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Vernier acuity of neurones in cat visual cortex.

The ability of human observers to detect Vernier breaks of as little as 5 s arc has been termed hyperacuity as this distance is substantially less than the angular separation of the bars of the highest spatial frequency of grating (approximately 1 arc min) that can be detected. Although the visual cortex is a likely candidate for the location of detectors involved in this performance, it is not known whether there are cells sensitive enough to detect deviations from co-linearity that are small compared with their spatial resolution (defined in terms of the highest spatial frequency that the cell can detect). We report here the results of physiological experiments on single units in area 17 of the cat visual cortex in which we studied the effect of introducing a Vernier break into a bar stimulus moved across the receptive field of the cell at a constant velocity. Our results show that the responses of most simple and complex cells are significantly reduced by the introduction of a Vernier break that is substantially smaller than the spatial resolution of the cell. The most sensitive cells in our sample could discriminate Vernier offsets of 3-6 arc min with a reliability of approximately 70%. This was much smaller than their spatial resolution, which was in the range 25-30 arc min. We interpret these results in terms of mechanisms that could underly the orientation selectivity of cortical neurones and suggest how our results relate to human Vernier acuity.

Animals↗

Molecular differences among neurons reveal an organization of human visual cortex.

Monoclonal antibody Cat-301 recognizes a cell-surface proteoglycan on subsets of neurons in several areas of the cat and macaque monkey central nervous system. In striate and extrastriate visual cortex of the macaque, the distribution of Cat-301-positive neurons demonstrates features of cellular organization that correlate with previously described functional subdivisions. Here we show that Cat-301 recognizes an antigen in human cortex that is closely related, if not identical, to the antigen in laboratory animals. Further, we use Cat-301 to demonstrate an organization of molecularly defined neurons in primary and secondary visual cortex (cortical areas V1 and V2) of the human. The organization demonstrated with Cat-301 in human area V1 correlates with the organization of ocular dominance columns demonstrated by cytochrome oxidase histochemistry. The organization demonstrated with Cat-301 in human area V2 correlates with the thick stripes of the cytochrome oxidase pattern. The present observations provide evidence for a visual pathway in human cortex homologous to the magnocellular pathway in macaque, a pathway involved in processing the low-contrast, achromatic, and moving components of visual stimuli.

Animals↗

Functional segregation of color and motion processing in the human visual cortex: clinical evidence.

Anatomical and physiological investigations indicate two major distinct functional streams within the extrastriate visual cortex of the macaque monkey, and behavioral observations suggest that the ventral (occipitotemporal) pathway is the cornerstone for object recognition whereas the dorsal (occipitoparietal) pathway is primarily involved in visuospatial perception and visuomotor performance. In the context of this dichotomy we conducted a psychophysical and neuropsychological study of visual perceptual abilities in two stroke patients, each with lesions involving several extrastriate areas. Magnetic resonance imaging demonstrated bilateral lesions; in one patient (E.W.) the lesion involves the ventral medial portions of the occipital and temporal lobes, and in the other (A.F.) the lesion involves dorsally the occipital-parietal area, including the region of the temporal-parietal-occipital junction. E.W. suffers from achromatopsia of central origin, prosopagnosia, visual agnosia, and alexia without agraphia. His depth and motion perception, including recognition of moving objects, are normal. He has superior visual field loss bilaterally, and slightly impaired acuity, and complains that the world appears in a deep twilight even on a sunny day. In contrast, A.F. shows specific deficits of stereopsis, spatial localization, and several aspects of motion perception. He is also impaired at recognizing objects presented from unconventional views, but recognition of prototypical views of objects, and color and form discrimination are normal, as is his ability to recognize faces. The anatomical characteristics of the lesions of these two patients permit a direct experimental comparison of the effects of lesions confined to the parietal or temporal pathways. E.W.'s and A.F.'s performance on the psychophysical and neuropsychological tasks discussed here supports the functional distinction between a dorsal and a ventral extrastriate system but additionally suggests the existence of a pathway involved in identification-from-motion that is separate from both the dorsal early motion/spatial analysis pathway and the ventral color/static-form pathway.

Cerebrovascular Disorders↗

Exposure to lines of only one orientation modifies dendritic morphology of cells in the visual cortex of the cat.

