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Sex differences in cortical thickness and the dendritic tree in the monocular and binocular subfields of the rat visual cortex at weaning age.

The visual cortex of adult rats is sexually dimorphic at both the gross size and dendritic levels [Brain Res., 295 (1984) 27-34; J. Comp. Neurol., in press]. In addition, sex differences in the dendritic tree are dynamic and can be altered by environmental conditions imposed at weaning [Brain Res., 295 (1984) 27-34]. The present study examines sex differences in cortical thickness and in the dendritic tree of the monocular (Oc1M) and binocular (Oc1B) subfields in littermate male and female pairs of Long-Evans rats at weaning age (25 days). From Nissl-stained tissue, it was found that the whole cortex and layer II-IV of Oc1B was thicker in males than females. No sex differences were found in the thickness of Oc1M. Golgi-Cox-stained pyramidal neurons in layer III from the Oc1M and Oc1B regions were quantified in 7 littermate pairs of weaning-age rats. There were no sex differences in the basilar tree, while the apical oblique branches were sexually dimorphic, especially in the monocular region. Females had greater total dendritic length and longer terminal branches in Oc1M compared to males. Females also had longer bifurcating branches in both Oc1M and Oc1B than males. The present study found that sex differences at weaning age do not completely mirror the dimorphisms found in the visual cortex of the adult rat. This study also indicates that related subfields can differ in their morphology and should be examined separately.

Analysis of Variance↗

Effects of check size on visual cortex activation studied by functional magnetic resonance imaging.

We studied functional magnetic resonance imaging (fMRI) of visual cortex during checkerboard visual stimulation with three standard check sizes to examine whether activation in the visual cortex varied among these sizes. We acquired fMRI at 1.5 T in 8 normal subjects, each receiving the best refractive correction. Each subject underwent an experiment consisting of four conditions: black and white checkerboards with three check sizes (0.25-, 0.5-, and 1.0-degree) flickering at 8 Hz, and a black screen. SPM96 was used for a group data analysis with a random effects model after each of the subject's data was motion-corrected and spatially normalized to a standard brain. The activation in the visual cortex showed the greatest signal changes with the 0.5-degree check among the three check sizes. When standard check sizes are used to stimulate visual cortex in fMRI experiments, our results suggest that 0.5-degree checks flickering at 8 Hz produce the most vigorous activation in visual cortex.

Adult↗

Orientation selectivity of synaptic potentials in neurons of cat primary visual cortex.

Neurons of the visual cortex of the cat were penetrated with intracellular electrodes and postsynaptic potentials evoked by visual stimuli recorded. By alternately polarizing the cell with steady current injected through the recording electrode, IPSPs and EPSPs could be recorded and analyzed independently. Hyperpolarizing current suppressed IPSPs and enhanced EPSPs by moving the membrane potential toward the IPSP equilibrium potential. Depolarizing the cell toward the EPSP equilibrium potential enhanced IPSP. The responses to electrical stimulation of the LGN, where EPSPs and IPSPs could be distinguished easily by virtue of their characteristic latencies and shapes, were used to set the current injection to the appropriate level to view the two types of synaptic potential. EPSPs were found to be well oriented in that maximal depolarizing responses could be evoked at only one stimulus orientation; rotating the stimulus orientation in either direction produced a fall in the EPSP response. IPSPs were also well tuned to orientation, and invariably the preferred orientations of EPSPs and IPSPs in any one cell were identical. In addition, no systematic difference in the width of tuning of the two types of potential was seen. This result has been obtained from penetrations of over 30 cortical cells, including those with simple and complex receptive fields. It is concluded that orientation of cortical receptive fields is neither created nor sharpened by inhibition between neurons with different orientation preference. The function of inhibition evoked simultaneously with excitation by optimally oriented stimuli has yet to be determined, though it is likely to be the mechanism underlying other cortical receptive field properties, such as direction selectivity and end-stopping.

Animals↗

The human visual cortex.

The discovery and analysis of cortical visual areas is a major accomplishment of visual neuroscience. In the past decade the use of noninvasive functional imaging, particularly functional magnetic resonance imaging (fMRI), has dramatically increased our detailed knowledge of the functional organization of the human visual cortex and its relation to visual perception. The fMRI method offers a major advantage over other techniques applied in neuroscience by providing a large-scale neuroanatomical perspective that stems from its ability to image the entire brain essentially at once. This bird's eye view has the potential to reveal large-scale principles within the very complex plethora of visual areas. Thus, it could arrange the entire constellation of human visual areas in a unified functional organizational framework. Here we review recent findings and methods employed to uncover the functional properties of the human visual cortex focusing on two themes: functional specialization and hierarchical processing.

