Abstract

Hue sensations resulting from the selective stimulation of short-wavelength-sensitive (S) and middle-wavelength-sensitive (M) cones were deduced from measurements of spectral unique green and unique blue under conditions of high or low S-cone sensitivity relative to M- and long-wavelength-sensitive-cone sensitivity. Selective reduction of S-cone stimulation shifted unique blue toward shorter wavelengths and unique green toward longer wavelengths, implying losses of perceived yellowness and short-wavelength redness relative to perceived blueness. The results imply that, under acromatic adaptation conditions, M-cone stimulation yields a sensation of predominately bluish cyan and S-cone stimulation yields a sensation of predominately reddish magenta. S-cone stimulation also appears to be indirectly responsible for yellowish sensations at long wavelengths and, by cancellation of the M-cone blueness signal, for greenish sensations at middle wavelengths.

© 1989 Optical Society of America

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  1. Representative examples of this model can be found in L. M. Hurvich, Color Vision (Sinauer, New York, 1982), pp. 128–135, and R. M. Boynton, Human Color Vision (Holt, Rinehart and Winston, New York, 1979), pp. 211–215. A notable exception is the model of C. R. Ingling, B. H.-P. Tsou [“Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977)], in which M cones also contribute to the blueness signal.
    [Crossref] [PubMed]
  2. The univariance of single cone types implies only that the responses to any two wavelengths can be made identical with suitable adjustments of intensity. It does not imply a lack of perceived color when only one cone type is stimulated. For example, it is easy to demonstrate with long-wavelength foveal stimuli in the dark-adapted eye that exclusive L-cone stimulation produces a predominately reddish hue sensation.
  3. The unique hues are defined as lights that produce perceptually pure sensations containing only one of the four fundamental hues of red, yellow, green, or blue. Along with the achromatic colors (white, gray, and black), they compose the set of equilibrium hues for which the response of at least one of the color-opponent systems is zero. Thus unique red and unique green are yellow–blue equilibria and unique yellow and unique blue are red–green equilibria.
  4. This paper is concerned with manipulating relative cone contributions to color perception, which is not necessarily the same as manipulating relative responses of the cones themselves. Perception does not occur at the photoreceptors but only after many intervening stages of neural processing that may substantially alter the space-, time-, and intensity-dependent properties of the visual signal. Thus the term cone sensitivity is used here to mean the sensitivity of the entire visual system to stimuli that produce responses in one or more of the L-, M-, and S-cone classes.
  5. C. H. Graham, Y. Hsia, F. F. Stephan, “Visual discriminations of a subject with acquired unilateral tritanopia,” Vision Res. 6, 469–479 (1966); N. Ohba, T. Tanino, “Unilateral colour vision defect resembling tritanopia,” Mod. Probl. Ophthalmol. 17, 331–335 (1976); M. Alpern, K. Kitahara, D. H. Krantz, “Perception of colour in unilateral tritanopia,”J. Physiol. (London) 335, 683–697 (1983).
    [PubMed]
  6. W. E. K. Middleton, M. C. Holmes, “The apparent colors of surfaces of small subtense—a preliminary report,”J. Opt. Soc. Am. 39, 582–592 (1949); D. O. Weitzman, J. A. S. Kinney, “Appearance of color for small, brief, spectral stimuli, in the central fovea,”J. Opt. Soc. Am. 57, 665–670 (1967); C. R. Ingling, H. M. O. Scheibner, R. M. Boynton, “Color naming of small foveal fields,” Vision Res. 10, 501–511 (1970); J. D. Mollon, “A taxonomy of tritanopias,” Doc. Ophthalmol. Proc. Ser. 33, 87–101 (1982).
    [Crossref] [PubMed]
  7. R. M. Boynton, W. Schafer, M. A. Neun, “Hue-wavelength relation measured by color-naming method for three retinal locations,” Science 146, 666–668 (1964); J. Gordon, I. Abramov, “Color vision in the peripheral retina. II. Hue and saturation,”J. Opt. Soc. Am. 67, 202–207 (1977).
    [Crossref] [PubMed]
  8. W. E. K. Middleton, E. G. Mayo, “The appearance of colors in twilight,”J. Opt. Soc. Am. 42, 116–121 (1951).
    [Crossref]
  9. M. Dagher, A. Cruz, L. Plaza, “Colour thresholds with monochromatic stimuli in the spectral region 530–630 μ m,” in Visual Problems of Colour III, National Physical Laboratory Symposium 8, H. M. Stationery Office, London (1958), pp. 387–398; C. R. Ingling, B. H.-P. Tsou, “Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977).
    [Crossref] [PubMed]
  10. The phenomena of small field tritanopia [G. S. Brindley, “The summation areas of human colour-receptive mechanisms at increment threshold,” J. Physiol. (London), 124, 400–408 (1954)] and foveal tritanopia [D. R. Williams, D. I. A. MacLeod, M. Hayhoe, “Foveal tritanopia,” Vision Res. 21, 1341–1356 (1981)] are well-known consequences of the relative scarcity of S cones in comparison with L and M cones, especially in the central foveola. Brief stimulus duration takes advantage of the supposed slow temporal response of S cones [D. O. Weitzman, J. A. S. Kinney, “Appearance of color for small, brief, spectral stimuli in the central fovea,”J. Opt. Soc. Am. 57, 665–670 (1967). Some recent evidence [A. Stockman, D. I. A. Mac-Leod, “Visible beats from invisible flickering lights: evidence that blue-sensitive cones respond to rapid flicker,” Invest. Ophthalmol. Vis. Sci. Suppl. 27, 71 (1986); A. Stockman, D. I. A. MacLeod, D. D. DePriest, “An inverted S-cone input to the luminance channel: evidence for two processes in S-cone flicker detection,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 92 (1987)] suggests that much of the limitation on temporal response is postreceptoral. However, the stage at which the limitation occurs is immaterial to the present study.
