Macaque monkeys were shown retinotopically-specific visual stimuli during 14C-2-deoxy-d-glucose (DG) infusion in a study of the retinotopic organization of primary visual cortex (V1). In the central half of V1, the cortical magnification was found to be greater along the vertical than along the horizontal meridian, and overall magnification factors appeared to be scaled proportionate to brain size across different species. The cortical magnification factor (CMF) was found to reach a maximum of about 15 mm/deg at the representation of the fovea, at a point of acute curvature in the V1-V2 border. We find neither a duplication nor an overrepresentation of the vertical meridian. The magnification factor did not appear to be doubled in a direction perpendicular to the ocular dominance strips; it may not be increased at all. The DG borders in parvorecipient layer 4Cb were found to be as sharp as 140 micron (half-amplitude, half width), corresponding to a visual angle of less than 2' of arc at the eccentricity measured. In other layers (including magnorecipient layer 4Ca), the retinotopic borders are broader. The retinotopic spread of activity is greater when produced by a low-spatial-frequency grating than when produced by a high-spatial-frequency grating. Orientation-specific stimuli produced a pattern of activation that spread further than 1 mm across cortex in some layers. Some DG evidence suggests that the spread of functional activity is greater near the foveal representation than near 5 degrees eccentricity.

We have anatomically analyzed retinotopic organization using the 14C-labeled 2-deoxy-D-glucose method. The method has several advantages over conventional electrophysiological mapping techniques. In the striate cortex, the anatomical substrate for retinotopic organization is surprisingly well ordered, and there seems to be a systematic relationship between ocular dominance strips and cortical magnification. The 2-deoxyglucose maps in this area appear to be largely uninfluenced by known differences in long-term metabolic activity. This method should prove useful in analyzing retinotopic organization in various visual areas of the brain and in different species.

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When macaque monkeys view achromatic, sinusoidal gratings of a single spatial frequency, the pattern of 14C-2-deoxy-d-glucose (DG) uptake produced by the gratings is shown to depend on the spatial frequency chosen. When a relatively high (5-7 cycles/deg) spatial frequency is shown binocularly at systematically varied orientations, uptake in parafoveal striate cortex is highest between the cytochrome oxidase blobs (that is, in the interblobs) in layers 1, 2, and 3. In layers 4B, 5, and 6, where the cytochrome oxidase blobs are faint or absent, DG uptake is highest in a periodic pattern that lies in register with the interblobs of layers 2 + 3. When the grating is, instead, of relatively low (1-1.5 cycles/deg) spatial frequency, DG uptake is highest in the blobs, in the blob-aligned portions of layers 1-4B, and in the lower-layer blobs as well. These variations in DG topography are confirmed in stimulus comparisons within a single hemisphere. Presumably, this shift in functional topography within the extra-granular layer is the primate homolog of "spatial frequency columns" shown earlier in the cat (Tootell et al., 1981; Silverman, 1984). In the well-differentiated architecture of primate striate cortex, laminar differences produced by high- versus low-spatial-frequency gratings are visible as well. Gratings of very high spatial frequency produce much higher uptake in 4Cb (which receives input from the parvocellular LGN layers) than in 4Ca (which gets its input from the magnocellular LGN layers). Gratings of low spatial frequency produce the converse result. Presumably, cells in the magnocellular LGN layers and/or in the magnocellular-dominated layer 4Ca have lower average spatial frequency tuning (larger receptive fields) than their counterparts in the parvocellular LGN and/or in striate layer 4Cb. The DG patterns produced by various spatial frequencies also vary with eccentricity, in a manner consistent with known, eccentricity-dependent variations of receptive-field size and spatial frequency tuning. Thus, gratings of a "middle"-spatial-frequency range (4-5 cycles/deg) produce high uptake in the blobs near the foveal representation and high uptake in the interblobs at more peripheral eccentricities, including 5 degrees. This shift in DG topography also includes the transition zone near 3 degrees, where the level of stimulus-driven uptake is as high in the blob regions as it is in interblob regions. Variations in uptake between layers 4Ca and 4Cb, as a function of eccentricity, shift in parallel with the changes in the upper-layer topography.