Animals↗

Functional compensation in the lateral suprasylvian visual area following bilateral visual cortex damage in kittens.

Previous studies have shown that functional compensation is present in the cat's posteromedial lateral suprasylvian (PMLS) area of cortex after damage to areas 17, 18, and 19 (visual cortex) early in life but not after damage in adults. These studies all have investigated animals with a unilateral visual cortex lesion, whereas all behavioral studies of compensation for early visual cortex damage have investigated animals with a bilateral lesion. In the present experiment, we investigated whether functional compensation also is present in PMLS cortex after a bilateral visual cortex lesion early in life. We recorded from single neurons in the PMLS cortex of adult cats that had received a bilateral lesion of areas 17, 18, and 19 on the day of birth or at 8 weeks of age. We found that PMLS cells in both groups of cats had functional compensation (normal direction selectivity and ocular dominance) similar to that seen after a unilateral lesion at the same ages. These results are consistent with the hypothesis that PMLS cortex is involved in the behavioral compensation seen after early visual cortex damage. In addition, the results indicate that inputs from contralateral visual cortex are not necessary for the development of functional compensation seen in PMLS cortex.

Age Factors↗

Linearity and normalization in simple cells of the macaque primary visual cortex.

Simple cells in the primary visual cortex often appear to compute a weighted sum of the light intensity distribution of the visual stimuli that fall on their receptive fields. A linear model of these cells has the advantage of simplicity and captures a number of basic aspects of cell function. It, however, fails to account for important response nonlinearities, such as the decrease in response gain and latency observed at high contrasts and the effects of masking by stimuli that fail to elicit responses when presented alone. To account for these nonlinearities we have proposed a normalization model, which extends the linear model to include mutual shunting inhibition among a large number of cortical cells. Shunting inhibition is divisive, and its effect in the model is to normalize the linear responses by a measure of stimulus energy. To test this model we performed extracellular recordings of simple cells in the primary visual cortex of anesthetized macaques. We presented large stimulus sets consisting of (1) drifting gratings of various orientations and spatiotemporal frequencies; (2) plaids composed of two drifting gratings; and (3) gratings masked by full-screen spatiotemporal white noise. We derived expressions for the model predictions and fitted them to the physiological data. Our results support the normalization model, which accounts for both the linear and the nonlinear properties of the cells. An alternative model, in which the linear responses are subject to a compressive nonlinearity, did not perform nearly as well.

Animals↗

Non-uniform distribution of the NMDAR1 receptor subunit in kitten visual cortex at the peak of the critical period.

PURPOSE: The development of columnar systems in the visual cortex, in particular ocular dominance columns, is dependent on experiential activity in conjunction with NMDA-mediated plasticity mechanisms. Recent experiments, however, have shown that certain aspects of the columnar organization of the visual cortex, such as the spacing of columns, are not changed by manipulations that affect the pattern of retinal activity. This raises the possibility that features intrinsic to the visual cortex may play a crucial role in the development of cortical columns and that a non-uniform distribution of NMDA receptors in the developing visual cortex could form the link between activity and intrinsic cortical modularity. METHODS: To examine this possibility we used immunohistochemical techniques to label the NMDAR1 receptor subunit protein in kitten visual cortex. The arrangement of the NMDAR1 subunit was visualized (using a monoclonal antibody) in flattened and coronal sections through visual cortex. The tangential and laminar distributions of NMDAR1 immunoreactivity (NMDAR1ir) were studied at the peak of the critical period for plasticity (4-5 weeks of age) in the developing kitten visual cortex. RESULTS: At the ages examined there was a non-uniform distribution of NMDAR1 immunoreactivity in the visual cortex. These patches of darker NMDAR1 label were found in layers 2/3 and extended up into layer 1. Thus, during development neurons expressing the NMDAR1 receptor subunit were distributed in a patchy fashion in the upper layers of the kitten visual cortex. CONCLUSIONS: This suggests that NMDA-mediated activity-dependent plasticity may not occur uniformly across the tangential extent of the visual cortex, and raises the possibility that the arrangement of NMDAR1 patches may guide the emergence of nascent columns in the developing visual cortex.

Animals↗

GABA-inactivation attenuates colinear facilitation in cat primary visual cortex.