    [Crossref] [PubMed]
  11. D. V. Norren, P. Padmos, “Human and macaque blue cones studied with electroretinography,” Vision Res. 13, 1241–1254 (1973); D. Hood, N. I. Benimoff, V. C. Greenstein, “The response range of the blue-cone pathways: a source of vulnerability to disease,” Invest. Ophthalmol. Vis. Sci. 25, 864–867 (1984); M. Sawusch, J. Pokorny, V. C. Smith, “Clinical electroretinography for short wavelength sensitive cones,” Invest. Ophthalmol. Vis. Sci. 28, 966–974 (1987); A. Valberg, B. B. Lee, D. A. Tigwell, “Neurones with strong inhibitory S-cone inputs in the macaque lateral geniculate nucleus,” Vision Res. 26, 1061–1064 (1986).
    [Crossref] [PubMed]
  12. P. W. Trezona, “Rod participation in the ‘blue’ mechanism and its effect on colour matching,” Vision Res. 10, 317–332 (1970).
    [Crossref] [PubMed]
  13. All stimuli were presented in Maxwellian view through a 3-mm exit pupil. Test stimuli were produced with an Oriel monochromator. The foveally centered 15°-diameter background field had a 5500° color temperature (3400° tungsten light corrected with a Kodak Wratten 80B filter) and a retinal illuminance of 1000 photopic trolands (td) (~1300 scotopic Td). Rods become saturated, or unresponsive to increments of any magnitude, in the presence of adapting lights brighter than 1000 scotopic td [M. Aguilar, W. S. Stiles, “Saturation of the rod mechanism of the retina at high levels of stimulation,” Opt. Acta 1, 59–65 (1954); C. Blakemore, W. A. H. Rushton, “Dark adaptation and increment threshold in a rod monochromat,” J. Physiol. (London), 181, 612–628 (1965); B. Stabell, K. Nordby, U. Stabell, “Light-adaptation of the human rod system,” Clin. Vision Sci. 2, 83–91 (1987)].
    [Crossref]
  14. With one exception, the unique hue shifts for the other subjects were in the same direction as those in Fig. 2. One subject (DO) showed only small unique green shifts (5 – 15 nm) in response to reduced size of the foveal target but a large shift (~50 nm) in response to reduced duration. This relative lack of foveal tritanopia was confirmed by measurement of DO’s foveal S-cone sensitivity profile with a 10′-diameter, 0.5-sec, 440-nm stimulus on a 4-log-td, 580-nm background. His S-cone sensitivity, not including a measured macular pigment density of 0.25, is only 0.26 log unit lower in the foveal center than at 2° eccentricity, whereas subject BD, whose unique hue data are shown in Fig. 2, has a relative foveal S-cone sensitivity loss of more than 1.3 log. units, plus a macular pigment density of 0.63. These results imply a much higher foveal S-cone density for DO than for BD. (Macular pigment densities were measured at 440 nm by comparing the 2– 0° differences in 20-Hz flicker sensitivity for 440-and 550-nm stimuli of the same size and overall duration as. above.)
  15. H. G. Sperling, “The distribution of blue receptors in primates’ eyes revealed by spectral photic damage and by histochemical response experiments,” in Frontiers in Visual Science, S. J. Cool, E. L. Smith, eds. (Springer-Verlag, New York, 1978), Vol. 8, pp. 138–153.
    [Crossref]
  16. G. Haegerstrom-Portnoy, A. J. Adams, “Spatial sensitization of the B cone pathways,” Vision Res. 28, 629–638 (1988).
    [Crossref] [PubMed]
  17. D. McL. Purdy, “Spectral hue as a function of intensity,” Am. J. Psychol. 43, 541–559 (1931); R. M. Boynton, J. Gordon, “Bezold–Brücke hue shift measured by color-naming technique,”J. Opt. Soc. Am. 55, 78–86 (1965); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—I. Red/green equilibria,” Vision Res. 14, 1127–1140 (1974); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—II. Yellow/blue equilibria and nonlinear models,” Vision Res. 15, 723–731 (1975); A. L. Nagy, “Short-flash Bezold–Brücke hue shifts,” Vision Res., 20, 361–368 (1980); M. Ayama, T. Nakatsue, P. K. Kaiser, “Constant hue loci of unique and binary balanced hues at 10, 100, and 1000 td,” J. Opt. Soc. Am. A 4, 1136–1144 (1987).
    [Crossref] [PubMed]
  18. D. B. Judd, “Basic correlates of the visual stimulus,” in Handbook of Experimental Psychology, S. S. Stevens, ed. (Wiley, New York, 1951), pp. 811–867; L. M. Hurvich, D. Jameson, “Some quantitative aspects of an opponent-colors theory. II. Brightness, saturation, and hue in normal and dichromatic vision,”J. Opt. Soc. Am. 45, 602–616 (1955).
    [Crossref] [PubMed]
  19. J. D. Mollon, P. G. Polden [“Saturation of a retinal cone mechanism,” Nature 265, 243–246 (1977)] previously suggested that S-cone saturation might explain the loss of short-wavelength redness at high intensities.
    [Crossref] [PubMed]
  20. B. Drum, “Color naming of near-threshold monochromatic increments on a white background,” in Digest of Topical Meeting on Color Appearance (Optical Society of America, Washington, D.C., 1987), pp. 50–53; “Color naming of chromatic increments on a white background: implications for hue signals from single classes of cone,” Color Res. Appl. (to be published).
  21. P. Gouras, H. Eggers, “Ganglion cells mediating the signals of blue sensitive cones in primate retina detect white–yellow borders independently of brightness,” Vision Res. 22, 675–679 (1982); P. Gouras, H. U. Evers, “The role of the blue-sensitive cones in color vision,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 216 (1987).
    [Crossref]
  22. C. R. Ingling, “The spectral sensitivity of the opponent-color channels,” Vision Res. 17, 1083–1089 (1977).
    [Crossref] [PubMed]

