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.

Topography of ganglion cells in human retina.

I. A consideration of methods for assessing contrast sensitivity leads to the conclusion that, for a clinical test, letters are more suitable than gratings. 2. A letter chart is described in which letters decrease in contrast but not in size. The letters are arranged in groups of three; successive groups decrease in contrast by a factor of IfJ2 from a very high contrast down to a contrast below the threshold of normal observers. A subject's threshold is taken to be the lowest contrast for which at least two letters in a group are correctly reported. 3. A mathematical model of the observer and the chart-testing procedure has been used to predict how the accuracy and repeatability of the test score depend on the parameters of the chart and observer. This reveals that even a low probability of misreporting supra threshold letters will seriously bias the test score if the passing criterion is strict, requiring correct report of all letters in each group, but will have little effect if the passing criterion is less strict. This effect of the passing criterion may explain Rubin's [Clin. Vision Sci. 2, No. I (1987)] finding that the new test, which uses a lenient criterion, has excellent test-retest reliability, much higher than the Ginsburg [Am. J. Optom. Physiol. Opt. 61,403-407 (1984)] chart with its strict criterion.

The design of a new letter chart for measuring contrast sensitivity

Comparisons of the published data on the density D of receptive fields of retinal ganglion cells and on the cortical magnification factor M indicated that M2 is directly proportional to D in primates. Therefore, the human M can be estimated for the principal meridians of the visual field from the density-distribution of retinal ganglion cells and from the density of the centralmost cones. Using the previously published empirical data, we estimated the values of the human M and express the values in four simple equations that can be used for finding the value of M for any location of the visual field. The monocular values of M are not radially symmetric.

An estimation and application of the human cortical magnification factor

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.

Peripheral vision and pattern recognition: a review.

https://visionlab.harvard.edu/Members/Patrick/PDF.files/CogPsychR.pdf

The spatial resolution of visual attention

THE area of visual cortex devoted to the analysis of a con-slant-size region in the visual field diminishes progressively for more peripheral locations. The change is described quantitatively by the cortical magnification factor, which indicates the linear extent of cortex in mm corresponding to one degree of visual angle at various eccentricities (angular distances from the middle fovea). The human cortical magnification factor has been estimated by Cowey and Rolls1 from the data of Brindley and Lewin2, who mapped the phosphenes (sensations of light) caused in the lower nasal visual field by electrical stimulation of the human visual cortex. Building on these results, we have studied the spatial contrast sensitivity functions in man at various eccentricities. We used two methods: in one the retinal image sizes of the test gratings were kept similar at different eccentricities and in the other, the calculated sizes of cortical projections of the test gratings were made similar at different locations. Our results indicate that peripheral contrast sensitivity and acuity are inferior to foveal performance, because of the diminished cortical projection area.

Cortical magnification factor predicts the photopic contrast sensitivity of peripheral vision.

Grating resolution was measured at various locations of the visual field for four grating orientations. As an instance of the oblique effect, vertical and horizontal gratings produced the highest resolution values in the central area. At eccentricities larger than about 20 deg, the oblique effect was replaced by a meridional resolution effect, in which resolution was systematically best for meridionally oriented grating bars and worst for grating bars perpendicular to the visual-field meridians. The origin of the effect seems to be neural because it was not caused by peripheral refractive errors or optical distortion of the peripheral retinal image.

Resolution of gratings oriented along and across meridians in peripheral vision.

Temporal contrast sensitivity and cortical magnification.

Spatial contrast sensitivity of 19 diabetics with different degrees of visual impairment was studied. It was found that contrast sensitivity at intermediate and low spatial frequencies may decrease without corresponding loss of visual acuity. In advanced cases of diabetes the opposite may be true: contrast sensitivity was better than expected on the basis of visual acuity. Thus both visual acuity and contrast sensitivity measurements are useful in the evaluation of the nature of visual impairment due to diabetic eye disease.

Jyrki Rovamo

Papers

Cortical magnification factor predicts the photopic contrast sensitivity of peripheral vision.

Resolution of gratings oriented along and across meridians in peripheral vision.

Temporal contrast sensitivity and cortical magnification.

Contrast sensitivity in evaluation of visual impairment due to diabetes

Critical flicker frequency and M-scaling of stimulus size and retinal illuminance.