To determine whether selective exposure to lines of one orientation modifies the shape of the dendritic fields of cells in visual cortex, we examined the dendritic morphology of neurons in area 17 of five normally reared cats, five cats reared viewing only vertical lines, and three cats reared viewing only horizontal lines. Kittens were placed with their mothers into a totally dark room before their eyes had opened. Beginning at 4 weeks of age, the kittens were brought out for daily periods of exposure wearing masks that limited the vision of each eye to a field of three vertical lines or three horizontal lines. After a minimum of 170 hours of exposure, the animals were killed and blocks of visual cortex were impregnated by the Golgi-Kopsch procedure and cut tangential to the pial surface. Complete neurons from layers III and IV were drawn with the aid of a camera lucida, and the orientations of the dendritic fields wer analyzed using Sholl diagrams. In normal cats, the distributions of the orientations of dendritic fields were uniform, whereas in strip-reared cats, the distributions for the layer III pyramidal cells were shifted. The direction of this shift varied with the experience of the cat: In cats reared viewing only vertical lines, the dendritic fields were oriented orthogonal to the representation of the vertical meridian, and in cats reared viewing only horizontal lines, the fields were oriented parallel to the representation of the vertical meridian. In contrast, the distribution of dentritic orientations for the stellate cells was not affected by stripe-rearing. These results demonstrate a morphological effect of early visual experience that is specific to the particular stimulus presented during rearing and suggest that (1) cortical cells differ in the degree to which they can be modified by such experience, and (2) the dendritic morphology of cortical neurons is related to their preferred orientations.

Animals↗

Comparison of the selectivity of postsynaptic potentials and spike responses in cat visual cortex.

Intracellular recordings were made from neurons in the cat visual cortex (area 17) to compare the orientation and direction selectivities of the output of a cell with those of the input the cell receives. The input to a cell was estimated from the PSPs (postsynaptic potentials) evoked by visual stimulation, and the output estimated from the number of spikes generated during the same responses. For the whole sample, selectivity of the output of cells was significantly higher than selectivity of their input. Upon PSP to spike transformation, the selectivity index was, on average, doubled. However, the degree of the selectivity improvement in individual cells was very different, varying from cases in which highly selective output was created from a poorly selective input and thus selectivity was greatly improved, to little or no improvement in other neurons. The improvement of selectivity was not correlated with resting membrane potential, threshold for action potential generation, background discharge rate or amplitude of optimal PSP response. Further, no systematic difference was found between simple and complex cells in the input-output relations, indicating that the 'tip of the iceberg' effect on shaping the response selectivity was cell specific, but not cell type specific. This supports the notion that multiple mechanisms are responsible for generation of the response selectivity, and that the contribution of any particular mechanism may vary from one cell to the other. The heterogeneity of the input-output relations in visual cortical cells could indicate different functions of cells in the cortical network; some cells are creating selectivity de novo, the function of other neurons probably being repetition and amplification of the selected signal and arrangement of the output of a whole column.

Action Potentials↗

Coincidental appearance of the alpha 1 subunit of the GABA-A receptor and the type I benzodiazepine receptor near birth in macaque monkey visual cortex.

The expression of subtypes of the GABA-A/benzodiazepine receptor complex has been studied during pre- and postnatal development of Macaca monkey visual cortex using complementary radioligand and immunocytochemical labeling. Type I benzodiazepine receptors were labeled directly by [3H]zolpidem. Type II receptors were determined by the amount of binding for [3H]flunitrazepam (FZ) persisting in the presence of the type I-specific ligand CL218872. Monoclonal antibody bd24 was used to label alpha 1 subunits and bd17 to label beta 2 and beta 3 subunits of the GABA-A receptor. Radioligand binding data and bd17 immunoreactivity indicated that type II benzodiazepine receptors were present by fetal day (Fd) 74 (44% of gestation). Immunoreactivity for the beta 2/beta 3 subunits increased until 3-6 weeks after birth, and then declined somewhat into adulthood. Neither radioligand labeling for type I receptors nor immunocytochemical staining for the alpha 1 subunit were apparent until mid-gestation. Both markers appeared shortly before birth in layer 4C, and then in other cortical layers after birth. Immunoreactivity for the alpha 1 subunit increased steadily after birth until it became more intense than that for beta 2/3 subunits in the adult. Quantitative densitometry of CL218872 competition for [3H]FZ binding showed that type I/II distribution was 22%/78% at Fd103; 42%/58% at Fd131; 67%/33% at 9 months; and 61%/39% in adult visual cortex. This "switch" between benzodiazepine receptor subtypes overlaps the postnatal critical period for geniculostriate development, suggesting that the change from type II to type I receptors and the appearance of alpha 1 subunits may play a decisive role in the maturation of geniculocortical axon terminations and cortical response properties. It remains to be shown whether this "switch" is dependent on functional visual input.