1988 (1)

G. Haegerstrom-Portnoy, A. J. Adams, “Spatial sensitization of the B cone pathways,” Vision Res. 28, 629–638 (1988).
[Crossref] [PubMed]

1982 (1)

P. Gouras, H. Eggers, “Ganglion cells mediating the signals of blue sensitive cones in primate retina detect white–yellow borders independently of brightness,” Vision Res. 22, 675–679 (1982); P. Gouras, H. U. Evers, “The role of the blue-sensitive cones in color vision,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 216 (1987).
[Crossref]

1977 (2)

C. R. Ingling, “The spectral sensitivity of the opponent-color channels,” Vision Res. 17, 1083–1089 (1977).
[Crossref] [PubMed]

J. D. Mollon, P. G. Polden [“Saturation of a retinal cone mechanism,” Nature 265, 243–246 (1977)] previously suggested that S-cone saturation might explain the loss of short-wavelength redness at high intensities.
[Crossref] [PubMed]

1973 (1)

D. V. Norren, P. Padmos, “Human and macaque blue cones studied with electroretinography,” Vision Res. 13, 1241–1254 (1973); D. Hood, N. I. Benimoff, V. C. Greenstein, “The response range of the blue-cone pathways: a source of vulnerability to disease,” Invest. Ophthalmol. Vis. Sci. 25, 864–867 (1984); M. Sawusch, J. Pokorny, V. C. Smith, “Clinical electroretinography for short wavelength sensitive cones,” Invest. Ophthalmol. Vis. Sci. 28, 966–974 (1987); A. Valberg, B. B. Lee, D. A. Tigwell, “Neurones with strong inhibitory S-cone inputs in the macaque lateral geniculate nucleus,” Vision Res. 26, 1061–1064 (1986).
[Crossref] [PubMed]

1970 (1)

P. W. Trezona, “Rod participation in the ‘blue’ mechanism and its effect on colour matching,” Vision Res. 10, 317–332 (1970).
[Crossref] [PubMed]

1966 (1)

C. H. Graham, Y. Hsia, F. F. Stephan, “Visual discriminations of a subject with acquired unilateral tritanopia,” Vision Res. 6, 469–479 (1966); N. Ohba, T. Tanino, “Unilateral colour vision defect resembling tritanopia,” Mod. Probl. Ophthalmol. 17, 331–335 (1976); M. Alpern, K. Kitahara, D. H. Krantz, “Perception of colour in unilateral tritanopia,”J. Physiol. (London) 335, 683–697 (1983).
[PubMed]

1964 (1)

R. M. Boynton, W. Schafer, M. A. Neun, “Hue-wavelength relation measured by color-naming method for three retinal locations,” Science 146, 666–668 (1964); J. Gordon, I. Abramov, “Color vision in the peripheral retina. II. Hue and saturation,”J. Opt. Soc. Am. 67, 202–207 (1977).
[Crossref] [PubMed]

1954 (2)

All stimuli were presented in Maxwellian view through a 3-mm exit pupil. Test stimuli were produced with an Oriel monochromator. The foveally centered 15°-diameter background field had a 5500° color temperature (3400° tungsten light corrected with a Kodak Wratten 80B filter) and a retinal illuminance of 1000 photopic trolands (td) (~1300 scotopic Td). Rods become saturated, or unresponsive to increments of any magnitude, in the presence of adapting lights brighter than 1000 scotopic td [M. Aguilar, W. S. Stiles, “Saturation of the rod mechanism of the retina at high levels of stimulation,” Opt. Acta 1, 59–65 (1954); C. Blakemore, W. A. H. Rushton, “Dark adaptation and increment threshold in a rod monochromat,” J. Physiol. (London), 181, 612–628 (1965); B. Stabell, K. Nordby, U. Stabell, “Light-adaptation of the human rod system,” Clin. Vision Sci. 2, 83–91 (1987)].
[Crossref]

