Abstract

Spectral sensitivity in the red–green spectral range typically reflects the joint influence of the middle-wavelength-sensitive cones (the M or green cones) and long-wavelength-sensitive cones (the L or red cones). The balance of M- and L-cone influence can be altered by presenting the test lights superimposed upon steady background fields of long or short wavelength. We find that presenting test stimuli just after an abrupt exchange between two colored backgrounds permits an easier and closer approach to cone isolation than presenting them either on a steady background or following an intense bleach. Background exchange drives the flicker detection or flicker photometric spectral sensitivities measured at 17 Hz to a limiting condition at lower intensities than do steady backgrounds. This condition is consistent with either M- or L-cone isolation. Steady backgrounds do not produce complete cone isolation: even on backgrounds that push spectral sensitivity closest to M or L, there are substantial phase differences between flickering lights of different color. In contrast, no phase differences remain following background exchange. The improvement in cone isolation produced by the exchange procedure is not confined to flicker measurements: the spectral range over which subjects are temporarily monochromatic is more extended following background exchange than on steady fields.

© 1993 Optical Society of America

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References

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  32. G. S. Brindley, “The colour of light of very long wavelength,”J. Physiol. (London) 130, 35–44 (1955).
  33. A. Eisner, D. I. A. MacLeod, “Blue-sensitive cones do not contribute to luminance,”J. Opt. Soc. Am. 70, 121–123 (1980).
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  34. A. Stockman, D. I. A. MacLeod, “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).
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    [CrossRef]
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    [CrossRef]
  39. W. A. H. Rushton, G. H. Henry, “Bleaching and regeneration of cone pigments in man,” Vision Res. 8, 617–631 (1968).
    [CrossRef]
  40. H. L. De Vries, “The luminosity curve of the eye as determined by measurements with the flicker photometer,” Physika 14, 319–348 (1948).
  41. A. Eisner, D. I. A. MacLeod, “Flicker photometric study of chromatic adaptation: selective suppression of cone inputs by colored backgrounds,”J. Opt. Soc. Am. 71, 705–718 (1981).
    [CrossRef] [PubMed]
  42. G. S. Brindley, “The effects on colour vision of adaptation to very bright lights,”J. Physiol. (London) 122, 332–350 (1953).
  43. G. Burch, “On artificial temporary colour-blindness, with an examination of the colour sensations of 109 persons,” Phil. Trans. R. Soc. London Ser. B 191, 1–34 (1898).
    [CrossRef]
  44. D. I. A. MacLeod, A. Stockman, J. A. Vivian, “Flicker photometric approximations to cone sensitivities under bleaching and transient adaptation conditions,” Perception 14, A17 (1985).
  45. A. Stockman, E. D. Montag, D. I. A. MacLeod, “Large changes in phase delay on intense bleaching backgrounds,” Invest. Ophthalmol. Vis. Sci. Suppl. 32, 841 (1991).
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    [CrossRef]
  47. C. E. Sternheim, G. A. Gorinson, N. Markovitz, “Visual sensitivity during successive chromatic contrast: evidence for interactions between photopic mechanisms,” Vision Res. 17, 45–49 (1977).
    [CrossRef]
  48. A. Reeves, “Exchange thresholds for green tests,” Vision Res. 22, 961–966 (1982).
    [CrossRef] [PubMed]
  49. A. Reeves, “Exchange threshold for long-wavelength incremental flashes,”J. Opt. Soc. Am. 72, 565–570 (1982).
    [CrossRef]
  50. W. H. Swanson, J. Pokorny, V. C. Smith, “Effects of chromatic adaptation on phase-dependent sensitivity to heterochromatic flicker,” J. Opt. Soc. Am. A 5, 1976–1982 (1988).
    [CrossRef] [PubMed]
  51. C. F. Stromeyer, G. R. Cole, R. E. Kronauer, “Chromatic suppression of cone inputs to the luminance flicker mechanisms,” Vision Res. 27, 1113–1137 (1987).
    [CrossRef]
  52. P. E. King-Smith, D. Carden, “Luminance and opponent-color contributions to visual detection and adaptation and to temporal and spatial integration,”J. Opt. Soc. Am. 66, 709–717 (1976).
    [CrossRef] [PubMed]
  53. G. Wald, “The receptors of human color vision,” Science 145, 1007–1016 (1964).
    [CrossRef] [PubMed]
  54. J. M. Enoch, “The two-color threshold technique of Stiles and derived component color mechanisms,” in Handbook of Sensory Physiology, D. Jameson, L. H. Hurvich, eds. (Springer-Verlag, Berlin, 1972).
    [CrossRef]
  55. R. M. Boynton, M. Ikeda, W. S. Stiles, “Interactions among chromatic mechanisms as inferred from positive and negative increment thresholds,” Vision Res. 4, 87–117 (1964).
    [CrossRef] [PubMed]
  56. M. Ikeda, M. Urakubo, “Flicker HRTF as test of color vision,”J. Opt. Soc. Am. 58, 27–31 (1968).
    [CrossRef] [PubMed]
  57. T. Yeh, V. C. Smith, J. Pokorny, “The effect of background luminance on cone sensitivity function,” Invest. Ophthalmol. Vis. Sci. 30, 2077–2086 (1989).
    [PubMed]
  58. W. H. Swanson, J. Pokorny, V. C. Smith, “Effects of temporal frequency on phase-dependent sensitivity to heterochromatic flicker,” J. Opt. Soc. Am. A 4, 2266–2273 (1988).
    [CrossRef]

1993 (1)

1991 (2)

A. Stockman, D. I. A. MacLeod, D. D. DePriest, “The temporal properties of the human short-wave photoreceptors and their associated pathways,” Vision Res. 31, 189–208 (1991).
[CrossRef]

A. Stockman, E. D. Montag, D. I. A. MacLeod, “Large changes in phase delay on intense bleaching backgrounds,” Invest. Ophthalmol. Vis. Sci. Suppl. 32, 841 (1991).