Neurons in primary visual cortex (V1) respond preferentially to stimuli of a particular orientation falling within a circumscribed region of visual space known as their receptive field (RF). However, the response to an optimally oriented stimulus presented within the RF can be enhanced by the simultaneous presentation of co-oriented, co-linearly aligned flank stimuli falling outside the RF which, when presented alone, fail to activate the cell. This type of contextual effect, termed colinear facilitation, presumably forms the physiological substrate for the integration of the line elements of a contour and the perceptual saliency of a contour in a complex environment. Here we show that colinear facilitation in single cells of cat area V1 can be substantially reduced or abolished by focal inactivation of laterally remote cells in the same area which respond strongly to the co-oriented, colinear flank stimulus inducing the facilitatory effect. The results provide evidence that horizontal intrinsic connections between cells with co-oriented and co-linearly aligned RFs make a major contribution to colinear facilitation in V1. They imply that the neuronal circuitry underlying contour integration and saliency is already present at the earliest stage of visual cortical information processing.

Action Potentials↗

Relation of cortical cell orientation selectivity to alignment of receptive fields of the geniculocortical afferents that arborize within a single orientation column in ferret visual cortex.

Neurons in the primary visual cortex of higher mammals are arranged in columns, and the neurons in each column respond best to light-dark borders of particular orientations. The basis of cortical cell orientation selectivity is not known. One possible mechanism would be for cortical cells to receive input from several lateral geniculate nucleus (LGN) neurons with receptive fields that are aligned in the visual field (Hubel and Wiesel, 1962). We have investigated the relationship between the arrangement of the receptive fields of geniculocortical afferents and the orientation preferences of cortical cells in the orientation columns to which the afferents provide visual input. Radial microelectrode penetrations were made into primary visual cortex of anesthetized adult sable ferrets. Cortical cells were recorded throughout the depth of the cortex, and their orientation preferences were determined. Cortical cell responses were then eliminated by superfusion of the cortex with either kainic acid (Zahs and Stryker, 1988) or muscimol. After the drug treatment, responses from many single units with distinct receptive fields were recorded. These responses were presumed to be those of geniculocortical afferents, because they had the response properties characteristic of LGN neurons, and because they could be recorded only in cortical layers that receive geniculate input. In 16 of 18 cases, the afferent receptive fields recorded in a single penetration covered an elongated region of visual space. In these penetrations, the best-fit line through the centers of the afferent receptive fields generally paralleled the preferred orientation of cortical cells recorded at the same site in cortex. These results are consistent with the Hubel and Wiesel (1962) model for the construction of oriented visual cortical receptive fields from geniculate inputs with aligned receptive fields.

Afferent Pathways↗

Synaptic development in the rabbit superior colliculus and visual cortex.

The development of synapses in the visual cortex (VC) and superior colliculus (SC) of the rabbit has been examined with the electron microscope. In both areas, the number of synapses reaches adult levels by 20--25 days of postnatal age, but the development in the visual cortex is delayed in comparison to that in the superior colliculus. When S synapses (spheroidal vesicles, asymmetric thickening) are compared with F synapses (flattened vesicles, symmetric thickening), even greater differences are seen. In both the VC and SC, S synapses develop earlier than F synapses, though there is considerable overlap. Of interest is that fact that synapses in the visual cortex seem to overshoot their adult levels late in development, suggesting that an excess of synapses may be formed in this system. Multiple synapses, probably of retinal origin, increase in the first 3 weeks of synaptic development in the SC, but never are present in significant proportions in the VC. Synapse formation most often is characterized by formation of a junction and a postsynaptic thickening, followed by acquisition of synaptic vesicles. After 15 days, there is only a small number of such "non-vesicle synapses" in either the SC or VC.

Age Factors↗

'Real-motion' cells in area V3A of macaque visual cortex.

The stability of visual perception despite eye movements suggests the existence, in the visual system, of neural elements able to recognize whether a movement of an image occurring in a particular part of the retina is the consequence of an actual movement that occurred in the visual field, or self-induced by an ocular movement while the object was still in the field of view. Recordings from single neurons in area V3A of awake macaque monkeys were made to check the existence of such a type of neurons (called 'real-motion' cells; see Galletti et al. 1984, 1988) in this prestriate area of the visual cortex. A total of 119 neurons were recorded from area V3A. They were highly sensitive to the orientation of the visual stimuli, being on average more sensitive than V1 and V2 neurons. Almost all of them were sensitive to a large range of velocities of stimulus movement and about one half to the direction of it. In order to assess whether they gave different responses to the movement of a stimulus and to that of its retinal image alone (self-induced by an eye movement while the stimulus was still), a comparison was made between neuronal responses obtained when a moving stimulus swept a stationary receptive field (during steady fixation) and when a moving receptive field swept a stationary stimulus (during tracking eye movement). The receptive field stimulation at retinal level was physically the same in both cases, but only in the first was there actual movement of the visual stimulus. Control trials, where the monkeys performed tracking eye movements without any intentional receptive field stimulation, were also carried out. For a number of neurons, the test was repeated in darkness and against a textured visual background. Eighty-seven neurons were fully studied to assess whether they were real-motion cells. About 48% of them (42/87) showed significant differences between responses to stimulus versus eye movement. The great majority of these cells (36/42) were real-motion cells, in that they showed a weaker response to visual stimulation during tracking than to the actual stimulus movement during steady fixation. On average, the reduction in visual response during eye movement was 64.0 +/- 15.7% (SD).(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗

