Human photoreceptor topography

We have measured the spatial density of cones and rods in eight wholemounted 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°). 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.

[1]  Max Johann Sigismund Schultze,et al.  Zur Anatomie und Physiologie der Retina , 1866 .

[2]  G. Fritsch Über Bau und Bedeutung der Area centralis des Menschen , 1908 .

[3]  Henrik Sjögren,et al.  ZUR KENNTNIS DER KERATOCONJUNCTIVITIS SICCA II , 1935 .

[4]  E. Ludvigh EXTRAFOVEAL VISUAL ACUITY AS MEASURED WITH SNELLEN TEST-LETTERS , 1941 .

[5]  H HARTRIDGE,et al.  Recent Advances in the Physiology of Vision , 1946, British medical journal.

[6]  B. O'Brien,et al.  Vision and resolution in the central retina. , 1951, Journal of the Optical Society of America.

[7]  Maynard C. Wheeler,et al.  VISUAL ACUITY WITHIN THE AREA CENTRALIS AND ITS RELATION TO EYE MOVEMENTS AND FIXATION , 1928 .

[8]  A. Cowey,et al.  Striate cortex lesions and visual acuity of the rhesus monkey. , 1963, Journal of comparative and physiological psychology.

[9]  D. G. Green,et al.  Optical and retinal factors affecting visual resolution. , 1965, The Journal of physiology.

[10]  F. Campbell,et al.  Optical quality of the human eye , 1966, The Journal of physiology.

[11]  M. H. Pirenne,et al.  VISION AND THE EYE , 1949 .

[12]  A. Laties,et al.  Photoreceptor Orientation in the Primate Eye , 1968, Nature.

[13]  L L Sloan,et al.  The photopic acuity-luminance function with special reference to parafoveal vision. , 1968, Vision research.

[14]  L Frisén The cartographic deformations of the visual field. , 1970, Ophthalmologica. Journal international d'ophtalmologie. International journal of ophthalmology. Zeitschrift fur Augenheilkunde.

[15]  D. G. Green Regional variations in the visual acuity for interference fringes on the retina , 1970, The Journal of physiology.

[16]  D. Worthen,et al.  Histology of the Human Eye. , 1972 .

[17]  F. Ederer,et al.  Shall we count numbers of eyes or numbers of subjects? , 1973, Archives of ophthalmology.

[18]  N Drasdo,et al.  Non-linear projection of the retinal image in a wide-angle schematic eye. , 1974, The British journal of ophthalmology.

[19]  Rod and cone densities in the Rhesus. , 1974, Investigative ophthalmology.

[20]  W. H. Miller,et al.  Photoreceptor diameter and spacing for highest resolving power. , 1977, Journal of the Optical Society of America.

[21]  Central vision of man and macaque: cone and rod sensitivity , 1977, Brain Research.

[22]  William H. Miller,et al.  Ocular Optical Filtering , 1979 .

[23]  T Wertheim,et al.  Peripheral visual acuity: Th. Wertheim. , 1980, American journal of optometry and physiological optics.

[24]  D. Borwein,et al.  The ultrastructure of monkey foveal photoreceptors, with special reference to the structure, shape, size, and spacing of the foveal cones. , 1980, The American journal of anatomy.

[25]  P. Henkind,et al.  Aging and degeneration of the human macula. 1. Outer nuclear layer and photoreceptors. , 1981, The British journal of ophthalmology.

[26]  J Hirsch,et al.  Limits of spatial-frequency discrimination as evidence of neural interpolation. , 1982, Journal of the Optical Society of America.

[27]  J. Yellott Spectral analysis of spatial sampling by photoreceptors: Topological disorder prevents aliasing , 1982, Vision Research.

[28]  G. Westheimer The spatial grain of the perifoveal visual field , 1982, Vision Research.

[29]  D. Williams,et al.  Consequences of spatial sampling by a human photoreceptor mosaic. , 1983, Science.

[30]  Gary D. Bernard,et al.  Averaging over the foveal receptor aperture curtails aliasing , 1983, Vision Research.

[31]  A. Hendrickson,et al.  The morphological development of the human fovea. , 1984, Ophthalmology.

[32]  J. Hirsch,et al.  Orientation dependence of visual hyperacuity contains a component with hexagonal symmetry. , 1984, Journal of the Optical Society of America. A, Optics and image science.

[33]  J. Hirsch,et al.  Quality of the primate photoreceptor lattice and limits of spatial vision , 1984, Vision Research.