1989 (2)

J. Lee, C. F. Stromeyer, “Contribution of human short-wave cones to luminance and motion detection,”J. Physiol. (London) 413, 563–593 (1989).

T. Yeh, V. C. Smith, J. Pokorny, “The effect of background luminance on cone sensitivity function,” Invest. Ophthalmol. Vis. Sci. 30, 2077–2086 (1989).
[PubMed]

1988 (2)

1987 (2)

C. F. Stromeyer, G. R. Cole, R. E. Kronauer, “Chromatic suppression of cone inputs to the luminance flicker mechanisms,” Vision Res. 27, 1113–1137 (1987).
[CrossRef]

A. Stockman, D. I. A. MacLeod, “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).

1986 (1)

A. Stockman, J. D. Mollon, “The spectral sensitivities of the middle- and long-wavelength cones: an extension of the two-colour threshold technique of W. S. Stiles,” Perception 15, 729–754 (1986).
[CrossRef]

1985 (1)

D. I. A. MacLeod, A. Stockman, J. A. Vivian, “Flicker photometric approximations to cone sensitivities under bleaching and transient adaptation conditions,” Perception 14, A17 (1985).

1983 (1)

H. J. A. Dartnall, J. K. Bowmaker, J. D. Mollon, “Human visual pigments: microspectrophotometric results from the eyes of seven persons,” Proc. R. Soc. London Ser. B 220, 115–130 (1983).
[CrossRef]

1982 (3)

J. D. Mollon, “Color vision,” Ann. Rev. Psychol. 33, 41–85 (1982).
[CrossRef]

A. Reeves, “Exchange thresholds for green tests,” Vision Res. 22, 961–966 (1982).
[CrossRef] [PubMed]

A. Reeves, “Exchange threshold for long-wavelength incremental flashes,”J. Opt. Soc. Am. 72, 565–570 (1982).
[CrossRef]

1981 (1)

1980 (3)

1978 (1)

J. J. Vos, “Colorimetric and photometric properties of a 2-deg fundamental observer,” Color Res. Appl. 3, 125–128 (1978).
[CrossRef]

1977 (1)

C. E. Sternheim, G. A. Gorinson, N. Markovitz, “Visual sensitivity during successive chromatic contrast: evidence for interactions between photopic mechanisms,” Vision Res. 17, 45–49 (1977).
[CrossRef]

1976 (1)

1975 (2)

R. S. Harwerth, H. G. Sperling, “Effects of intense visible radiation on the increment threshold spectral sensitivity of the Rhesus monkey eye,” Vision Res. 15, 1193–1204 (1975).
[CrossRef] [PubMed]

V. C. Smith, J. Pokorny, “Spectral sensitivity of the foveal cone photopigments between 400 and 500 nm,” Vision Res. 15, 161–171 (1975).
[CrossRef] [PubMed]

1974 (1)

P. E. King-Smith, J. R. Webb, “The use of photopic saturation in determining the fundamental spectral sensitivity curves,” Vision Res. 14, 421–429 (1974).
[CrossRef] [PubMed]

1971 (2)

D. E. Mitchell, W. A. H. Rushton, “Visual pigments in dichromats,” Vision Res. 11, 1033–1043 (1971).
[CrossRef] [PubMed]

J. J. Vos, P. L. Walraven, “On the derivation of the foveal receptor primaries,” Vision Res. 11, 799–818 (1971).
[CrossRef] [PubMed]

1968 (2)

W. A. H. Rushton, G. H. Henry, “Bleaching and regeneration of cone pigments in man,” Vision Res. 8, 617–631 (1968).
[CrossRef]

M. Ikeda, M. Urakubo, “Flicker HRTF as test of color vision,”J. Opt. Soc. Am. 58, 27–31 (1968).
[CrossRef] [PubMed]

1966 (1)

1964 (3)

W. S. Stiles, “Foveal threshold sensitivity on fields of different colors,” Science 145, 1016–1018 (1964).
[CrossRef] [PubMed]

G. Wald, “The receptors of human color vision,” Science 145, 1007–1016 (1964).
[CrossRef] [PubMed]

R. M. Boynton, M. Ikeda, W. S. Stiles, “Interactions among chromatic mechanisms as inferred from positive and negative increment thresholds,” Vision Res. 4, 87–117 (1964).
[CrossRef] [PubMed]

1961 (1)

1959 (1)

W. S. Stiles, “Color vision: the approach through increment threshold sensitivity,” Proc. Natl. Acad. Sci. (USA) 45, 100–114 (1959).
[CrossRef]

1958 (1)

1957 (2)

C. V. Truss, “Chromatic flicker fusion frequency as a function of chromaticity difference,”J. Opt. Soc. Am. 47, 1130–1134 (1957).
[CrossRef] [PubMed]

Y. Hsia, C. H. Graham, “Spectral luminosity curves for protanopic, deuteranopic, and normal subjects,” Proc. Natl. Acad. Sci. (USA) 43, 1011–1019 (1957).
[CrossRef]

1955 (2)

1953 (2)

G. S. Brindley, “The effects on colour vision of adaptation to very bright lights,”J. Physiol. (London) 122, 332–350 (1953).

W. S. Stiles, “Further studies of visual mechanisms by the two-colour threshold technique,” Coloquio Probl. Opt. Vision 1, 65–103 (1953).

1952 (1)

1950 (1)

E. N. Willmer, “Further observations on the properties of the central fovea in colour-blind and normal subjects,”J. Physiol. (London) 110, 422–446 (1950).

1949 (1)

S. Hecht, “Brightness, visual acuity and color blindness,” Doc. Ophthalmol. 3, 289–306 (1949).
[CrossRef]

1948 (1)

H. L. De Vries, “The luminosity curve of the eye as determined by measurements with the flicker photometer,” Physika 14, 319–348 (1948).

