The Representation of Orientation in Macaque V2: Four Stripes Not Three (original) (raw)
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Orientation selectivity in macaque VI: diversity and laminar dependence. J Neurosci
The Journal of Neuroscience : The Official Journal of the Society for Neuroscience
We studied the steady-state orientation selectivity of single neurons in macaque primary visual cortex (V1). To analyze the data, two measures of orientation tuning selectivity, circular variance and orientation bandwidth, were computed from the tuning curves. Circular variance is a global measure of the shape of the tuning curve, whereas orientation bandwidth is a local measure of the sharpness of the tuning curve around its peak. Circular variance in V1 was distributed broadly, indicating a great diversity of orientation selectivity. This diversity was also reflected in the individual cortical layers. However, there was a tendency for neurons with high circular variance, meaning low selectivity for orientation, to be concentrated in layers 4C, 3B, and 5. The relative variation of orientation bandwidth across the cortical layers was less than for circular variance, but it showed a similar laminar dependence. Neurons with large orientation bandwidth were found predominantly in layers 4C and 3B. There was a weak correlation between orientation selectivity and the level of spontaneous activity of the neurons. We also assigned a response modulation ratio for each cell, which is a measure of the linearity of spatial summation. Cells with low modulation ratios tended to have higher circular variance and bandwidth than those with high modulation ratios. These findings suggest a revision to the classical view that nonoriented receptive fields are principally found in layer 4C and the cytochrome oxidase-rich blobs in layer 2/3. Instead, a broad distribution of tuning selectivity is found in all cortical layers, and neurons that are weakly tuned for orientation are ubiquitous in V1 cortex.
Orientation selectivity in macaque V1: diversity and laminar dependence
The Journal of …, 2002
We studied the steady-state orientation selectivity of single neurons in macaque primary visual cortex (V1). To analyze the data, two measures of orientation tuning selectivity, circular variance and orientation bandwidth, were computed from the tuning curves. Circular variance is a ...
Orientation selectivity without orientation maps in visual cortex of a highly visual mammal
2005
In mammalian neocortex, the orderly arrangement of columns of neurons is thought to be a fundamental organizing principle. In primary visual cortex (V1), neurons respond preferentially to bars of a particular orientation, and, in many mammals, these orientationselective cells are arranged in a semiregular, smoothly varying map across the cortical surface. Curiously, orientation maps have not been found in rodents or lagomorphs. To explore whether this lack of organization in previously studied rodents could be attributable to low visual acuity, poorly differentiated visual brain areas, or small absolute V1 size, we examined V1 organization of a larger, highly visual rodent, the gray squirrel. Using intrinsic signal optical imaging and single-cell recordings, we found no evidence of an orientation map, suggesting that formation of orientation maps depends on mechanisms not found in rodents. We did find robust orientation tuning of single cells, and this tuning was invariant to stimulus contrast. Therefore, it seems unlikely that orientation maps are important for orientation tuning or its contrast invariance in V1. In vertical electrode penetrations, we found little evidence for columnar organization of orientation-selective neurons and little evidence for local anisotropy of orientation preferences. We conclude that an orderly and columnar arrangement of functional response properties is not a universal characteristic of cortical architecture.
Orientation domain diversity in macaque area V2
Eye and Brain, 2014
Single orientation domains in primary (V1) and second (V2) visual cortical areas are known to encode the orientation of visual contours. However, the visual world contains multiple and complex contour types. How do these domains handle such complexity? Using optical imaging methods, we have examined orientation response to two types of contours: real (luminance-defined) and illusory (inferred). We find that, unlike area V1, there are multiple types of orientation domain in V2. These include "real only" domains, "higher-order" domains (which respond to an orientation whether real or illusory), and other domains with nonmatching real/illusory orientation preference. We suggest that this plurality of orientation domains in V2 enables the complexities of local and global contour extraction.
Orientation tuning of cytochrome oxidase patches in macaque primary visual cortex
Nature Neuroscience, 2011
The abundant concentration of cytochrome oxidase in patches or blobs of primate striate cortex has never been explained. Patches are thought to contain unoriented, color-opponent neurons. Lacking orientation selectivity, these cells might endow patches with a high level of metabolic activity because they respond to all contours in visual scenes. To test this idea, orientation tuning was measured in layer 2/3 of macaque V1 using acutely implanted 100-electrode arrays. Each electrode recording site was identified, and assigned to the patch or interpatch compartment. The mean orientation bandwidth of cells was 28.4° in patches and 25.8° in interpatches. Neurons in patches were indeed less orientation selective, but the difference was subtle, indicating that the processing of form and color is not strictly segregated in V1. The most conspicuous finding was that patch cells had a 49% greater overall firing rate. This global difference in neuronal responsiveness, rather than an absence of orientation tuning, may account for the rich mitochondrial enzyme activity that defines patches.
