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The arrangement of the three cone classes in the living human eye

Nature volume 397, pages 520522 (11 February 1999) | Download Citation



Human colour vision depends on three classes of receptor, the short- (S), medium- (M), and long- (L) wavelength-sensitive cones. These cone classes are interleaved in a single mosaic so that, at each point in the retina, only a single class of cone samples the retinal image. As a consequence, observers with normal trichromatic colour vision are necessarily colour blind on a local spatial scale1. The limits this places on vision depend on the relative numbers and arrangement of cones. Although the topography of human S cones is known2,3, the human L- and M-cone submosaics have resisted analysis. Adaptive optics, a technique used to overcome blur in ground-based telescopes4, can also overcome blur in the eye, allowing the sharpest images ever taken of the living retina5. Here we combine adaptive optics and retinal densitometry6 to obtain what are, to our knowledge, the first images of the arrangement of S, M and L cones in the living human eye. The proportion of L to M cones is strikingly different in two male subjects, each of whom has normal colour vision. The mosaics of both subjects have large patches in which either M or L cones are missing. This arrangement reduces the eye's ability to recover colour variations of high spatial frequency in the environment but may improve the recovery of luminance variations of high spatial frequency.

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  1. 1.

    , , , & in From Pigments to Perception.(eds Valberg, A. & Lee, B. B.) 11–22 (Plenum, New York, (1991)).

  2. 2.

    , & Punctate sensitivity of the blue sensitive mechanism.Vision Res. 21, 1357–1375 (1981).

  3. 3.

    et al. Distribution and morphology of human cone photoreceptors stained with anti-blue opsin.J. Comp. Neurol. 312, 610–624 (1991).

  4. 4.

    The possibility of compensating astronomical seeing.Publ. Astron. Soc. Pacif. 65, 229–236 (1953).

  5. 5.

    , & Supernormal vision and high-resolution retinal imaging through adaptive opticsJ. Opt. Soc. Am. A 14, 2884–2892 (1997).

  6. 6.

    & Measurement of the scotopic pigment in the living human eye.J.Physiol. (Lond.) 130, 131–147 (1955).

  7. 7.

    & Red/green sensitivity in normal vision.Vision Res. 4, 75–85 (1964).

  8. 8.

    , & in From Pigments to Perception(eds Valberg, A. & Lee, B. B.) 23–34 (Plenum, New York, (1991)).

  9. 9.

    & The relative numbers of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea.Vision Res. 26, 115–128 (1989).

  10. 10.

    , , & Foveal cone thresholds.Vision Res. 29, 61–78 (1989).

  11. 11.

    & in Colour Vision Deficiencies Vol. XI(ed. Drum, B.) 107–112 (Kluwer Acedemic, Netherlands, (1993)).

  12. 12.

    & Spectral sensitivity of macaque monkeys measured with ERG flicker photometry.Vis. Neurosci. 14, 921–928 (1997).

  13. 13.

    & Visual pigments of rods and cones in a human retina.J. Physiol. (Lond.) 298, 501–511 (1980).

  14. 14.

    , & Human visual pigments: microspectrophotometric results from the eyes of seven persons.Proc. R. Soc. Lond. B 220, 115–130 (1983).

  15. 15.

    , & Visual pigment gene structure and expression in human retinae.Hum. Mol. Genet. 6, 981–990 (1998).

  16. 16.

    , & Variation in cone populations for red-green color vision examined by analysis of mRNA.NeuroReport 9, 1963–1967 (1998).

  17. 17.

    & The spatial arrangement of cones in the primate fovea.Nature 360, 677–679 (1992).

  18. 18.

    , & Photoreceptor transmittance imaging of the primate photoreceptor mosaic.J. Neurosci. 16, 2251–2260 (1996).

  19. 19.

    Uber den Farbensinn.Comp. Rend. Congr. Period. Intern. Sci. Med. Copenhagen 1, 80–98 (1884).

  20. 20.

    Color appearance of small stimuli and the spatial distribution of color receptors.J. Opt. Soc. Am. 54, 1171 (1964).

  21. 21.

    On the undulations excited in the retina by the action of luminous points and lines.Lond. Edinb. Philos. Mag. J. Sci. 1, 169–174 (1832).

  22. 22.

    , & Efficiency in detection of isoluminant and isochromatic interference fringes.J. Opt. Soc. Am. A. 10, 2118–2133 (1993).

  23. 23.

    in Advances in Photoreception: Proc. Symp. Frontiers Visual Sci.135–148 (National Academy, Washington DC, (1990)).

  24. 24.

    , , , & Color vision in two observers with highly biased LWS/MWS cone ratios.Vision Res. 38, 601–612 (1998).

  25. 25.

    , & Molecular genetics of human colour vision: the genes encoding blue, green and red pigments.Science 232, 193–202 (1986).

  26. 26.

    “Tho' she kneeled in that place where they grew...”: The uses and origins of primate colour vision.J. Exp. Biol. 146, 21–38 (1989).

  27. 27.

    Statistical Analysis of Spatial Point Patterns(Academic, London, (1983)).

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We thank D. Brainard, D. Dacey, J. Jacobs, J. Liang, D. Miller and O. Packer for their assistance. We acknowledge financial support from the Fight for Sight research division of Prevent Blindness America (to A.R.) and the National Eye Institute and Research to Prevent Blindness (to D.R.W.).

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    • Austin Roorda

    Present address: University of Houston, College of Optometry, Houston, Texas 77204-6052, USA


  1. *Center for Visual Science, University of Rochester, Rochester, New York 14627, USA

    • Austin Roorda
    •  & David R. Williams


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