1947 (1)

B. H. Crawford, “Visual adaptation in relation to brief conditioning stimuli,” Proc. R. Soc. London Ser. B 134, 283–302 (1947).
[CrossRef]

1939 (1)

W. S. Stiles, “The directional sensitivity of the retina and the spectral sensitivity of the rods and cones,” Proc. R. Soc. London Ser. B 127, 64–105 (1939).
[CrossRef]

1938 (1)

K. J. W. Craik, “The effect of adaptation on differential brightness adaptation,”J. Physiol. (London) 92, 406–421 (1938).

1912 (1)

H. E. Ives, “Studies in the photometry of lights of different colours. I. Spectral luminosity curves obtained by the equality of brightness photometer and flicker photometer under similar conditions,” Phil. Mag. Ser. 6 24, 149–188 (1912).
[CrossRef]

1898 (1)

G. Burch, “On artificial temporary colour-blindness, with an examination of the colour sensations of 109 persons,” Phil. Trans. R. Soc. London Ser. B 191, 1–34 (1898).
[CrossRef]

1893 (1)

O. N. Rood, “On a photometric method that is independent of color,” Am. J. Sci. 46, 173–176 (1893).

Bowmaker, J. K.

H. J. A. Dartnall, J. K. Bowmaker, J. D. Mollon, “Human visual pigments: microspectrophotometric results from the eyes of seven persons,” Proc. R. Soc. London Ser. B 220, 115–130 (1983).
[CrossRef]

Boynton, R. M.

Brindley, G. S.

G. S. Brindley, “The colour of light of very long wavelength,”J. Physiol. (London) 130, 35–44 (1955).

G. S. Brindley, “The effects on colour vision of adaptation to very bright lights,”J. Physiol. (London) 122, 332–350 (1953).

Burch, G.

G. Burch, “On artificial temporary colour-blindness, with an examination of the colour sensations of 109 persons,” Phil. Trans. R. Soc. London Ser. B 191, 1–34 (1898).
[CrossRef]

Bush, W. R.

Carden, D.

Cole, G. R.

C. F. Stromeyer, G. R. Cole, R. E. Kronauer, “Chromatic suppression of cone inputs to the luminance flicker mechanisms,” Vision Res. 27, 1113–1137 (1987).
[CrossRef]

Craik, K. J. W.

K. J. W. Craik, “The effect of adaptation on differential brightness adaptation,”J. Physiol. (London) 92, 406–421 (1938).

Crawford, B. H.

B. H. Crawford, “Visual adaptation in relation to brief conditioning stimuli,” Proc. R. Soc. London Ser. B 134, 283–302 (1947).
[CrossRef]

Dartnall, H. J. A.

H. J. A. Dartnall, J. K. Bowmaker, J. D. Mollon, “Human visual pigments: microspectrophotometric results from the eyes of seven persons,” Proc. R. Soc. London Ser. B 220, 115–130 (1983).
[CrossRef]

Das, S. R.

De Lange, H.

De Vries, H. L.

H. L. De Vries, “The luminosity curve of the eye as determined by measurements with the flicker photometer,” Physika 14, 319–348 (1948).

DePriest, D. D.

A. Stockman, D. I. A. MacLeod, D. D. DePriest, “The temporal properties of the human short-wave photoreceptors and their associated pathways,” Vision Res. 31, 189–208 (1991).
[CrossRef]

Eisner, A.

Enoch, J. M.

J. M. Enoch, “The two-color threshold technique of Stiles and derived component color mechanisms,” in Handbook of Sensory Physiology, D. Jameson, L. H. Hurvich, eds. (Springer-Verlag, Berlin, 1972).
[CrossRef]

Gardiner, J.

Gorinson, G. A.

C. E. Sternheim, G. A. Gorinson, N. Markovitz, “Visual sensitivity during successive chromatic contrast: evidence for interactions between photopic mechanisms,” Vision Res. 17, 45–49 (1977).
[CrossRef]

Graham, C. H.

Y. Hsia, C. H. Graham, “Spectral luminosity curves for protanopic, deuteranopic, and normal subjects,” Proc. Natl. Acad. Sci. (USA) 43, 1011–1019 (1957).
[CrossRef]

Harwerth, R. S.

R. S. Harwerth, H. G. Sperling, “Effects of intense visible radiation on the increment threshold spectral sensitivity of the Rhesus monkey eye,” Vision Res. 15, 1193–1204 (1975).
[CrossRef] [PubMed]

Hecht, S.

S. Hecht, “Brightness, visual acuity and color blindness,” Doc. Ophthalmol. 3, 289–306 (1949).
[CrossRef]

Henry, G. H.

W. A. H. Rushton, G. H. Henry, “Bleaching and regeneration of cone pigments in man,” Vision Res. 8, 617–631 (1968).
[CrossRef]

Hsia, Y.

Y. Hsia, C. H. Graham, “Spectral luminosity curves for protanopic, deuteranopic, and normal subjects,” Proc. Natl. Acad. Sci. (USA) 43, 1011–1019 (1957).
[CrossRef]

Ikeda, M.

M. Ikeda, M. Urakubo, “Flicker HRTF as test of color vision,”J. Opt. Soc. Am. 58, 27–31 (1968).
[CrossRef] [PubMed]

R. M. Boynton, M. Ikeda, W. S. Stiles, “Interactions among chromatic mechanisms as inferred from positive and negative increment thresholds,” Vision Res. 4, 87–117 (1964).
[CrossRef] [PubMed]

Ives, H. E.

H. E. Ives, “Studies in the photometry of lights of different colours. I. Spectral luminosity curves obtained by the equality of brightness photometer and flicker photometer under similar conditions,” Phil. Mag. Ser. 6 24, 149–188 (1912).
[CrossRef]

Johnson, N. E.

Judd, D. B.

D. B. Judd, “Report of U.S. Secretariat Committee on Colorimetry and Artificial Daylight,” in Proceedings of the Twelfth Session of the CIE, Stockholm (Bureau Central de la CIE, Paris, 1951).

King-Smith, P. E.