Functional implications of orientation maps in visual cortex
Journal of Vision, 2016
Stimulus orientation in the primary visual cortex of primates and carnivores is mapped as iso-orientation domains radiating from pinwheel centres, where orientation preferences of neighbouring cells change circularly. Whether this orientation map has a function is currently debated, because many mammals, such as rodents, do not have such maps. Here we show that two fundamental properties of visual cortical responses, contrast saturation and crossorientation suppression, are stronger within cat iso-orientation domains than at pinwheel centres. These differences develop when excitation (not normalization) from neighbouring oriented neurons is applied to different cortical orientation domains and then balanced by inhibition from un-oriented neurons. The functions of the pinwheel mosaic emerge from these local intra-cortical computations: Narrower tuning, greater cross-orientation suppression and higher contrast gain of iso-orientation cells facilitate extraction of object contours from images, whereas broader tuning, greater linearity and less suppression of pinwheel cells generate selectivity for surface patterns and textures.
Cerebral Cortex, 2004
We studied orientation selectivity in V1 of alert monkeys and its relationship to other physiological parameters and to anatomical organization. Single neurons were stimulated with drifting bars or with sinusoidal gratings while compensating for eye position. Orientation selectivity based on spike counts was quantified by circular variance and by the bandwidth of the orientation tuning curve. The circular variance distribution was bimodal, suggesting groups with low and with high selectivity. Orientation selectivity was clearly correlated with spontaneous activity, classical receptive field (CRF) size and the strength of surround suppression. Laminar distributions of neuronal properties were distinct. Neurons in the output layers 2/3, 4B and 5 had low spontaneous activity, small CRFs and high orientation selectivity, while the input layers had greater diversity. Direction-selective cells were among the neurons most selective for orientation and most had small CRFs. A narrow band of direction-and orientation-selective cells with small CRFs was located in the middle of layer 4C, indicating appearance of very selective cells at an early stage of cortical processing. We suggest that these results reflect interactions between excitatory and inhibitory mechanisms specific to each sublamina. Regions with less inhibition have higher spontaneous activity, larger CRFs and broader orientation tuning. Where inhibition is stronger, spontaneous activity almost disappears, CRFs shrink, and orientation selectivity is high.
Effect of Stimulus Size on the Dynamics of Orientation Selectivity in Macaque V1
Journal of Neurophysiology, 2005
Previous research has established that orientation selectivity depends to a great extent on suppressive mechanisms in the visual cortex. In this study, we investigated the spatial organization and the time-course of these mechanisms. Stimuli were presented in circular windows of “optimal” and “large” radii. The two stimulus sizes were chosen based on an area-response function measured with drifting gratings at high contrast. The “optimal” size was defined as the smallest radius that elicited the peak response (average value of 0.45°), whereas “large” was defined as two to five times the optimal size. We found that the peak amplitude of tuned enhancement and untuned suppression varied <10% on average with stimulus radius, indicating that they are mainly concentrated in the classical receptive field. However, tuned suppression—in those cells that showed it—was significantly stronger with large stimuli, indicating that this component has a contribution from beyond the classical rece...
2005
We studied orientation selectivity in V1 of alert monkeys and its relationship to other physiological parameters and to anatomical organization. Single neurons were stimulated with drifting bars or with sinusoidal gratings while compensating for eye position. Orientation selectivity based on spike counts was quantified by circular variance and by the bandwidth of the orientation tuning curve. The circular variance distribution was bimodal, suggesting groups with low and with high selectivity. Orientation selectivity was clearly correlated with spontaneous activity, classical receptive field (CRF) size and the strength of surround suppression. Laminar distributions of neuronal properties were distinct. Neurons in the output layers 2/3, 4B and 5 had low spontaneous activity, small CRFs and high orientation selectivity, while the input layers had greater diversity. Direction-selective cells were among the neurons most selective for orientation and most had small CRFs. A narrow band of direction-and orientation-selective cells with small CRFs was located in the middle of layer 4C, indicating appearance of very selective cells at an early stage of cortical processing. We suggest that these results reflect interactions between excitatory and inhibitory mechanisms specific to each sublamina. Regions with less inhibition have higher spontaneous activity, larger CRFs and broader orientation tuning. Where inhibition is stronger, spontaneous activity almost disappears, CRFs shrink, and orientation selectivity is high.