Animals↗

Oligodendrocytes differentiate in organotypic cultures of rat visual cortex and myelinate efferent axons.

We have investigated the presence and function of glia cells, especially of oligodendrocytes (OL) in organotypic cultures of rat visual cortex grown for 1-6 weeks in vitro. OL identified by strong Galactocerebroside-immunoreactivity (GalC-ir) displayed rather small somata and elaborately ramified processes. They were most concentrated in layers VIa and VIb and the remnant of the white matter. Silver staining revealed long descending or oblique processes in layers V and VI, which were often arranged in patches, and horizontal processes in the white matter. Proximal processes of OL cell bodies were connected to these long processes. DiI-labeling revealed very similar patches of processes, termed OL domains. They were identified as membraneous sheaths formed by processes of single OL around axons passing the OL domain. Confocal microscopy revealed single axons running through the membrane sheaths. We compared the molecular differentiation of glial cells in cultures to the in vivo situation with protein blots and immunohistochemistry for glial cell marker molecules. In homogenates of visual cortex in vivo, protein blots revealed the increase in expression by OL of myelin basic protein (MBP) during the fourth postnatal week. The astrocytic marker glial fibrillary acidic protein (GFAP), blotted as a control, increased over time in vivo, beginning at P14, indicating the differentiation of astrocytes. In homogenates of organotypic cortex cultures, the times course of expression of GFAP was very similar: it increased dramatically during the first 10 DIV, and remained fairly constant in older cultures.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Responses of neurons in primary visual cortex are modulated by eye position.

1. We tested the effects of eye position on the visual excitability of 88 neurons in the primary visual cortex of awake cats trained in oculomotor tasks. For most cells, we examined responses evoked by retinotopically identical stimuli for centered gaze, 8 degrees to the left of center, and 8 degrees to the right of center. 2. An effect of eye position was observed for 40% of the cells. For 13%, responsiveness varied by a factor of 2 or more. Most commonly, response was maximal with gaze shifted to one side, minimal when shifted to the opposite side, and intermediate for centered fixation. The exceptions were four cells for which excitability varied symmetrically with fixations to either side of center. 3. Variability in excitability associated with eye position is a wide-spread phenomenon, having been observed in the lateral geniculate nucleus, V1, and extrastriate cortex. These results are consistent with the belief that such variability is utilized in constructing a head-centered frame of reference from a retinotopic input.

Animals↗

Development and regulation of alpha adrenoceptors in kitten visual cortex.

Alpha-1 and alpha-2 adrenergic receptors were localized in developing cat visual cortex by using [3H]prazosin and [3H]rauwolscine, respectively as selective ligands. The effects of neuronal input on the development of the two receptor subtypes were also studied in animals with lesions at various sites within the central visual pathways. Binding densities for both ligands increased during the first few postnatal weeks and declined thereafter. For both receptor subtypes, the highest concentration of binding sites was found in the subplate zone of the cortex in neonatal animals. Both ligands showed their highest concentrations in cortical layer IV beginning at postnatal day 30 and in the superficial cortical layers in adulthood. However, the developmental redistribution of alpha-1 receptors began at earlier ages than that of the alpha-2 sites. The alpha-1 sites were still concentrated in the subplate zone up to 60 days postnatal, while the alpha-2 sites in this region disappeared much earlier. Receptor binding densities were also examined in animals with quinolinic acid lesions within cortex, lesions of the lateral geniculate nucleus and lesions of the optic tract. The results indicate that both alpha-adrenoceptor subtypes were mainly located on cortical cells, and that the absence of neuronal activity during development resulted in a reduction of the binding density for both subtypes in the visual cortex. An additional major reduction in alpha-2 but not alpha-1 binding sites was observed following the lateral geniculate nucleus lesion, suggesting that the development of alpha-2 receptors is also dependent on input from the lateral geniculate nucleus. Removal of the lateral geniculate nucleus early in life resulted in a significant increase in alpha-1 receptors in the subplate region, indicating that receptor densities in this zone may be negatively regulated by the lateral geniculate nucleus afferents. These results show that adrenergic receptors reorganize during postnatal cortical development with a strong temporary concentration in the subplate zone. The reorganization process is heavily influenced by cortical inputs.