P. E. King-Smith, D. Carden, “Luminance and opponent-color contributions to visual detection and adaptation and to temporal and spatial integration,”J. Opt. Soc. Am. 66, 709–717 (1976).
[CrossRef] [PubMed]

P. E. King-Smith, J. R. Webb, “The use of photopic saturation in determining the fundamental spectral sensitivity curves,” Vision Res. 14, 421–429 (1974).
[CrossRef] [PubMed]

Kronauer, R. E.

C. F. Stromeyer, G. R. Cole, R. E. Kronauer, “Chromatic suppression of cone inputs to the luminance flicker mechanisms,” Vision Res. 27, 1113–1137 (1987).
[CrossRef]

Lee, J.

J. Lee, C. F. Stromeyer, “Contribution of human short-wave cones to luminance and motion detection,”J. Physiol. (London) 413, 563–593 (1989).

MacLeod, D. I. A.

A. Stockman, D. I. A. MacLeod, N. E. Johnson, “Spectral sensitivities of the human cones,” J. Opt. Soc. Am. A 10, 2491–2521 (1993).
[CrossRef]

A. Stockman, D. I. A. MacLeod, D. D. DePriest, “The temporal properties of the human short-wave photoreceptors and their associated pathways,” Vision Res. 31, 189–208 (1991).
[CrossRef]

A. Stockman, E. D. Montag, D. I. A. MacLeod, “Large changes in phase delay on intense bleaching backgrounds,” Invest. Ophthalmol. Vis. Sci. Suppl. 32, 841 (1991).

A. Stockman, D. I. A. MacLeod, “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).

D. I. A. MacLeod, A. Stockman, J. A. Vivian, “Flicker photometric approximations to cone sensitivities under bleaching and transient adaptation conditions,” Perception 14, A17 (1985).

A. Eisner, D. I. A. MacLeod, “Flicker photometric study of chromatic adaptation: selective suppression of cone inputs by colored backgrounds,”J. Opt. Soc. Am. 71, 705–718 (1981).
[CrossRef] [PubMed]

A. Eisner, D. I. A. MacLeod, “Blue-sensitive cones do not contribute to luminance,”J. Opt. Soc. Am. 70, 121–123 (1980).
[CrossRef] [PubMed]

Markovitz, N.

C. E. Sternheim, G. A. Gorinson, N. Markovitz, “Visual sensitivity during successive chromatic contrast: evidence for interactions between photopic mechanisms,” Vision Res. 17, 45–49 (1977).
[CrossRef]

Mitchell, D. E.

D. E. Mitchell, W. A. H. Rushton, “Visual pigments in dichromats,” Vision Res. 11, 1033–1043 (1971).
[CrossRef] [PubMed]

Mollon, J. D.

A. Stockman, J. D. Mollon, “The spectral sensitivities of the middle- and long-wavelength cones: an extension of the two-colour threshold technique of W. S. Stiles,” Perception 15, 729–754 (1986).
[CrossRef]

H. J. A. Dartnall, J. K. Bowmaker, J. D. Mollon, “Human visual pigments: microspectrophotometric results from the eyes of seven persons,” Proc. R. Soc. London Ser. B 220, 115–130 (1983).
[CrossRef]

J. D. Mollon, “Color vision,” Ann. Rev. Psychol. 33, 41–85 (1982).
[CrossRef]

Montag, E. D.

A. Stockman, E. D. Montag, D. I. A. MacLeod, “Large changes in phase delay on intense bleaching backgrounds,” Invest. Ophthalmol. Vis. Sci. Suppl. 32, 841 (1991).

Pitt, F. H. G.

F. H. G. Pitt, Characteristics of Dichromatic Vision, Medical Research Council Special Report Series, No. 200 (Her Majesty’s Stationery Office, London, 1935).

Pokorny, J.

T. Yeh, V. C. Smith, J. Pokorny, “The effect of background luminance on cone sensitivity function,” Invest. Ophthalmol. Vis. Sci. 30, 2077–2086 (1989).
[PubMed]

W. H. Swanson, J. Pokorny, V. C. Smith, “Effects of temporal frequency on phase-dependent sensitivity to heterochromatic flicker,” J. Opt. Soc. Am. A 4, 2266–2273 (1988).
[CrossRef]

W. H. Swanson, J. Pokorny, V. C. Smith, “Effects of chromatic adaptation on phase-dependent sensitivity to heterochromatic flicker,” J. Opt. Soc. Am. A 5, 1976–1982 (1988).
[CrossRef] [PubMed]

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Pugh, E. N.

C. Sigel, E. N. Pugh, “Stiles’s π5color mechanism: tests of field displacements and field additivity properties,”J. Opt. Soc. Am. 70, 71–81 (1980).
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B. A. Wandell, E. N. Pugh, “Detection of long-duration incremental flashes by a chromatically coded pathway,” Vision Res. 20, 625–635 (1980).
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O. N. Rood, “On a photometric method that is independent of color,” Am. J. Sci. 46, 173–176 (1893).

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D. E. Mitchell, W. A. H. Rushton, “Visual pigments in dichromats,” Vision Res. 11, 1033–1043 (1971).
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W. A. H. Rushton, G. H. Henry, “Bleaching and regeneration of cone pigments in man,” Vision Res. 8, 617–631 (1968).
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T. Yeh, V. C. Smith, J. Pokorny, “The effect of background luminance on cone sensitivity function,” Invest. Ophthalmol. Vis. Sci. 30, 2077–2086 (1989).
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W. H. Swanson, J. Pokorny, V. C. Smith, “Effects of temporal frequency on phase-dependent sensitivity to heterochromatic flicker,” J. Opt. Soc. Am. A 4, 2266–2273 (1988).
[CrossRef]

W. H. Swanson, J. Pokorny, V. C. Smith, “Effects of chromatic adaptation on phase-dependent sensitivity to heterochromatic flicker,” J. Opt. Soc. Am. A 5, 1976–1982 (1988).
[CrossRef] [PubMed]

V. C. Smith, J. Pokorny, “Spectral sensitivity of the foveal cone photopigments between 400 and 500 nm,” Vision Res. 15, 161–171 (1975).
[CrossRef] [PubMed]

Sperling, H. G.

