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      Asymmetries in visual acuity around the visual field

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          Abstract

          Human vision is heterogeneous around the visual field. At a fixed eccentricity, performance is better along the horizontal than the vertical meridian and along the lower than the upper vertical meridian. These asymmetric patterns, termed performance fields, have been found in numerous visual tasks, including those mediated by contrast sensitivity and spatial resolution. However, it is unknown whether spatial resolution asymmetries are confined to the cardinal meridians or whether and how far they extend into the upper and lower hemifields. Here, we measured visual acuity at isoeccentric peripheral locations (10 deg eccentricity), every 15° of polar angle. On each trial, observers judged the orientation (± 45°) of one of four equidistant, suprathreshold grating stimuli varying in spatial frequency (SF). On each block, we measured performance as a function of stimulus SF at 4 of 24 isoeccentric locations. We estimated the 75%-correct SF threshold, SF cutoff point (i.e., chance-level), and slope of the psychometric function for each location. We found higher SF estimates (i.e., better acuity) for the horizontal than the vertical meridian and for the lower than the upper vertical meridian. These asymmetries were most pronounced at the cardinal meridians and decreased gradually as the angular distance from the vertical meridian increased. This gradual change in acuity with polar angle reflected a shift of the psychometric function without changes in slope. The same pattern was found under binocular and monocular viewing conditions. These findings advance our understanding of visual processing around the visual field and help constrain models of visual perception.

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          Human photoreceptor topography.

          We have measured the spatial density of cones and rods in eight whole-mounted human retinas, obtained from seven individuals between 27 and 44 years of age, and constructed maps of photoreceptor density and between-individual variability. The average human retina contains 4.6 million cones (4.08-5.29 million). Peak foveal cone density averages 199,000 cones/mm2 and is highly variable between individuals (100,000-324,000 cones/mm2). The point of highest density may be found in an area as large as 0.032 deg2. Cone density falls steeply with increasing eccentricity and is an order of magnitude lower 1 mm away from the foveal center. Superimposed on this gradient is a streak of high cone density along the horizontal meridian. At equivalent eccentricities, cone density is 40-45% higher in nasal compared to temporal retina and slightly higher in midperipheral inferior compared to superior retina. Cone density also increases slightly in far nasal retina. The average human retina contains 92 million rods (77.9-107.3 million). In the fovea, the average horizontal diameter of the rod-free zone is 0.350 mm (1.25 degrees). Foveal rod density increases most rapidly superiorly and least rapidly nasally. The highest rod densities are located along an elliptical ring at the eccentricity of the optic disk and extending into nasal retina with the point of highest density typically in superior retina (5/6 eyes). Rod densities decrease by 15-25% where the ring crosses the horizontal meridian. Rod density declines slowly from the rod ring to the far periphery and is highest in nasal and superior retina. Individual variability in photoreceptor density differs with retinal region and is similar for both cones and rods. Variability is highest near the fovea, reaches a minimum in the midperiphery, and then increases with eccentricity to the ora serrata. The total number of foveal cones is similar for eyes with widely varying peak cone density, consistent with the idea that the variability reflects differences in the lateral migration of photoreceptors during development. Two fellow eyes had cone and rod numbers within 8% and similar but not identical photoreceptor topography.
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            Peripheral vision and pattern recognition: a review.

            We summarize the various strands of research on peripheral vision and relate them to theories of form perception. After a historical overview, we describe quantifications of the cortical magnification hypothesis, including an extension of Schwartz's cortical mapping function. The merits of this concept are considered across a wide range of psychophysical tasks, followed by a discussion of its limitations and the need for non-spatial scaling. We also review the eccentricity dependence of other low-level functions including reaction time, temporal resolution, and spatial summation, as well as perimetric methods. A central topic is then the recognition of characters in peripheral vision, both at low and high levels of contrast, and the impact of surrounding contours known as crowding. We demonstrate how Bouma's law, specifying the critical distance for the onset of crowding, can be stated in terms of the retinocortical mapping. The recognition of more complex stimuli, like textures, faces, and scenes, reveals a substantial impact of mid-level vision and cognitive factors. We further consider eccentricity-dependent limitations of learning, both at the level of perceptual learning and pattern category learning. Generic limitations of extrafoveal vision are observed for the latter in categorization tasks involving multiple stimulus classes. Finally, models of peripheral form vision are discussed. We report that peripheral vision is limited with regard to pattern categorization by a distinctly lower representational complexity and processing speed. Taken together, the limitations of cognitive processing in peripheral vision appear to be as significant as those imposed on low-level functions and by way of crowding. © ARVO
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              Topography of ganglion cells in human retina.

              We quantified the spatial distribution of presumed ganglion cells and displaced amacrine cells in unstained whole mounts of six young normal human retinas whose photoreceptor distributions had previously been characterized. Cells with large somata compared to their nuclei were considered ganglion cells; cells with small somata relative to their nuclei were considered displaced amacrine cells. Within the central area, ganglion cell densities reach 32,000-38,000 cells/mm2 in a horizontally oriented elliptical ring 0.4-2.0 mm from the foveal center. In peripheral retina, densities in nasal retina exceed those at corresponding eccentricities in temporal retina by more than 300%; superior exceeds inferior by 60%. Displaced amacrine cells represented 3% of the total cells in central retina and nearly 80% in the far periphery. A twofold range in the total number of ganglion cells (0.7 to 1.5 million) was largely explained by a similar range in ganglion cell density in different eyes. Cone and ganglion cell number were not correlated, and the overall cone:ganglion cell ratio ranged from 2.9 to 7.5 in different eyes. Peripheral cones and ganglion cells have different topographies, thus suggesting meridianal differences in convergence onto individual ganglion cells. Low convergence of foveal cones onto individual ganglion cells is an important mechanism for preserving high resolution at later stages of neural processing. Our improved estimates for the density of central ganglion cells allowed us to ask whether there are enough ganglion cells for each cone at the foveal center to have a direct line to the brain. Our calculations indicate that 1) there are so many ganglion cells relative to cones that a ratio of only one ganglion cell per foveal cone would require fibers of Henle radiating toward rather than away from the foveal center; and 2) like the macaque, the human retina may have enough ganglion cells to transmit the information afforded by closely spaced foveal cones to both ON- and OFF-channels. Comparison of ganglion cell topography with the visual field representation in V1 reveals similarities consistent with the idea that cortical magnification is proportional to ganglion cell density throughout the visual field.
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                Author and article information

                Journal
                J Vis
                J Vis
                jovi
                JOVI
                Journal of Vision
                The Association for Research in Vision and Ophthalmology
                1534-7362
                05 January 2021
                January 2021
                : 21
                : 1
                : 2
                Affiliations
                [1 ]Department of Psychology, New York University, New York, NY, USA
                [2 ]Center for Neural Science, New York University, New York, NY, USA
                [3 ]Spinoza Centre for Neuroimaging, Amsterdam, Netherlands
                Author notes
                Article
                JOV-07414-2020
                10.1167/jov.21.1.2
                7794272
                33393963
                6dafa4a3-2098-4178-8594-672a2c56433a
                Copyright 2021 The Authors

                This work is licensed under a Creative Commons Attribution 4.0 International License.

                History
                : 10 November 2020
                : 13 April 2020
                Page count
                Pages: 23
                Categories
                Article
                Article

                spatial frequency,performance fields,visual acuity,spatial vision,isoeccentricity

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