The phenomena of small field tritanopia [G. S. Brindley, “The summation areas of human colour-receptive mechanisms at increment threshold,” J. Physiol. (London), 124, 400–408 (1954)] and foveal tritanopia [D. R. Williams, D. I. A. MacLeod, M. Hayhoe, “Foveal tritanopia,” Vision Res. 21, 1341–1356 (1981)] are well-known consequences of the relative scarcity of S cones in comparison with L and M cones, especially in the central foveola. Brief stimulus duration takes advantage of the supposed slow temporal response of S cones [D. O. Weitzman, J. A. S. Kinney, “Appearance of color for small, brief, spectral stimuli in the central fovea,”J. Opt. Soc. Am. 57, 665–670 (1967). Some recent evidence [A. Stockman, D. I. A. Mac-Leod, “Visible beats from invisible flickering lights: evidence that blue-sensitive cones respond to rapid flicker,” Invest. Ophthalmol. Vis. Sci. Suppl. 27, 71 (1986); A. Stockman, D. I. A. MacLeod, D. D. DePriest, “An inverted S-cone input to the luminance channel: evidence for two processes in S-cone flicker detection,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 92 (1987)] suggests that much of the limitation on temporal response is postreceptoral. However, the stage at which the limitation occurs is immaterial to the present study.
[Crossref] [PubMed]

1951 (1)

1949 (1)

1931 (1)

D. McL. Purdy, “Spectral hue as a function of intensity,” Am. J. Psychol. 43, 541–559 (1931); R. M. Boynton, J. Gordon, “Bezold–Brücke hue shift measured by color-naming technique,”J. Opt. Soc. Am. 55, 78–86 (1965); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—I. Red/green equilibria,” Vision Res. 14, 1127–1140 (1974); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—II. Yellow/blue equilibria and nonlinear models,” Vision Res. 15, 723–731 (1975); A. L. Nagy, “Short-flash Bezold–Brücke hue shifts,” Vision Res., 20, 361–368 (1980); M. Ayama, T. Nakatsue, P. K. Kaiser, “Constant hue loci of unique and binary balanced hues at 10, 100, and 1000 td,” J. Opt. Soc. Am. A 4, 1136–1144 (1987).
[Crossref] [PubMed]

Adams, A. J.

G. Haegerstrom-Portnoy, A. J. Adams, “Spatial sensitization of the B cone pathways,” Vision Res. 28, 629–638 (1988).
[Crossref] [PubMed]

Aguilar, M.

All stimuli were presented in Maxwellian view through a 3-mm exit pupil. Test stimuli were produced with an Oriel monochromator. The foveally centered 15°-diameter background field had a 5500° color temperature (3400° tungsten light corrected with a Kodak Wratten 80B filter) and a retinal illuminance of 1000 photopic trolands (td) (~1300 scotopic Td). Rods become saturated, or unresponsive to increments of any magnitude, in the presence of adapting lights brighter than 1000 scotopic td [M. Aguilar, W. S. Stiles, “Saturation of the rod mechanism of the retina at high levels of stimulation,” Opt. Acta 1, 59–65 (1954); C. Blakemore, W. A. H. Rushton, “Dark adaptation and increment threshold in a rod monochromat,” J. Physiol. (London), 181, 612–628 (1965); B. Stabell, K. Nordby, U. Stabell, “Light-adaptation of the human rod system,” Clin. Vision Sci. 2, 83–91 (1987)].
[Crossref]

Boynton, R. M.

R. M. Boynton, W. Schafer, M. A. Neun, “Hue-wavelength relation measured by color-naming method for three retinal locations,” Science 146, 666–668 (1964); J. Gordon, I. Abramov, “Color vision in the peripheral retina. II. Hue and saturation,”J. Opt. Soc. Am. 67, 202–207 (1977).
[Crossref] [PubMed]

Brindley, G. S.

The phenomena of small field tritanopia [G. S. Brindley, “The summation areas of human colour-receptive mechanisms at increment threshold,” J. Physiol. (London), 124, 400–408 (1954)] and foveal tritanopia [D. R. Williams, D. I. A. MacLeod, M. Hayhoe, “Foveal tritanopia,” Vision Res. 21, 1341–1356 (1981)] are well-known consequences of the relative scarcity of S cones in comparison with L and M cones, especially in the central foveola. Brief stimulus duration takes advantage of the supposed slow temporal response of S cones [D. O. Weitzman, J. A. S. Kinney, “Appearance of color for small, brief, spectral stimuli in the central fovea,”J. Opt. Soc. Am. 57, 665–670 (1967). Some recent evidence [A. Stockman, D. I. A. Mac-Leod, “Visible beats from invisible flickering lights: evidence that blue-sensitive cones respond to rapid flicker,” Invest. Ophthalmol. Vis. Sci. Suppl. 27, 71 (1986); A. Stockman, D. I. A. MacLeod, D. D. DePriest, “An inverted S-cone input to the luminance channel: evidence for two processes in S-cone flicker detection,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 92 (1987)] suggests that much of the limitation on temporal response is postreceptoral. However, the stage at which the limitation occurs is immaterial to the present study.
[Crossref] [PubMed]

Cruz, A.

M. Dagher, A. Cruz, L. Plaza, “Colour thresholds with monochromatic stimuli in the spectral region 530–630 μ m,” in Visual Problems of Colour III, National Physical Laboratory Symposium 8, H. M. Stationery Office, London (1958), pp. 387–398; C. R. Ingling, B. H.-P. Tsou, “Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977).
[Crossref] [PubMed]

Dagher, M.

M. Dagher, A. Cruz, L. Plaza, “Colour thresholds with monochromatic stimuli in the spectral region 530–630 μ m,” in Visual Problems of Colour III, National Physical Laboratory Symposium 8, H. M. Stationery Office, London (1958), pp. 387–398; C. R. Ingling, B. H.-P. Tsou, “Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977).
[Crossref] [PubMed]

Drum, B.