Animals↗

Myelinated axons and the pyramidal cell modules in monkey primary visual cortex.

In addition to the horizontal bands of myelinated axons that produce the line of Gennari and the inner band of Baillarger, the macaque primary visual cortex contains prominent vertical bundles of myelinated axons. In tangential sections through layer IVC, these axon bundles are regularly arranged. They have a mean center-to-center spacing of about 23 microns, and each one contains an average of 34 (S.D. +/- 13) myelinated axons. These bundles seem to be largely composed of efferent fibers, because in material in which pyramidal cells have been labelled in layer II/III and in layers IVA and IVB the axons of these neurons descend towards the white matter in bundles. However, it is doubtful whether all of the descending myelinated axons from the superficial layers emerge from the cortex, since counts show that the bundles contain maximum numbers of myelinated axons at the level of layer IVC, and that in layers V and VI their number is reduced by about 30%. Perhaps some of the axons enter the line of Baillarger, in layer V. When the bundles of myelinated axons and the clusters of apical dendrites of the layer V pyramidal cells are visualized simultaneously within layer IVC in electron microscopic preparations, it is apparent that their center-to-center spacing is similar, namely, about 23 microns and that a bundle of axons has a cluster of apical dendrites lying adjacent to it. Because of this association, and because axons from layer III pyramidal cells have been shown to enter the bundles, it is suggested that the myelinated axon bundles contain the efferent axons from the projection neurons in the individual pyramidal cell modules. However, in addition to the myelinated axons, the bundles contain unmyelinated axons, so that they also probably serve as the conduits for axons forming connections between layers. It is proposed that the pyramidal cell modules are the basic, functional neuronal units of the visual cortex, and since the neurons within a particular module can be expected to have slightly different inputs and response properties from those in neighboring modules, the individual axon bundles that emerge from each module would be expected to carry a unique set of efferent information.

Animals↗

Direct mapping of ocular dominance columns in human primary visual cortex.

Functional magnetic resonance imaging at 2.0T was employed to identify columnar structures in human visual cortex. Sagittal sections (4 mm thickness) covering the calcarine cortex were acquired with use of a multiecho low flip angle gradient-echo sequence at 4.0 s temporal resolution and 0.25 x 0.25 mm2 spatial resolution. Extending earlier attempts based on a differential paradigm contrasting left vs right eye stimulation, this work presents the first direct mapping of human ocular dominance columns by measuring separate activation maps with left and right eye stimulation. The resulting individual maps reveal patterns of ocular dominance as spots or bands of altered activity in calcarine cortex. Their superposition shows only little spatial overlap of eye-specific encoding which strongly supports the genuineness of these functional units.

Adult↗

Recovery of a temporally based visual discrimination after visual cortex lesion in the rat.

In Expt. 1, rats were conditioned to emit a shock avoidance response when the pulse rate of a light was increased. Then, after bilateral visual cortex lesions, the rats were given 10, 20, or 40 days recovery before relearning the discrimination. While all rats were able to relearn the discrimination response, lesion rats had a performance deficit after all recovery periods. Expt. 2 compared the effects of postoperative visual pulse rate training to those of auditory pulse rate training on relearning of the photic pulse-rate discrimination 10 days after visual decortication. Recovery of discrimination responding was better after auditory pulse rate training than after visual training. These data suggest that visual cortex lesions in the rat disrupt perceptual or associational functions involving the temporal features of a visual stimulus. In addition, generalization of relational properties during cross-modal training through multimodal CNS structures appears to enhance recovery of behavior after brain insult.