R. S. Harwerth, H. G. Sperling, “Effects of intense visible radiation on the increment threshold spectral sensitivity of the Rhesus monkey eye,” Vision Res. 15, 1193–1204 (1975).
[CrossRef] [PubMed]

Sternheim, C. E.

C. E. Sternheim, G. A. Gorinson, N. Markovitz, “Visual sensitivity during successive chromatic contrast: evidence for interactions between photopic mechanisms,” Vision Res. 17, 45–49 (1977).
[CrossRef]

Stiles, W. S.

R. M. Boynton, M. Ikeda, W. S. Stiles, “Interactions among chromatic mechanisms as inferred from positive and negative increment thresholds,” Vision Res. 4, 87–117 (1964).
[CrossRef] [PubMed]

W. S. Stiles, “Foveal threshold sensitivity on fields of different colors,” Science 145, 1016–1018 (1964).
[CrossRef] [PubMed]

W. S. Stiles, “Color vision: the approach through increment threshold sensitivity,” Proc. Natl. Acad. Sci. (USA) 45, 100–114 (1959).
[CrossRef]

W. S. Stiles, “Further studies of visual mechanisms by the two-colour threshold technique,” Coloquio Probl. Opt. Vision 1, 65–103 (1953).

W. S. Stiles, “The directional sensitivity of the retina and the spectral sensitivity of the rods and cones,” Proc. R. Soc. London Ser. B 127, 64–105 (1939).
[CrossRef]

W. S. Stiles, “The physical interpretation of the spectral sensitivity curve of the eye,” in Transactions of the Optical Convention of the Worshipful Company of Spectacle Makers (Spectacle Maker’s Company, London, 1948), pp. 97–107.

W. S. Stiles, Mechanisms of Colour Vision (Academic, London, 1978).

G. Wyszecki, W. S. Stiles, Color Science, 2nd ed. (Wiley, New York, 1982).

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A. Stockman, D. I. A. MacLeod, N. E. Johnson, “Spectral sensitivities of the human cones,” J. Opt. Soc. Am. A 10, 2491–2521 (1993).
[CrossRef]

A. Stockman, D. I. A. MacLeod, D. D. DePriest, “The temporal properties of the human short-wave photoreceptors and their associated pathways,” Vision Res. 31, 189–208 (1991).
[CrossRef]

A. Stockman, E. D. Montag, D. I. A. MacLeod, “Large changes in phase delay on intense bleaching backgrounds,” Invest. Ophthalmol. Vis. Sci. Suppl. 32, 841 (1991).

A. Stockman, D. I. A. MacLeod, “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).

A. Stockman, J. D. Mollon, “The spectral sensitivities of the middle- and long-wavelength cones: an extension of the two-colour threshold technique of W. S. Stiles,” Perception 15, 729–754 (1986).
[CrossRef]

D. I. A. MacLeod, A. Stockman, J. A. Vivian, “Flicker photometric approximations to cone sensitivities under bleaching and transient adaptation conditions,” Perception 14, A17 (1985).

Stromeyer, C. F.

J. Lee, C. F. Stromeyer, “Contribution of human short-wave cones to luminance and motion detection,”J. Physiol. (London) 413, 563–593 (1989).

C. F. Stromeyer, G. R. Cole, R. E. Kronauer, “Chromatic suppression of cone inputs to the luminance flicker mechanisms,” Vision Res. 27, 1113–1137 (1987).
[CrossRef]

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Truss, C. V.

Urakubo, M.

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D. I. A. MacLeod, A. Stockman, J. A. Vivian, “Flicker photometric approximations to cone sensitivities under bleaching and transient adaptation conditions,” Perception 14, A17 (1985).

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B. A. Wandell, E. N. Pugh, “Detection of long-duration incremental flashes by a chromatically coded pathway,” Vision Res. 20, 625–635 (1980).
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G. Wyszecki, W. S. Stiles, Color Science, 2nd ed. (Wiley, New York, 1982).

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T. Yeh, V. C. Smith, J. Pokorny, “The effect of background luminance on cone sensitivity function,” Invest. Ophthalmol. Vis. Sci. 30, 2077–2086 (1989).
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Am. J. Sci. (1)

O. N. Rood, “On a photometric method that is independent of color,” Am. J. Sci. 46, 173–176 (1893).

Ann. Rev. Psychol. (1)

J. D. Mollon, “Color vision,” Ann. Rev. Psychol. 33, 41–85 (1982).
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Coloquio Probl. Opt. Vision (1)

W. S. Stiles, “Further studies of visual mechanisms by the two-colour threshold technique,” Coloquio Probl. Opt. Vision 1, 65–103 (1953).

Color Res. Appl. (1)

J. J. Vos, “Colorimetric and photometric properties of a 2-deg fundamental observer,” Color Res. Appl. 3, 125–128 (1978).
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Doc. Ophthalmol. (1)

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A. Stockman, E. D. Montag, D. I. A. MacLeod, “Large changes in phase delay on intense bleaching backgrounds,” Invest. Ophthalmol. Vis. Sci. Suppl. 32, 841 (1991).

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J. Opt. Soc. Am. A (3)

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J. Lee, C. F. Stromeyer, “Contribution of human short-wave cones to luminance and motion detection,”J. Physiol. (London) 413, 563–593 (1989).

E. N. Willmer, “Further observations on the properties of the central fovea in colour-blind and normal subjects,”J. Physiol. (London) 110, 422–446 (1950).

K. J. W. Craik, “The effect of adaptation on differential brightness adaptation,”J. Physiol. (London) 92, 406–421 (1938).