The contrast dependence of simultaneous masking has been measured using isochromatic yellow-black luminance sinusoids and isoluminant red-green chrominance gratings. Masking functions for all four combinations of chromatic and luminance masks and tests are reported. In the two same-on-same conditions (luminance mask/luminance test and chromatic mask/chromatic test) these functions (increment threshold contrast versus mask contrast) have the typical dipper shape and are almost identical when test and mask contrasts are normalized to the unmasked contrast thresholds. The contrast dependence of the luminance mask/color test and color mask/luminance test functions are quite different. The luminance mask/color test shows facilitation over a broad range of both subthreshold and suprathreshold contrasts of the luminance mask. In the color mask/luminance test condition facilitation is never observed, but at suprathreshold contrasts a 2-cycle/degree (c/deg) chromatic grating masks a 2-c/deg luminance grating as strongly as does a luminance mask. The luminance mask/chromatic test results are invariant over the 0.25-2-c/deg spatial-frequency range, whereas the robust masking of luminance by color at 2 c/deg diminishes at lower spatial frequencies. The spatial-frequency selectivity of the luminance-facilitates-color interaction is much broader than facilitatory interactions in either the color-color or luminance-luminance conditions. Possible mechanisms of color-luminance interactions are considered. The lack of facilitation in the color mask/luminance test condition precludes a simple pedestal interpretation of this masking interaction. The data are, however, consistent with models that invoke inhibitory or more elaborate excitatory masking interactions.

A series of experiments was carried out using 14C-2-deoxy-d-glucose (DG) in order to examine the functional architecture of macaque striate (primary visual) cortex. This paper describes the results of experiments on uptake during various baseline (or reference) conditions of visual stimulation (described below), and on differences in the functional architecture following monocular versus binocular viewing conditions. In binocular “baseline” experiments, monkeys were stimulated either (1) in the dark, (2) with a diffuse gray screen, or (3) with a very general visual stimulus composed of gratings of varied orientation and spatial frequency. In all of these conditions, DG uptake was found to be topographically uniform within all layers of parafoveal striate cortex. In monocular experiments that were otherwise similar, uptake was topographically uniform within the full extent of the eye dominance strip, in all layers. Certain other visual stimuli produce high uptake in the blobs, and still another set of visual stimuli (including high-spatial-frequency gratings) produce highest uptake between the blobs at parafoveal eccentricities, even in an unanesthetized, unparalyzed monkey. Eye movements per se had no obvious effect on striate DG uptake. Endogenous uptake in the blobs (relative to that in the interblobs) appears higher in the squirrel monkey than in the macaque. The pattern of DG uptake produced by binocular viewing was found to deviate in a number of ways from that expected by linearly summing the component monocular DG patterns. One of the most interesting deviations was an enhancement of the representation of visual field borders between stimuli differing from each other in texture, orientation, direction, etc. This “border enhancement” was confined to striate layers 1–3 (not appearing in any of the striate input layers), and it only appeared following binocular, but not monocular, viewing conditions. The border enhancement may be related to a suppression of DG uptake that occurs during binocular viewing conditions in layers 2 + 3 (and perhaps layers 1 and 4B), but not in layers 4Ca, 4Cb, 5 or 6. Another major class of binocular interaction was a spread of neural activity into the “unstimulated” ocular dominance strips following monocular stimulation. Such an effect was prominent in striate layer 4Ca, but it did not occur in layer 4Cb. This “binocular” spread of DG uptake into the inappropriate eye dominance strip in 4Ca may be related to the appearance of orientation tuning and orientation columns in that layer. No DG effects were seen that depended on the absolute disparity of visual stimuli in macaque striate cortex.

Using a hue scaling technique, we have examined the appearance of colored spots produced by shifts from white to isoluminant stimuli along various color vectors in order to examine color appearance without the complications of the combined luminance and chromatic stimulation involved in most previous hue scaling studies, which have used flashes of monochromatic light. We also used spots lying along cone-isolating vectors in order to determine what hues would be reported with a change in activation of only single cone types or of only single geniculate opponent-cell types, an issue of direct relevance to any model of color vision. We find that: 1. Unique hues do not correspond either to the change in activation of single cone types or of single geniculate opponent-cell types. This is well known to be the case for yellow and blue, but we find it to be true for red and green as well. 2. These conclusions are not limited to the particular white (Illuminant C) used as an adapting background in most of the experiments. Shifts along the same cone-contrast vectors relative to different backgrounds lead to much the same hue percepts, independent of the starting white used. 3. The shifts of the perceptual colors from the geniculate axes are in the directions, and close to the absolute amounts, predicted by our [De Valois & De Valois (1993). Vision Research, 33, 1053-1065] multi-stage color model in which we postulate that the S-opponent cells are added to or subtracted from the M- and L-opponent cells to form the four perceptual color systems. 4. There are distinct asymmetries with respect to the extent to which various hues within each perceptual opponent system deviate from the geniculate opponent-cell axes. Blue is shifted more from the S-LM axis than is yellow; green is shifted more from the L-M axis than is red. There are also asymmetries in the angular extent of opponent color regions. Blue is seen over a larger range of color vectors than is yellow, and red over a slightly larger range than green. 5. Such asymmetries are not accounted for by any model that treats red-green and yellow-blue each as unitary, mirror-image opponent-color systems. Although red and green are perceptually opponent, the red and green perceptual systems do not appear to be constructed in a mirror-image fashion with respect to input from different cone types or from different geniculate opponent-cell types. The same is true for yellow and blue.

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