B. Drum, “Color naming of near-threshold monochromatic increments on a white background,” in Digest of Topical Meeting on Color Appearance (Optical Society of America, Washington, D.C., 1987), pp. 50–53; “Color naming of chromatic increments on a white background: implications for hue signals from single classes of cone,” Color Res. Appl. (to be published).

Eggers, H.

P. Gouras, H. Eggers, “Ganglion cells mediating the signals of blue sensitive cones in primate retina detect white–yellow borders independently of brightness,” Vision Res. 22, 675–679 (1982); P. Gouras, H. U. Evers, “The role of the blue-sensitive cones in color vision,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 216 (1987).
[Crossref]

Gouras, P.

P. Gouras, H. Eggers, “Ganglion cells mediating the signals of blue sensitive cones in primate retina detect white–yellow borders independently of brightness,” Vision Res. 22, 675–679 (1982); P. Gouras, H. U. Evers, “The role of the blue-sensitive cones in color vision,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 216 (1987).
[Crossref]

Graham, C. H.

C. H. Graham, Y. Hsia, F. F. Stephan, “Visual discriminations of a subject with acquired unilateral tritanopia,” Vision Res. 6, 469–479 (1966); N. Ohba, T. Tanino, “Unilateral colour vision defect resembling tritanopia,” Mod. Probl. Ophthalmol. 17, 331–335 (1976); M. Alpern, K. Kitahara, D. H. Krantz, “Perception of colour in unilateral tritanopia,”J. Physiol. (London) 335, 683–697 (1983).
[PubMed]

Haegerstrom-Portnoy, G.

G. Haegerstrom-Portnoy, A. J. Adams, “Spatial sensitization of the B cone pathways,” Vision Res. 28, 629–638 (1988).
[Crossref] [PubMed]

Holmes, M. C.

Hsia, Y.

C. H. Graham, Y. Hsia, F. F. Stephan, “Visual discriminations of a subject with acquired unilateral tritanopia,” Vision Res. 6, 469–479 (1966); N. Ohba, T. Tanino, “Unilateral colour vision defect resembling tritanopia,” Mod. Probl. Ophthalmol. 17, 331–335 (1976); M. Alpern, K. Kitahara, D. H. Krantz, “Perception of colour in unilateral tritanopia,”J. Physiol. (London) 335, 683–697 (1983).
[PubMed]

Hurvich, L. M.

Representative examples of this model can be found in L. M. Hurvich, Color Vision (Sinauer, New York, 1982), pp. 128–135, and R. M. Boynton, Human Color Vision (Holt, Rinehart and Winston, New York, 1979), pp. 211–215. A notable exception is the model of C. R. Ingling, B. H.-P. Tsou [“Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977)], in which M cones also contribute to the blueness signal.
[Crossref] [PubMed]

Ingling, C. R.

C. R. Ingling, “The spectral sensitivity of the opponent-color channels,” Vision Res. 17, 1083–1089 (1977).
[Crossref] [PubMed]

Judd, D. B.

D. B. Judd, “Basic correlates of the visual stimulus,” in Handbook of Experimental Psychology, S. S. Stevens, ed. (Wiley, New York, 1951), pp. 811–867; L. M. Hurvich, D. Jameson, “Some quantitative aspects of an opponent-colors theory. II. Brightness, saturation, and hue in normal and dichromatic vision,”J. Opt. Soc. Am. 45, 602–616 (1955).
[Crossref] [PubMed]

Mayo, E. G.

Middleton, W. E. K.

Mollon, J. D.

J. D. Mollon, P. G. Polden [“Saturation of a retinal cone mechanism,” Nature 265, 243–246 (1977)] previously suggested that S-cone saturation might explain the loss of short-wavelength redness at high intensities.
[Crossref] [PubMed]

Neun, M. A.

R. M. Boynton, W. Schafer, M. A. Neun, “Hue-wavelength relation measured by color-naming method for three retinal locations,” Science 146, 666–668 (1964); J. Gordon, I. Abramov, “Color vision in the peripheral retina. II. Hue and saturation,”J. Opt. Soc. Am. 67, 202–207 (1977).
[Crossref] [PubMed]

Norren, D. V.

D. V. Norren, P. Padmos, “Human and macaque blue cones studied with electroretinography,” Vision Res. 13, 1241–1254 (1973); D. Hood, N. I. Benimoff, V. C. Greenstein, “The response range of the blue-cone pathways: a source of vulnerability to disease,” Invest. Ophthalmol. Vis. Sci. 25, 864–867 (1984); M. Sawusch, J. Pokorny, V. C. Smith, “Clinical electroretinography for short wavelength sensitive cones,” Invest. Ophthalmol. Vis. Sci. 28, 966–974 (1987); A. Valberg, B. B. Lee, D. A. Tigwell, “Neurones with strong inhibitory S-cone inputs in the macaque lateral geniculate nucleus,” Vision Res. 26, 1061–1064 (1986).
[Crossref] [PubMed]

Padmos, P.