Acoustic Stimulation↗

Stereoscopic mechanisms in monkey visual cortex: binocular correlation and disparity selectivity.

The neural signals in visual cortex associated with positional disparity and contrast texture correlation of binocular images are the subject of this study. We have analyzed the effects of stereoscopically presented luminous bars and of dynamic random-dot patterns on the activity of single neurons in cortical visual areas V1, V2, and V3-V3A of the alert, visually trained rhesus macaque. The interpretation of the results and considerations of possible neural mechanisms led us to recognize 2 functional sets of stereoscopic neurons. (1) A set of neurons, tuned excitatory (T0) or tuned inhibitory (TI), which respond sharply to images of zero or near-zero disparity. Objects at or about the horopter drive the T0 neurons and suppress the TI, while objects nearer and farther have the opposite effects on each type, inhibition of the T0 and excitation of the TI. The activity of these neurons may provide, in a reciprocal way, the definition of the plane of fixation, and the basic reference for binocular single vision and depth discrimination. (2) A second set of neurons includes tuned excitatory at larger crossed or uncrossed disparities (TN/TF) and neurons with reciprocal excitatory and inhibitory disparity sensitivity with cross-over at the horopter (NE/FA). Binocularly uncorrelated image contrast drives these neurons to a maintained level of activity, which shifts, in response to correlated images, toward facilitation or suppression as a function of positional disparity. These neurons may operate in the neural processing leading to stereopsis, both coarse and fine, and also provide signals for the system controlling binocular vergence. These results indicate that cortical visual neurons are binocularly linked to respond to the relative position and contrast of the images over their receptive fields, and also that both these aspects of binocular stimulation may be utilized by the brain as a source of stereoscopic information.

Animals↗

Effects of luminance and flicker on ocular dominance shift in kitten visual cortex.

We raised monocularly deprived kittens in visual environments with low level illumination that was either steady or flickering. With steady scotopic luminance ocular dominance shifted as it does in normal photopic lighting. In flickering light with an average frequency of 2 Hz there was virtually no ocular dominance shift, while in flickering light averaging 0.1 Hz there was a significant shift. Recordings from the 2 Hz flicker-reared were similar to the dark-reared recordings. The flickering illumination was produced in one case by a high contrast-low brightness TV near the cage, and in another case, by a low voltage incandescent bulb driven by a pseudo-random sequence generator. This circuit delivered either a maximum ON time of 1.7 s or a maximum of 40 s, for the 2 Hz and 0.1 Hz respectively. Both the TV and flickering bulb produced average illumination comparable to the dim (0.01 cd/m2) steady scotopic illumination. We conclude that dim flickering light is not a sufficient stimulus for promoting ocular dominance shift in kittens in the critical period unless the flicker rate approaches 0.1 Hz. Furthermore results from the TV rearing suggest that flicker may be capable of preventing an ocular dominance shift expected from a concurrent steady low light level background.

Animals↗

Implantation of retina stimulation electrodes and recording of electrical stimulation responses in the visual cortex of the cat.

BACKGROUND: Simple basic visual perception may be restored by epiretinal electrical stimulation in patients that are blind due to photoreceptor loss. To stimulate ganglion cells, epiretinally flat platinum microelectrodes embedded in thin polyimide film were developed and tested in the cat. METHODS: After removal of the lens and the vitreous body a thin microfilm electrode array was implanted through a corneoscleral incision in the cat eye (n = 4). In two eyes no further attempt was made to fixate the tip of the electrode, which was pressed onto the retinal surface due to the tension of the curved polyimide film. In two eyes the tip of the electrode was fixed with cyanoacrylate adhesive. The exterior part of the microelectrode film was directed under the skin towards the forehead which allowed fixation of the microplug to a head fixation bolt. Retinal stimulation experiments were performed within 1 week after implantation. Success of stimulation was assessed by recording neuronal activities from areas 17 and 18. Retinal microelectrodes were removed 2 weeks or longer after implantation. RESULTS: Intraocular inflammation or retinal detachment were not observed after implantation of the microelectrode film. In two eyes the tip of the microelectrodes dislocated spontaneously within the first few days. The lowest threshold of electrical stimulation was 35 microA, corresponding to a charge transfer of 14 nC per phase. These values were ten times higher than those obtained by needle electrodes used in prior experiments. CONCLUSIONS: Intraocular implanted flat microelectrodes made of platinum and polyimide were well tolerated. Because of the flat configuration of the microelectrodes higher stimulation thresholds than for needle electrodes were found, indicating insufficient contact to the retinal surface. An alternative shape and fixation technique is required to minimise electrodes' threshold of stimulation.