G. S. Brindley, “The colour of light of very long wavelength,”J. Physiol. (London) 130, 35–44 (1955).

Perception (2)

A. Stockman, J. D. Mollon, “The spectral sensitivities of the middle- and long-wavelength cones: an extension of the two-colour threshold technique of W. S. Stiles,” Perception 15, 729–754 (1986).
[CrossRef]

D. I. A. MacLeod, A. Stockman, J. A. Vivian, “Flicker photometric approximations to cone sensitivities under bleaching and transient adaptation conditions,” Perception 14, A17 (1985).

Phil. Mag. Ser. 6 (1)

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[CrossRef]

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[CrossRef]

Physika (1)

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Proc. Natl. Acad. Sci. (USA) (2)

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[CrossRef]

W. S. Stiles, “Color vision: the approach through increment threshold sensitivity,” Proc. Natl. Acad. Sci. (USA) 45, 100–114 (1959).
[CrossRef]

Proc. R. Soc. London Ser. B (3)

W. S. Stiles, “The directional sensitivity of the retina and the spectral sensitivity of the rods and cones,” Proc. R. Soc. London Ser. B 127, 64–105 (1939).
[CrossRef]

H. J. A. Dartnall, J. K. Bowmaker, J. D. Mollon, “Human visual pigments: microspectrophotometric results from the eyes of seven persons,” Proc. R. Soc. London Ser. B 220, 115–130 (1983).
[CrossRef]

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[CrossRef]

Science (2)

W. S. Stiles, “Foveal threshold sensitivity on fields of different colors,” Science 145, 1016–1018 (1964).
[CrossRef] [PubMed]

G. Wald, “The receptors of human color vision,” Science 145, 1007–1016 (1964).
[CrossRef] [PubMed]

Vision Res. (12)

R. M. Boynton, M. Ikeda, W. S. Stiles, “Interactions among chromatic mechanisms as inferred from positive and negative increment thresholds,” Vision Res. 4, 87–117 (1964).
[CrossRef] [PubMed]

C. F. Stromeyer, G. R. Cole, R. E. Kronauer, “Chromatic suppression of cone inputs to the luminance flicker mechanisms,” Vision Res. 27, 1113–1137 (1987).
[CrossRef]

C. E. Sternheim, G. A. Gorinson, N. Markovitz, “Visual sensitivity during successive chromatic contrast: evidence for interactions between photopic mechanisms,” Vision Res. 17, 45–49 (1977).
[CrossRef]

A. Reeves, “Exchange thresholds for green tests,” Vision Res. 22, 961–966 (1982).
[CrossRef] [PubMed]

W. A. H. Rushton, G. H. Henry, “Bleaching and regeneration of cone pigments in man,” Vision Res. 8, 617–631 (1968).
[CrossRef]

A. Stockman, D. I. A. MacLeod, D. D. DePriest, “The temporal properties of the human short-wave photoreceptors and their associated pathways,” Vision Res. 31, 189–208 (1991).
[CrossRef]

R. S. Harwerth, H. G. Sperling, “Effects of intense visible radiation on the increment threshold spectral sensitivity of the Rhesus monkey eye,” Vision Res. 15, 1193–1204 (1975).
[CrossRef] [PubMed]

P. E. King-Smith, J. R. Webb, “The use of photopic saturation in determining the fundamental spectral sensitivity curves,” Vision Res. 14, 421–429 (1974).
[CrossRef] [PubMed]

D. E. Mitchell, W. A. H. Rushton, “Visual pigments in dichromats,” Vision Res. 11, 1033–1043 (1971).
[CrossRef] [PubMed]

V. C. Smith, J. Pokorny, “Spectral sensitivity of the foveal cone photopigments between 400 and 500 nm,” Vision Res. 15, 161–171 (1975).
[CrossRef] [PubMed]

J. J. Vos, P. L. Walraven, “On the derivation of the foveal receptor primaries,” Vision Res. 11, 799–818 (1971).
[CrossRef] [PubMed]

B. A. Wandell, E. N. Pugh, “Detection of long-duration incremental flashes by a chromatically coded pathway,” Vision Res. 20, 625–635 (1980).
[CrossRef]

Other (6)

F. H. G. Pitt, Characteristics of Dichromatic Vision, Medical Research Council Special Report Series, No. 200 (Her Majesty’s Stationery Office, London, 1935).

W. S. Stiles, Mechanisms of Colour Vision (Academic, London, 1978).

W. S. Stiles, “The physical interpretation of the spectral sensitivity curve of the eye,” in Transactions of the Optical Convention of the Worshipful Company of Spectacle Makers (Spectacle Maker’s Company, London, 1948), pp. 97–107.

G. Wyszecki, W. S. Stiles, Color Science, 2nd ed. (Wiley, New York, 1982).

D. B. Judd, “Report of U.S. Secretariat Committee on Colorimetry and Artificial Daylight,” in Proceedings of the Twelfth Session of the CIE, Stockholm (Bureau Central de la CIE, Paris, 1951).

J. M. Enoch, “The two-color threshold technique of Stiles and derived component color mechanisms,” in Handbook of Sensory Physiology, D. Jameson, L. H. Hurvich, eds. (Springer-Verlag, Berlin, 1972).
[CrossRef]

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Figures (12)

Fig. 1
Fig. 1

Temporal sequence of stimuli. (a) Background condition: the two 17-Hz flicker photometric test lights and the steady background were exposed continuously. (b) Bleach condition: the bleaching background was alternated with the two flicker photometric test lights. The bleaching background was exposed for 15 s and the two test lights for 5 s. (Not to scale.)

Fig. 2
Fig. 2

17-Hz flicker photometric spectral sensitivities for subjects (a) JAV and (b) AS measured following a 5.43-log10-Td, 485-nm bleach (open diamonds); on a steady 3.00-log10-Td, 485-nm background (filled diamonds); following a 5.75-log10-Td, 678-nm bleach (open circles); and on a steady 4.00-log10-Td, 678-nm background (filled circles). Standard stimulus: 561 nm at approximately 8 log quanta sec−1 deg−2. Curves are Smith–Pokorny L-cone (upper) and M-cone (lower) fundamentals. (c) Filled circles, 17-Hz flicker photometric sensitivity differences between 545- and 668-nm lights for JAV measured as function of the 678-nm background luminance. (d) Open circles, 545-668-nm sensitivity differences for AS as a function of the luminance of the 678-nm bleach. The horizontal lines in (c) and (d) represent the Smith–Pokorny M-cone-sensitivity difference. ±1 Standard deviation across sessions is shown when it is larger than the size of the plotted symbol.