D. V. Norren, P. Padmos, “Human and macaque blue cones studied with electroretinography,” Vision Res. 13, 1241–1254 (1973); D. Hood, N. I. Benimoff, V. C. Greenstein, “The response range of the blue-cone pathways: a source of vulnerability to disease,” Invest. Ophthalmol. Vis. Sci. 25, 864–867 (1984); M. Sawusch, J. Pokorny, V. C. Smith, “Clinical electroretinography for short wavelength sensitive cones,” Invest. Ophthalmol. Vis. Sci. 28, 966–974 (1987); A. Valberg, B. B. Lee, D. A. Tigwell, “Neurones with strong inhibitory S-cone inputs in the macaque lateral geniculate nucleus,” Vision Res. 26, 1061–1064 (1986).
[Crossref] [PubMed]

Plaza, L.

M. Dagher, A. Cruz, L. Plaza, “Colour thresholds with monochromatic stimuli in the spectral region 530–630 μ m,” in Visual Problems of Colour III, National Physical Laboratory Symposium 8, H. M. Stationery Office, London (1958), pp. 387–398; C. R. Ingling, B. H.-P. Tsou, “Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977).
[Crossref] [PubMed]

Polden, P. G.

J. D. Mollon, P. G. Polden [“Saturation of a retinal cone mechanism,” Nature 265, 243–246 (1977)] previously suggested that S-cone saturation might explain the loss of short-wavelength redness at high intensities.
[Crossref] [PubMed]

Purdy, D. McL.

D. McL. Purdy, “Spectral hue as a function of intensity,” Am. J. Psychol. 43, 541–559 (1931); R. M. Boynton, J. Gordon, “Bezold–Brücke hue shift measured by color-naming technique,”J. Opt. Soc. Am. 55, 78–86 (1965); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—I. Red/green equilibria,” Vision Res. 14, 1127–1140 (1974); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—II. Yellow/blue equilibria and nonlinear models,” Vision Res. 15, 723–731 (1975); A. L. Nagy, “Short-flash Bezold–Brücke hue shifts,” Vision Res., 20, 361–368 (1980); M. Ayama, T. Nakatsue, P. K. Kaiser, “Constant hue loci of unique and binary balanced hues at 10, 100, and 1000 td,” J. Opt. Soc. Am. A 4, 1136–1144 (1987).
[Crossref] [PubMed]

Schafer, W.

R. M. Boynton, W. Schafer, M. A. Neun, “Hue-wavelength relation measured by color-naming method for three retinal locations,” Science 146, 666–668 (1964); J. Gordon, I. Abramov, “Color vision in the peripheral retina. II. Hue and saturation,”J. Opt. Soc. Am. 67, 202–207 (1977).
[Crossref] [PubMed]

Sperling, H. G.

H. G. Sperling, “The distribution of blue receptors in primates’ eyes revealed by spectral photic damage and by histochemical response experiments,” in Frontiers in Visual Science, S. J. Cool, E. L. Smith, eds. (Springer-Verlag, New York, 1978), Vol. 8, pp. 138–153.
[Crossref]

Stephan, F. F.

C. H. Graham, Y. Hsia, F. F. Stephan, “Visual discriminations of a subject with acquired unilateral tritanopia,” Vision Res. 6, 469–479 (1966); N. Ohba, T. Tanino, “Unilateral colour vision defect resembling tritanopia,” Mod. Probl. Ophthalmol. 17, 331–335 (1976); M. Alpern, K. Kitahara, D. H. Krantz, “Perception of colour in unilateral tritanopia,”J. Physiol. (London) 335, 683–697 (1983).
[PubMed]

Stiles, W. S.

All stimuli were presented in Maxwellian view through a 3-mm exit pupil. Test stimuli were produced with an Oriel monochromator. The foveally centered 15°-diameter background field had a 5500° color temperature (3400° tungsten light corrected with a Kodak Wratten 80B filter) and a retinal illuminance of 1000 photopic trolands (td) (~1300 scotopic Td). Rods become saturated, or unresponsive to increments of any magnitude, in the presence of adapting lights brighter than 1000 scotopic td [M. Aguilar, W. S. Stiles, “Saturation of the rod mechanism of the retina at high levels of stimulation,” Opt. Acta 1, 59–65 (1954); C. Blakemore, W. A. H. Rushton, “Dark adaptation and increment threshold in a rod monochromat,” J. Physiol. (London), 181, 612–628 (1965); B. Stabell, K. Nordby, U. Stabell, “Light-adaptation of the human rod system,” Clin. Vision Sci. 2, 83–91 (1987)].
[Crossref]

Trezona, P. W.

P. W. Trezona, “Rod participation in the ‘blue’ mechanism and its effect on colour matching,” Vision Res. 10, 317–332 (1970).
[Crossref] [PubMed]

Am. J. Psychol. (1)

D. McL. Purdy, “Spectral hue as a function of intensity,” Am. J. Psychol. 43, 541–559 (1931); R. M. Boynton, J. Gordon, “Bezold–Brücke hue shift measured by color-naming technique,”J. Opt. Soc. Am. 55, 78–86 (1965); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—I. Red/green equilibria,” Vision Res. 14, 1127–1140 (1974); J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent process additivity—II. Yellow/blue equilibria and nonlinear models,” Vision Res. 15, 723–731 (1975); A. L. Nagy, “Short-flash Bezold–Brücke hue shifts,” Vision Res., 20, 361–368 (1980); M. Ayama, T. Nakatsue, P. K. Kaiser, “Constant hue loci of unique and binary balanced hues at 10, 100, and 1000 td,” J. Opt. Soc. Am. A 4, 1136–1144 (1987).
[Crossref] [PubMed]