An explanation of contextual modulation by short-range isotropic connections and orientation map geometry in the primary visual cortex.

Recent experimental studies on the primary visual cortex have revealed complicated nonclassical neuronal activities. Contextual modulation on orientation-contrast is one typical example of nonclassical neuronal behavior. This modulation by surrounding stimuli in a nonclassical receptive field is mainly thought to be mediated by short- and long-range horizontal connections within the primary visual cortex. Short-range connections are circularly symmetrical and relatively independent of orientation preferences, while long-range connections are patchy, asymmetrical, and orientation specific. Although this modulation can be explained by long-range specific connections qualitatively, recent studies suggest that long-range connections alone may be insufficient with respect to the balance between two types of connections. Here, in order to clarify the role of short-range connections in the process of contextual modulation, we propose a model of the primary visual cortex with isotropic short-range connections and a geometric orientation map. Computational simulations using the model have demonstrated that contextual modulation can be explained by short-range connections alone. This is due to the interaction between the spatial periodicity of orientation domains and the excitatory-inhibitory regions arising from the propagation of activities.

Action Potentials↗

Intracortical connections are not required for oscillatory activity in the visual cortex.

Synchronized oscillatory discharge in the visual cortex has been proposed to underlie the linking of retinotopically disparate features into perceptually coherent objects. These proposals have largely relied on the premise that the oscillations arise from intracortical circuitry. However, strong oscillations within both the retina and the lateral geniculate nucleus (LGN) have been reported recently. To evaluate the possibility that cortical oscillations arise from peripheral pathways, we have developed two plausible models of single cell oscillatory discharge that specifically exclude intracortical networks. In the first model, cortical oscillatory discharge near 50 Hz in frequency arises from the integration of signals from strongly oscillatory cells within the LGN. The model also predicts the incidence of 50-Hz oscillatory cells within the cortex. Oscillatory discharge around 30 Hz is explained in a second model by the presence of intrinsically oscillatory cells within cortical layer 5. Both models generate spike trains whose power spectra and mean firing rates are in close agreement with experimental observations of simple and complex cells. Considered together, the two models can largely account for the nature and incidence of oscillatory discharge in the cat's visual cortex. The validity of these models is consistent with the possibility that oscillations are generated independently of intracortical interactions. Because these models rely on intrinsic stimulus-independent oscillators within the retina and cortex, the results further suggest that oscillatory activity within the cortex is not necessarily associated with the processing of high-order visual information.

Action Potentials↗

Intracortical connections are not required for oscillatory activity in the visual cortex.

Synchronized oscillatory discharge in the visual cortex has been proposed to underlie the linking of retinotopically disparate features into perceptually coherent objects. These proposals have largely relied on the premise that the oscillations arise from intracortical circuitry. However, strong oscillations within both the retina and the lateral geniculate nucleus (LGN) have been reported recently. To evaluate the possibility that cortical oscillations arise from peripheral pathways, we have developed two plausible models of single cell oscillatory discharge that specifically exclude intracortical networks. In the first model, cortical oscillatory discharge near 50 Hz in frequency arises from the integration of signals from strongly oscillatory cells within the LGN. The model also predicts the incidence of 50-Hz oscillatory cells within the cortex. Oscillatory discharge around 30 Hz is explained in a second model by the presence of intrinsically oscillatory cells within cortical layer 5. Both models generate spike trains whose power spectra and mean firing rates are in close agreement with experimental observations of simple and complex cells. Considered together, the two models can largely account for the nature and incidence of oscillatory discharge in the cat's visual cortex. The validity of these models is consistent with the possibility that oscillations are generated independently of intracortical interactions. Because these models rely on intrinsic stimulus-independent oscillators within the retina and cortex, the results further suggest that oscillatory activity within the cortex is not necessarily associated with the processing of high-order visual information.

