so, excitation from the axons of subplate neurons may be required in addition to that from lateral geniculate nucleus neurons for the activity-dependent synaptic interactions that lead to the formation of ocular dominance columns during development.

Animals↗

Dynamics of receptive field size in primary visual cortex.

Recent studies have shown that the initial responses evoked by a stimulus in neurons of primary visual cortex are dominated by low spatial frequency information in the image, whereas finer spatial scales dominate later in the response. Such phenomena could arise from the dynamics of receptive field (RF) size at early stages of cortical processing. We measured changes in RF size in simple cells recorded from the primary visual cortex of anesthetized macaques by measuring their first-order spatio-temporal kernels and fitting them with two-dimensional Gabor functions at different time slices. We found that the width and length of the RF envelope and the period of the carrier tend to decrease during the time-course of the response. The most pronounced changes are seen in the width and spatial period of the RFs, which decrease by 15% during the central 20 ms of the response. These results show a novel form of spatio-temporal inseparability in simple cells and are consistent with the notion of a coarse-to-fine processing of information in early visual cortex.

Action Potentials↗

Identification of proteins downregulated during the postnatal development of the cat visual cortex.

To identify proteins that play a role in the development of the mammalian visual cortex, we have used an immunosuppression and rapid immunization strategy to generate monoclonal antibodies to antigens that are present in area 17 of the cat during the peak of cortical plasticity but are downregulated near the end of the plastic period. We report here the immunohistochemical and immunobiochemical characterization of six monoclonal antibodies that identify antigens preferentially expressed in the cat visual cortex at 5 weeks of age. Monoclonal antibodies Cat-305 and Cat-306 detect three immunoreactive elements that are not present at birth but are present at 5 weeks. The majority of immunoreactivity is associated with a population of cells in the white matter that are absent at 15 weeks of age. At both 5 and 15 weeks, a very small number of neurons show intense immunoreactivity throughout all processes, resembling that achieved with a Golgi stain. In addition, a diffuse band of immunoreactivity in layer IV is largely restricted to cortical areas 17 and 18. Cat-307 recognizes a 150 kDa soluble protein present in small cytoplasmic inclusions. These cytoplasmic "dot" are present in all layers, but are most prominent in layer V. Cat-307 immunoreactivity is present at birth and is completely downregulated by 15 weeks. Cat-104 and Cat-105 recognize a 200 kDa insoluble protein present at birth and at 5 weeks, but markedly downregulated by 15 weeks. At birth, the white matter, subplate, and layer I are most densely labeled, while at 5 weeks labeling is densest in layers II, III, and V. Cat-402 recognizes a number of high-molecular-weight antigens that are differentially expressed at 5 and 15 weeks of age. Stained non-neuronal cells that resemble protoplasmic astrocytes are present in all layers at both 5 and 15 weeks. At 5 weeks, but not at birth or 15 weeks, darkly immunoreactive radial processes are observed that run through the full depth of the cortex. We show here that immunoreactivity for several different monoclonal antibodies is detected selectively during the period of maximal developmental plasticity. The results demonstrate that the cat visual cortex at 5 weeks of age is molecularly distinct from the cortex at 15 weeks.

Animals↗

[Neurons in the visual cortex of Microtus brandti].

Neurons were described in the visual Cortex of Microtus brandti, a Mongolian harmful rodent living in day-activity. We find following types of neurons in our Golgi-material: 1. spiny neurons: pyramidal and stellate neurons. 2. smooth or sparsely spined neurons: smooth, large neurons, sparsely spined small neurons with descending axons, sparsely spined neurons with ascending axons. Double-bouquet-, chandelier and neuroglioforme cells are not impregnated. There are no bipolare neurons (Martinotti cells) among the neurons with ascending axons. The small, sparsely spined neurons are not only in lamina IV - like in other species - but they can also be found in laminae II to IV. Their distribution of spines on the distal parts of dendrites seems to be characteristical for rodents. The lamination of the visual cortex of Microtus brandti is the same like in the rat. All cells are of large size in relation to the body mass of the animal.

Animals↗

Computations in the early visual cortex.

This paper reviews some of the recent neurophysiological studies that explore the variety of visual computations in the early visual cortex in relation to geometric inference, i.e. the inference of contours, surfaces and shapes. It attempts to draw connections between ideas from computational vision and findings from awake primate electrophysiology. In the classical feed-forward, modular view of visual processing, the early visual areas (LGN, V1 and V2) are modules that serve to extract local features, while higher extrastriate areas are responsible for shape inference and invariant object recognition. However, recent findings in primate early visual systems reveal that the computations in the early visual cortex are rather complex and dynamic, as well as interactive and plastic, subject to influence from global context, higher order perceptual inference, task requirement and behavioral experience. The evidence argues that the early visual cortex does not merely participate in the first stage of visual processing, but is involved in many levels of visual computation.

Animals↗

The effect of reversible cooling of cat's primary visual cortex on the responses of area 21a neurons.

1. Responses of sixty-four neurons in cortical area 21a were studied with areas 17 and 18 reversibly deactivated by cooling. From anatomical studies, most of area 21a input in the cat originates from these primary areas with no input from the A laminae of the lateral geniculate nucleus. 2. Both responses and spontaneous activity of all sixty-four area 21a neurons were markedly reduced when primary areas were cooled. In sixteen cells the responses were totally blocked. Temperatures of primary cortex required to produce total blockade varied between 25 and 4.5 degrees C. 3. The effect of cooling the primary visual cortex on the shape of orientation tuning curves was analysed in thirteen neurons from area 17 and in fifty-eight neurons from area 21a. In both areas, the width of the curve, when measured at its half-height, was preserved even when spike activity was reduced to below 10% of the original level. All neurons also retained their original directional preferences during cooling of the primary visual cortex. 4. The responsiveness of forty-seven neurons from area 21a was tested after rewarming of primary cortex. All but one neuron recovered their initial responsiveness within half an hour of restoring physiological temperature of primary visual cortex. 5. The results of the present study give an indication of the extent to which area 21a is sequentially related to the primary visual cortex in the processing of information.

Action Potentials↗

Postnatal development of dye-coupling among astrocytes in rat visual cortex.

Intercellular coupling among astrocytes was studied in rat visual cortex slices from animals aged 1 week to 4 months. Cell coupling via gap junctions was determined by the dye spread of the low molecular weight dye Lucifer Yellow CH injected into electrophysiologically identified cells to adjacent cells. Coupling among glial cells was first detected at postnatal day 11 and was thereafter consistently observed until adulthood. Dye spread was observed up to 300 microns radially from the injected cell covering multiple cortical layers. Following dye injection into a single cell up to several hundred Lucifer Yellow-positive cells could be observed. Quantitative analysis revealed a similar extent of dye spread at different developmental stages including a quite constant number of dye-coupled astrocytes from the end of the second postnatal week to adulthood. Double labelling of Lucifer Yellow-filled cells with an antiserum against the glial fibrillary acidic protein confirmed the astrocytic nature of the injected and coupled cells. Comparison of the density of dye-coupled cells in a given area and the total number of astrocytes as revealed by immunocytochemical staining suggests that dye-coupling includes the entire local astrocytic population. It is concluded that coupling among astrocytes via gap junctions in rat visual cortex occurs shortly after birth and reflects one of the first steps in astroglial maturation.

Animals↗