J. Opt. Soc. Am. (2)

J. Physiol. (London) (1)

The phenomena of small field tritanopia [G. S. Brindley, “The summation areas of human colour-receptive mechanisms at increment threshold,” J. Physiol. (London), 124, 400–408 (1954)] and foveal tritanopia [D. R. Williams, D. I. A. MacLeod, M. Hayhoe, “Foveal tritanopia,” Vision Res. 21, 1341–1356 (1981)] are well-known consequences of the relative scarcity of S cones in comparison with L and M cones, especially in the central foveola. Brief stimulus duration takes advantage of the supposed slow temporal response of S cones [D. O. Weitzman, J. A. S. Kinney, “Appearance of color for small, brief, spectral stimuli in the central fovea,”J. Opt. Soc. Am. 57, 665–670 (1967). Some recent evidence [A. Stockman, D. I. A. Mac-Leod, “Visible beats from invisible flickering lights: evidence that blue-sensitive cones respond to rapid flicker,” Invest. Ophthalmol. Vis. Sci. Suppl. 27, 71 (1986); A. Stockman, D. I. A. MacLeod, D. D. DePriest, “An inverted S-cone input to the luminance channel: evidence for two processes in S-cone flicker detection,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 92 (1987)] suggests that much of the limitation on temporal response is postreceptoral. However, the stage at which the limitation occurs is immaterial to the present study.
[Crossref] [PubMed]

Nature (1)

J. D. Mollon, P. G. Polden [“Saturation of a retinal cone mechanism,” Nature 265, 243–246 (1977)] previously suggested that S-cone saturation might explain the loss of short-wavelength redness at high intensities.
[Crossref] [PubMed]

Opt. Acta (1)

All stimuli were presented in Maxwellian view through a 3-mm exit pupil. Test stimuli were produced with an Oriel monochromator. The foveally centered 15°-diameter background field had a 5500° color temperature (3400° tungsten light corrected with a Kodak Wratten 80B filter) and a retinal illuminance of 1000 photopic trolands (td) (~1300 scotopic Td). Rods become saturated, or unresponsive to increments of any magnitude, in the presence of adapting lights brighter than 1000 scotopic td [M. Aguilar, W. S. Stiles, “Saturation of the rod mechanism of the retina at high levels of stimulation,” Opt. Acta 1, 59–65 (1954); C. Blakemore, W. A. H. Rushton, “Dark adaptation and increment threshold in a rod monochromat,” J. Physiol. (London), 181, 612–628 (1965); B. Stabell, K. Nordby, U. Stabell, “Light-adaptation of the human rod system,” Clin. Vision Sci. 2, 83–91 (1987)].
[Crossref]

Science (1)

R. M. Boynton, W. Schafer, M. A. Neun, “Hue-wavelength relation measured by color-naming method for three retinal locations,” Science 146, 666–668 (1964); J. Gordon, I. Abramov, “Color vision in the peripheral retina. II. Hue and saturation,”J. Opt. Soc. Am. 67, 202–207 (1977).
[Crossref] [PubMed]

Vision Res. (6)

D. V. Norren, P. Padmos, “Human and macaque blue cones studied with electroretinography,” Vision Res. 13, 1241–1254 (1973); D. Hood, N. I. Benimoff, V. C. Greenstein, “The response range of the blue-cone pathways: a source of vulnerability to disease,” Invest. Ophthalmol. Vis. Sci. 25, 864–867 (1984); M. Sawusch, J. Pokorny, V. C. Smith, “Clinical electroretinography for short wavelength sensitive cones,” Invest. Ophthalmol. Vis. Sci. 28, 966–974 (1987); A. Valberg, B. B. Lee, D. A. Tigwell, “Neurones with strong inhibitory S-cone inputs in the macaque lateral geniculate nucleus,” Vision Res. 26, 1061–1064 (1986).
[Crossref] [PubMed]

P. W. Trezona, “Rod participation in the ‘blue’ mechanism and its effect on colour matching,” Vision Res. 10, 317–332 (1970).
[Crossref] [PubMed]

C. H. Graham, Y. Hsia, F. F. Stephan, “Visual discriminations of a subject with acquired unilateral tritanopia,” Vision Res. 6, 469–479 (1966); N. Ohba, T. Tanino, “Unilateral colour vision defect resembling tritanopia,” Mod. Probl. Ophthalmol. 17, 331–335 (1976); M. Alpern, K. Kitahara, D. H. Krantz, “Perception of colour in unilateral tritanopia,”J. Physiol. (London) 335, 683–697 (1983).
[PubMed]

G. Haegerstrom-Portnoy, A. J. Adams, “Spatial sensitization of the B cone pathways,” Vision Res. 28, 629–638 (1988).
[Crossref] [PubMed]

P. Gouras, H. Eggers, “Ganglion cells mediating the signals of blue sensitive cones in primate retina detect white–yellow borders independently of brightness,” Vision Res. 22, 675–679 (1982); P. Gouras, H. U. Evers, “The role of the blue-sensitive cones in color vision,” Invest. Ophthalmol. Vis. Sci. Suppl. 28, 216 (1987).
[Crossref]

C. R. Ingling, “The spectral sensitivity of the opponent-color channels,” Vision Res. 17, 1083–1089 (1977).
[Crossref] [PubMed]

Other (9)

B. Drum, “Color naming of near-threshold monochromatic increments on a white background,” in Digest of Topical Meeting on Color Appearance (Optical Society of America, Washington, D.C., 1987), pp. 50–53; “Color naming of chromatic increments on a white background: implications for hue signals from single classes of cone,” Color Res. Appl. (to be published).