Animals↗

Oscillatory and non-oscillatory synchronizations in the visual cortex and their possible roles in associations of visual features.

It was postulated that the perceived association of visual features is based on the synchronization of those neural signals that are activated by a coherent visual object. Two types of synchronized cortical signals were found by us in cat and monkey visual cortex, and were proposed as candidates for feature association: (1) stimulus-locked signals, evoked by transient retinal stimulation, and typically non-rhythmic; (2) oscillatory signals, induced by sustained stimuli, and typically not locked in their oscillation phases to stimulus events. Both types of signals can occur synchronously in those neurons that are activated by a common stimulus. Synchronized activities were found in paired recordings within vertical cortex columns, in separate columns of the same cortical area, and even between different cortical areas or hemispheres. The average phase difference between such common oscillatory events was typically close to zero (< 1 msec mean +/- 2 msec S.D.). For the dependence of synchronization from stimulus and receptive field properties, a preliminary 'rule' can be given: the coherence of fast oscillations in separate cortical assemblies depends inversely on the 'coding distance' between the assemblies' RF properties, but directly on the degree of overlap between the assemblies' respective coding properties and the features of a common stimulus. This means that oscillatory events in any two assemblies, in the same or in different cortical areas or hemispheres, are more closely correlated the more similar are their receptive field properties, and the better a common stimulus activates the assemblies simultaneously. Our results can explain some neural mechanisms of perceptual feature-linking, including mutual enhancement among similar, spatially and temporally dispersed features, definitions of spatial and temporal continuity, scene segmentation, and figure-ground discrimination. We further propose that mutual enhancement and synchronization of cell activities are general principles of temporal coding by assemblies, that are also used within and among other sensory modalities as well as between cortical sensory and motor systems.

Animals↗

Ocular dominance peaks at pinwheel center singularities of the orientation map in cat visual cortex.

In the primary visual cortex of monkey and cat, ocular dominance and orientation are represented continuously and simultaneously, so that most neighboring neurons respond optimally to visual stimulation of the same eye and orientation. Maps of stimulus orientation are punctuated by singularities referred to as "pinwheel centers," around which all orientations are represented. Given that the orientation map is mostly continuous, orientation singularities are a mathematical necessity unless the map consists of perfectly parallel rows, and there is no evidence that the singularities play a role in normal function or development. We report here that in cats there is a strong tendency for peaks of ocular dominance to lie on the pinwheel center singularities of the orientation map. This relationship predicts but is not predicted by the tendencies, previously reported, for pinwheels to lie near the center lines of ocular dominance bands and for iso-orientation bands to cross ocular dominance boundaries at right angles. The coincidence of ocular dominance peaks with orientation singularities is likely to reflect a strong underlying functional link between the two visual cortical maps.

Animals↗

Activation of visual cortex in REM sleep measured by 24-channel NIRS imaging.

To visualize dreaming brain functions we studied hemodynamic changes in the visual cortex during the transition from non-rapid eye movement (NREM) to rapid eye movement (REM) sleep, using a 24-channel Near-Infrared Spectroscopy (NIRS) imaging method. Results were compared to the activation in visual cortex by visual stimulation during wakefulness. Subjects were four healthy males between 25 and 49 years of age. Five all-night polysomnographic and NIRS recordings were made. Increases in the oxygenated hemoglobin concentration in visual cortex were observed from nine of 14 REM periods. The activated areas were broader during REM sleep than during visual stimulation. These findings suggest that activation of visual cortex in REM sleep might represent dream-related brain activity.

Adult↗

Excitatory synaptic inputs to spiny stellate cells in cat visual cortex.

In layer 4 of cat visual cortex, the monocular, concentric receptive fields of thalamic neurons, which relay retinal input to the cortex, are transformed into 'simple' cortical receptive fields that are binocular and selective for the precise orientation, direction of motion, and size of the visual stimulus. These properties are thought to arise from the pattern of connections from thalamic neurons, although anatomical studies show that most excitatory inputs to layer 4 simple cells are from recurrently connected circuits of cortical neurons. We examined single fibre inputs to spiny stellate neurons. We examined single fibre inputs to spiny stellate neurons in slices of cat visual cortex, and conclude that thalamocortical synapses are powerful and the responses they evoke are unusually invariant for central synapses. However, the responses to intracortical inputs, although less invariant, are strong enough to provide most of the excitation to simple cells in vivo. Our results suggest that the recurrent excitatory circuits of cortex may amplify the initial feedforward thalamic signal, subserving dynamic modifications of the functional properties of cortical neurons.

Animals↗