Fig. 3
Fig. 3

Temporal sequence of stimuli. (a) Exchange condition: the two backgrounds, each on for 1 s, were exchanged once every 2 s. The two 17-Hz flicker photometric test lights were exposed only during the 500 ms immediately following the transition from the preceding background to the concurrent one. (b) Pulsed condition: like the exchange condition, except that the preceding background was absent. (Not to scale.)

Fig. 4
Fig. 4

Differences in flicker photometric sensitivity for 17-Hz, 545-nm, and 668-nm lights. (a) 545–668-nm sensitivity differences for JAV as a function of 678-nm background luminance, following an exchange of background from 485 to 678 nm (circles) or on a steady 678-nm field (triangles). The luminance of the preceding 485-nm background was 3.32 log10 Td. The horizontal line is the Smith–Pokorny M-cone sensitivity. The triangular symbols plotted along the abscissa denote the concurrent background luminances at which the exchange was invisible to the L cones (3.08 log10 Td) and to the M cones (4.46 log10 Td). (b) 545–668-nm sensitivity differences for JAV measured at two concurrent 678-nm luminances (open circles, 2.88 log10 Td; filled circles, 3.50 log10 Td) as a function of the preceding 485-nm background luminance. The squares denote the flashed condition, when the 485-nm background is extinguished. The triangular symbols on the abscissa denote the preceding background luminances at which the exchange was invisible to the M cones (1.74 and 2.36 log10 Td for the low and the high levels, respectively) and to the L cones (3.12 and 3.74 log10 Td for the low and the high levels, respectively).

Fig. 5
Fig. 5

545–668-nm, 17-Hz flicker photometric sensitivity differences for (a) JAV and (b) AS measured with a concurrent 678-nm background of 2.88 log10 Td, as a function of the preceding background luminance for five preceding background wavelengths: 485 nm (filled circles), 530 nm (open circles), 574 nm (filled triangles), 617 nm (open squares) and a deep red, homochromatic with the concurrent field (filled squares). The vertical line corresponds to the luminance at which the deep-red concurrent and preceding backgrounds are equal in luminance. The horizontal line is the Smith–Pokorny M-cone sensitivity. The filled diamond is the average of the sensitivity differences obtained when the preceding backgrounds were absent. Error bars are ±1 standard deviation. The triangular symbols on the abscissa denote the preceding background luminances (485 nm, 574 nm, and deep red only) at which the exchange is M- or L-cone equated. (c) 545–668-nm, 17-Hz flicker photometric sensitivity differences for JAV (filled symbols) and AS (open symbols) for background exchanges of equal luminance (2.88 log10 Td) as a function of ΔL and ΔM, where ΔL and ΔM are the change in L- and M-cone excitations (expressed as a fraction of the total luminance) caused by the background exchange (see text for details). Preceding background wavelengths: 485 nm (circles), 530 nm (diamonds), 574 nm (inverted triangles), 617 nm (upright triangles), and deep red (squares).

Fig. 6
Fig. 6

Detection thresholds for 17-Hz, 545-nm (triangles) and 668-nm (circles) flicker measured as a function of background luminance on a steady 678-nm field (open symbols) or following an exchange of background from 485 to 678 nm (filled symbols). The luminance of the 485-nm preceding background was 3.26 log10 Td. The triangular symbols plotted along the abscissa denote the concurrent background luminances at which the exchange was invisible to the L cones (3.02 log10 Td) and to the M cones (4.40 log10 Td). For each subject, we shifted the 668-nm f.t.v.i. curve 2.46 log10 units downward relative to the 545-nm curve to equate the test lights for their effects on the M cones. The 546-nm f.t.v.i. curves for AS and the 668-nm f.t.v.i. curve for JAV are plotted correctly with respect to the scale of the ordinate. To obtain the correct threshold levels for the 668-nm f.t.v.i. curve for AS add 2.46 log10 units, and for the 545-nm f.t.v.i. curve for JAV subtract 2.46 log10 units.

Fig. 7
Fig. 7

(a) Upper curves are the difference in log10 sensitivity for detection of 545- and 668-nm, 17-Hz flicker for JAV either on a steady 678-nm background (filled triangles) or following an exchange of background from 485 to 678 nm (filled circles) plotted as a function of background luminance. The sensitivity differences are derived from the data of Fig. 6. The upper horizontal dashed line is the Smith–Pokorny M-cone sensitivity. The lower curves are the phase advances (from 180°) of the 545-nm flickering light required for a flicker photometric null of the 668-nm light on the steady 678-nm background (open triangles) or following the exchange of background from 485 to 678 nm (open circles), also plotted as a function of background luminance. The lower dashed line is the 0° phase advance that is expected if the two lights are detected by the same cone type. In the exchange procedure the preceding 485-nm background was 3.26 log10 Td. The triangular symbols plotted along the abscissa denote the concurrent background luminances at which the exchange was invisible to the L cones (3.02 log10 Td) and to the M cones (4.40 log10 Td). The flicker-detection sensitivity differences shown in (a) for JAV can be compared with his flicker photometric sensitivity differences shown in Fig. 4(a). (b) Like (a) but for subject AS. (c) L-cone isolation. The upper curves are the difference in log10 sensitivity for detection of 638- and 470-nm, 17-Hz flicker for JAV either on a steady 485-nm background (filled triangles) or following an exchange of background from 678 to 485 nm (filled circles) plotted as a function of background luminance. The upper horizontal dashed line is our estimate of the L-cone spectral sensitivity. The lower curves are the phase advances (from 180°) of the 638-nm flickering light required for flicker photometric null of the 470-nm light on the steady 485-nm background (open triangles) or following the exchange of background from 678 to 485 nm (open circles). The preceding 678-nm background was 3.56 log10 Td. The triangular symbols plotted along the abscissa denote the concurrent background luminances at which the exchange was invisible to the M cones (2.42 log10 Td) and to the L cones (3.80 log10 Td). (d) Like (c) but for subject AS. [See Figs. 1(a) and 1(b) of Ref. 9 for exchange sensitivity measurements in other subjects.]