[34]  S. Schein,et al.  Density profile of blue-sensitive cones along the horizontal meridian of macaque retina. , 1985, Investigative ophthalmology & visual science.

[35]  J. Flannery,et al.  Distribution patterns of photoreceptors, protein, and cyclic nucleotides in the human retina. , 1985, Investigative ophthalmology & visual science.

[36]  L. Reymond Spatial visual acuity of the eagle Aquila audax: a behavioural, optical and anatomical investigation , 1985, Vision Research.

[37]  I. Klock,et al.  Zonulae adherentes pore size in the external limiting membrane of the rabbit retina. , 1985, Investigative ophthalmology & visual science.

[38]  R. Nussenblatt,et al.  Effects of cyclosporine on T-cell subsets in experimental autoimmune uveitis. , 1985, Investigative ophthalmology & visual science.

[39]  A. Cowey,et al.  The ganglion cell and cone distributions in the monkey's retina: Implications for central magnification factors , 1985, Vision Research.

[40]  David Williams Aliasing in human foveal vision , 1985, Vision Research.

[41]  C A Curcio,et al.  Computer‐assisted morphometry using video‐mixed microscopic images and computer graphics , 1986, The Anatomical record.

[42]  A. Hendrickson,et al.  A qualitative and quantitative analysis of the human fovea during development , 1986, Vision Research.

[43]  W S Geisler,et al.  Sampling-theory analysis of spatial vision. , 1986, Journal of the Optical Society of America. A, Optics and image science.

[44]  David R. Williams,et al.  Seeing through the photoreceptor mosaic , 1986, Trends in Neurosciences.

[45]  Terry Bossomaier,et al.  Optical image quality and the cone mosaic. , 1986, Science.

[46]  P K Ahnelt,et al.  Identification of a subtype of cone photoreceptor, likely to be blue sensitive, in the human retina , 1987, The Journal of comparative neurology.

[47]  D. Williams,et al.  Cone spacing and the visual resolution limit. , 1987, Journal of the Optical Society of America. A, Optics and image science.

[48]  R A Smith,et al.  Aliasing in the parafovea with incoherent light. , 1987, Journal of the Optical Society of America. A, Optics and image science.

[49]  F. Fitzke,et al.  Retinal magnification factor at the ora terminalis: A structural study of human and animal eyes , 1987, Vision Research.

[50]  A J Ahumada,et al.  Cone sampling array models. , 1987, Journal of the Optical Society of America. A, Optics and image science.

[51]  Christine A. Curcio,et al.  A whole mount method for sequential analysis of photoreceptor and ganglion cell topography in a single retina , 1987, Vision Research.

[52]  W. H. Miller,et al.  Does cone positional disorder limit resolution? , 1987, Journal of the Optical Society of America. A, Optics and image science.

[53]  L. Thibos,et al.  Retinal limits to the detection and resolution of gratings. , 1987, Journal of the Optical Society of America. A, Optics and image science.

[54]  A. Hendrickson,et al.  Distribution of cones in human and monkey retina: individual variability and radial asymmetry. , 1987, Science.

[55]  L. Reymond,et al.  Spatial visual acuity of the falcon, Falco berigora: A behavioural, optical and anatomical investigation , 1987, Vision Research.

[56]  J. Hirsch,et al.  Two-dot vernier discrimination within 2.0 degrees of the foveal center. , 1987, Journal of the Optical Society of America. A, Optics and image science.

[57]  N J Coletta,et al.  Psychophysical estimate of extrafoveal cone spacing. , 1987, Journal of the Optical Society of America. A, Optics and image science.

[58]  S. Schein Anatomy of macaque fovea and spatial densities of neurons in foveal representation , 1988, The Journal of comparative neurology.

[59]  David Williams Topography of the foveal cone mosaic in the living human eye , 1988, Vision Research.

[60]  M Tessier-Lavigne,et al.  The effect of photoreceptor coupling and synapse nonlinearity on signal : noise ratio in early visual processing , 1988, Proceedings of the Royal Society of London. Series B. Biological Sciences.

[61]  Kenneth R. Sloan,et al.  Computer methods for sampling, reconstruction, display and analysis of retinal whole mounts , 1989, Vision Research.

[62]  Changes in rod sensitivity through adulthood. , 1989, Investigative ophthalmology & visual science.

[63]  Christine A. Curcio,et al.  The spatial resolution capacity of human foveal retina , 1989, Vision Research.

[64]  A. Hendrickson,et al.  Photoreceptor topography of the retina in the adult pigtail macaque (Macaca nemestrina) , 1989, The Journal of comparative neurology.