D. B. Judd, “Basic correlates of the visual stimulus,” in Handbook of Experimental Psychology, S. S. Stevens, ed. (Wiley, New York, 1951), pp. 811–867; L. M. Hurvich, D. Jameson, “Some quantitative aspects of an opponent-colors theory. II. Brightness, saturation, and hue in normal and dichromatic vision,”J. Opt. Soc. Am. 45, 602–616 (1955).
[Crossref] [PubMed]

With one exception, the unique hue shifts for the other subjects were in the same direction as those in Fig. 2. One subject (DO) showed only small unique green shifts (5 – 15 nm) in response to reduced size of the foveal target but a large shift (~50 nm) in response to reduced duration. This relative lack of foveal tritanopia was confirmed by measurement of DO’s foveal S-cone sensitivity profile with a 10′-diameter, 0.5-sec, 440-nm stimulus on a 4-log-td, 580-nm background. His S-cone sensitivity, not including a measured macular pigment density of 0.25, is only 0.26 log unit lower in the foveal center than at 2° eccentricity, whereas subject BD, whose unique hue data are shown in Fig. 2, has a relative foveal S-cone sensitivity loss of more than 1.3 log. units, plus a macular pigment density of 0.63. These results imply a much higher foveal S-cone density for DO than for BD. (Macular pigment densities were measured at 440 nm by comparing the 2– 0° differences in 20-Hz flicker sensitivity for 440-and 550-nm stimuli of the same size and overall duration as. above.)

H. G. Sperling, “The distribution of blue receptors in primates’ eyes revealed by spectral photic damage and by histochemical response experiments,” in Frontiers in Visual Science, S. J. Cool, E. L. Smith, eds. (Springer-Verlag, New York, 1978), Vol. 8, pp. 138–153.
[Crossref]

M. Dagher, A. Cruz, L. Plaza, “Colour thresholds with monochromatic stimuli in the spectral region 530–630 μ m,” in Visual Problems of Colour III, National Physical Laboratory Symposium 8, H. M. Stationery Office, London (1958), pp. 387–398; C. R. Ingling, B. H.-P. Tsou, “Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977).
[Crossref] [PubMed]

Representative examples of this model can be found in L. M. Hurvich, Color Vision (Sinauer, New York, 1982), pp. 128–135, and R. M. Boynton, Human Color Vision (Holt, Rinehart and Winston, New York, 1979), pp. 211–215. A notable exception is the model of C. R. Ingling, B. H.-P. Tsou [“Orthogonal combination of the three visual channels,” Vision Res. 17, 1075–1082 (1977)], in which M cones also contribute to the blueness signal.
[Crossref] [PubMed]

The univariance of single cone types implies only that the responses to any two wavelengths can be made identical with suitable adjustments of intensity. It does not imply a lack of perceived color when only one cone type is stimulated. For example, it is easy to demonstrate with long-wavelength foveal stimuli in the dark-adapted eye that exclusive L-cone stimulation produces a predominately reddish hue sensation.

The unique hues are defined as lights that produce perceptually pure sensations containing only one of the four fundamental hues of red, yellow, green, or blue. Along with the achromatic colors (white, gray, and black), they compose the set of equilibrium hues for which the response of at least one of the color-opponent systems is zero. Thus unique red and unique green are yellow–blue equilibria and unique yellow and unique blue are red–green equilibria.

This paper is concerned with manipulating relative cone contributions to color perception, which is not necessarily the same as manipulating relative responses of the cones themselves. Perception does not occur at the photoreceptors but only after many intervening stages of neural processing that may substantially alter the space-, time-, and intensity-dependent properties of the visual signal. Thus the term cone sensitivity is used here to mean the sensitivity of the entire visual system to stimuli that produce responses in one or more of the L-, M-, and S-cone classes.

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Figures (2)

Fig. 1
Fig. 1

A, Conventional wiring-diagram model of transformations from cone responses to hue sensations. Triangular boxes indicate L-, M-, and S-cone types. Centers of red (R), yellow (Y), green (G), and blue (B) color sensations are indicated by square boxes linked by opponent operators, which are shown as circled minuses. These operators do not necessarily represent linear subtraction but merely mutual exclusivity between opponent hues. Connecting lines indicate contributions from cone types to hue centers. Transformations for achromatic sensations are not included. B, Modified color-transformation model, incorporating an M-cone contribution to blueness (heavy solid line) and a special indirect S-cone contribution to yellowness (heavy dotted line).

Fig. 2
Fig. 2

Unique blue (open symbols) and unique green (filled symbols) wavelengths for one normal subject. Each data symbol indicates the mean of three to seven measurements. Error bars indicate standard deviations greater than 2 nm. A, Foveal measurements for l-sec-duration stimuli as a function of diameter (squares) and 50-msec-duration stimuli at 10′ diameter (circles). B, Measurements for 10′-diameter, l-sec-duration stimuli at 0 and 1° eccentricity. Foveal data are copied from A. C, Measurements for foveal, 10′-diameter, 1-sec-duration stimuli as a function of luminance above threshold.

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