Fig. 8
Fig. 8

Open circles are the phase advances (from 180°) of the 545-nm flickering light required for flicker photometric nulling of the 668-nm light following an exchange from a 3.26-log10-Td, 485-nm background to a 4.16-log10-Td, 678-nm background. Filled circles are the phase advances of the 638-nm flickering light required for nulling of the 470-nm light following an exchange from a 3.29-log10-Td, 678-nm background to a 3.91-log10-Td, 485-nm background. Subject: JAV

Fig. 9
Fig. 9

Spectral sensitivities for 17-Hz flicker (circles) and flashes (triangles). (a) Subject JAV M-cone flicker sensitivities (lower circles) measured following an exchange from a 3.26-log10-Td, 485-nm background to a 4.14-log10-Td, 678-nm background. L-cone flicker sensitivities (upper circles) measured following an exchange from a 3.56-log10-Td, 678-nm background to a 3.88-log10-Td, 485-nm background. Test sensitivities at 545 nm are −8.20 and −8.85 log10, quanta sec−1 deg−2 for the M- and the L-cone data, respectively. Flash sensitivities measured following an exchange from a 3.30-log10-Td, 485-nm background to a 3.79-log10-Td, 678-nm background. Test sensitivities at 545 nm are −7.52 and −8.07 log10 quanta sec−1 deg−2 for the 10- and the 200-ms functions, respectively. (b) Subject AS. M-cone flicker sensitivities measured following an exchange from a 2.94-log10-Td, 485-nm background to a 3.98-log10-Td, 678-nm background, and L-cone flicker sensitivities measured following an exchange from a 3.29-log10-Td, 678-nm background to a 3.91-log10-Td, 485-nm background. Test sensitivities at 545 nm are −7.96 and −8.69 log10 quanta sec−1 deg−2 for the M- and the L-cone estimates, respectively. The diamonds and squares are test sensitivity measurements for AS made in a different laboratory. They were obtained with a foveally fixated, 17-ms-duration, 3-min-diameter test flash presented in the center of a 7-min-diameter background (see text for details). Error bars in both panels are ±1 standard deviation; curves are Smith–Pokorny M- and L-cone sensitivities.

Fig. 10
Fig. 10

(a) Bipartite field color-matching data for AS, JAV, and NEJ measured either following an exchange of background from 678 to 485 nm (circles) or on a steady 678-nm field (triangles), with Smith–Pokorny M-cone sensitivity (continuous curves). A standard of 561 nm was used. If there was a perfect match between the standard and the test light, an open symbol is plotted. If there was not, a filled symbol is plotted. The concurrent or steady 678-nm background luminances were 4.23, 4.16, and 4.16 log10 Td, and the preceding 485-nm background luminances were 2.94, 3.26, and 3.26 log10 Td for AS, JAV, and NEJ, respectively. The absolute sensitivities at 545 nm for the exchange condition are −8.36, −8.21, and −8.70 log10 quanta sec−1 deg−2 for AS, JAV, and NEJ, respectively, and for the steady condition are −8.06 and −8.42 log10 quanta sec−1 deg−2 for AS and NEJ, respectively. (b) Color-matching data for the three subjects measured either following an exchange of background from 485 to 678 nm (circles) or on a steady 485-nm field (triangles), with Smith–Pokorny L-cone sensitivity (continuous curves). The concurrent or steady 485-nm background luminances were 3.91, 3.47, and 4.21 log10 Td, and the preceding 678-nm background luminances were 3.29, 3.56, and 3.56 log10 Td for AS, JAV, and NEJ, respectively. The test sensitivities at 545 nm for the exchange condition are −9.52, −8.59, and −9.37 log10 quanta sec−1 deg−2 for AS, JAV, and NEJ, respectively, and for the steady condition are −9.34 and −9.17 log10 quanta sec−1 deg−2 for AS and NEJ, respectively. Other details as for (a). Subject JAV made matches only following background exchange.

Fig. 11
Fig. 11

How an opponent attenuation might independently suppress M- and L-cone 17-Hz flicker signals.

Fig. 12
Fig. 12

Vector diagram illustrating how a 16° phase difference between 545- and 668-nm flickering lights and a near-M-cone spectral sensitivity implies a much larger phase difference between the underlying M- and L-cone signals. Each cone’s response to a stimulus component is represented by a vector, the length of which represents its amplitude and the direction its phase. OP, pure M-cone response to 545-nm stimulus; PQ, M-cone response to 668-nm stimulus (retarded from opposite phase by 16°), QO, L-cone response to 668-nm stimulus (advanced by 60° relative to the M-cone response); PO, resultant vector for 668 nm stimulus exactly cancels M-cone response to 545-nm stimulus. We assume that under near-M-cone isolation the M-cone response to the 668 nm light (QP) is 2.5 times that of the L-cone response (QO).

Tables (2)

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Table 1 Percentage L- and M-Cone Contributions That Best Describe the L- or M-Cone Spectral-Sensitivity Estimates for Subjects AS and JAV Obtained Either on Steady Backgrounds or Following Intense Bleaches

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Table 2 Percentage L- and M-Cone Contributions That Best Describe the L- or M-Cone Spectral-Sensitivity Estimates for Subjects AS and JAV Obtained Following an Exchange of Background Color and Intensity

Equations (2)

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log S M = log ( M λ + w L w M L λ ) + k M , log S L = log ( w M w L M λ + L λ ) + k L .
log S λ = log ( 100 w M w L + w M M λ + 100 w L w L + w M L λ ) + c ,

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