Animals↗

The statistical reliability of signals in single neurons in cat and monkey visual cortex.

The variability of the discharge of visual cortical neurons in cats and macaque monkeys limits the reliability with which such neurons can relay signals about weak visual stimuli. In general, the variance of a neuron's firing rate is directly proportional to its mean firing rate. The probability that a neuron will fire a criterion number of impulses on a stimulus trial grows monotonically with the contrast of a sinusoidal grating stimulus. Neural probability functions prepared either by computing the probability of criterion response or by integrating receiver operating characteristics to yield the probability of correct choice in a two-alternative forced-choice situation resemble psychometric functions obtained in psychophysical and behavioral experiments on humans and animals, but are shallower in slope. The slopes of neuronal probability functions are slightly higher when they are estimated over short time periods, but even so do not equal the slopes measured psychophysically in human and monkey observers. This discrepancy in slope could be explained if the whole observer responded only when about four neurons were active together.

Animals↗

Functional streams and local connections of layer 4C neurons in primary visual cortex of the macaque monkey.

The primate visual system is composed of multiple, functionally specialized cortical areas. The functional diversity among areas is thought to reflect different contributions from early parallel visual pathways to the area V1 neurons providing input to "higher" cortical areas. The M pathway is believed to provide information about motion and contrast, via layer 4B of V1, to dorsal visual areas. The P pathway is believed to provide information about shape and color, via layer 2/3 of V1, to ventral visual areas, with specialized contributions from cytochrome-oxidase (CO) blob versus interblob neurons. However, the detailed anatomical relationships between the M and P pathways and the neurons in V1 that provide input to higher extrastriate cortical areas are poorly understood. To study these relationships, spiny stellate neurons in the M- and P-recipient layers of V1, 4Calpha and 4Cbeta, respectively, were intracellularly labeled, and their axonal and dendritic arbors were reconstructed. We find that neurons with dendrites in upper layer 4Calpha project axons to layer 4B and CO blobs in layer 2/3, thus relaying M input to these regions. Other neurons in lower layer 4Calpha provide M input to interblobs. These cells have either (1) dendrites restricted to lower layer 4Calpha and axons specifically targeting layer 2/3 interblobs, or (2) dendrites in lower 4Calpha and 4Cbeta and axons targeting blobs and interblobs. P-recipient layer 4Cbeta neurons have dense axonal arbors in both blobs and interblobs but not layer 4B. Quantitative analyses reveal that 4Calpha cells provide approximately five times more synapses than 4Cbeta cells to layer 4B, whereas 4Cbeta cells provide five times more synapses than 4Calpha cells to layer 2/3. These observations imply that M input is dominant in layer 4B. In layer 2/3, both blobs and interblobs receive M and P input, but the P input is dominant, and M input to interblobs derives exclusively from a subpopulation of M afferents that targets lower 4Calpha, not from afferents targeting only upper 4Calpha (cf. Blasdel and Lund, 1983). We speculate that the M and P pathways to interblobs are "X-like" linear systems, whereas blobs also receive nonlinear "Y-like" M input.

Animals↗

Lateral interactions in primary visual cortex: a model bridging physiology and psychophysics.

Recent physiological studies show that the spatial context of visual stimuli enhances the response of cells in primary visual cortex to weak stimuli and suppresses the response to strong stimuli. A model of orientation-tuned neurons was constructed to explore the role of lateral cortical connections in this dual effect. The differential effect of excitatory and inhibitory current and noise conveyed by the lateral connections explains the physiological results as well as the psychophysics of pop-out and contour completion. Exploiting the model's property of stochastic resonance, the visual context changes the model's intrinsic input variability to enhance the detection of weak signals.

Animals↗