Abstract

The relative numerosity of the long-wavelength-sensitive (L) and middle-wavelength-sensitive (M) cones and the red–green color appearance, as assessed by means of unique yellow, are stable from fovea to midperiphery (±28 deg nasotemporal). As foveal tests decrease in size, unique yellow progressively shifts toward longer wavelengths, favoring a model of red–green opponency carried by cells whose centers receive input from either L or M cones and whose surrounds receive mixed contributions from both. Individual differences in unique yellow over a 20-nm range and the relative numerosity of L and M cones can be linked by means of this model, suggesting that the relative number of L and M cones is a factor that regulates individual variations in red–green color appearance.

© 2000 Optical Society of America

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1999 (2)

A. Roorda, D. R. Williams, “The arrangement of the three cone classes in the living human eye,” Nature 397, 520–522 (1999).
[CrossRef] [PubMed]

J. Mollon, “Color vision: opsins and options,” Proc. Natl. Acad. Sci. USA 96, 4743–4745 (1999).
[CrossRef] [PubMed]

1998 (3)

S. A. Hagstrom, M. Neitz, J. Neitz, “Variations in cone populations for red–green color vision examined by analysis of mRNA,” NeuroReport 9, 1963–1967 (1998).
[CrossRef] [PubMed]

P. D. Gowdy, C. M. Cicerone, “The spatial arrangement of L and M cones in the central fovea of the living human eye,” Vision Res. 38, 2575–2589 (1998).
[CrossRef]

E. Miyahara, J. Pokorny, V. C. Smith, R. Baron, E. Baron, “Color vision in two observers with highly biased LWS/MWS cone ratios,” Vision Res. 38, 601–612 (1998).
[CrossRef] [PubMed]

1996 (3)

M. J. Sankeralli, K. T. Mullen, “Estimation of the L-, M-, and S-cone weights of the postreceptoral detection mechanisms,” J. Opt. Soc. Am. A 13, 906–915 (1996).
[CrossRef]

O. S. Packer, D. R. Williams, D. G. Bensinger, “Photopigment transmittance imaging of the primate photoreceptor mosaic,” J. Neurosci. 16, 2251–2260 (1996).
[PubMed]

D. M. Dacey, “Circuitry for color coding in the primate retina,” Proc. Natl. Acad. Sci. USA 93, 582–588 (1996).
[CrossRef] [PubMed]

1995 (3)

1994 (2)

D. J. Calkins, S. J. Schein, Y. Tsukamoto, P. Sterling, “M and L cones in macaque fovea connect to midget ganglion cells by different numbers of excitatory synapses,” Nature 371, 70–72 (1994).
[CrossRef] [PubMed]

A. Chaparro, C. F. Stromeyer, R. E. Kronauer, R. T. Eskew, “Separable red–green and luminance detectors for small flashes,” Vision Res. 34, 751–762 (1994).
[CrossRef] [PubMed]

1993 (3)

R. L. De Valois, K. K. De Valois, “A multi-stage color model,” Vision Res. 33, 1053–1065 (1993).
[CrossRef] [PubMed]

J. Neitz, M. Neitz, G. H. Jacobs, “More than three different cone pigments among people with normal color vision,” Vision Res. 33, 117–122 (1993).
[CrossRef] [PubMed]

R. Navarro, P. Artal, D. R. Williams, “Modulation transfer of the human eye as a function of retinal eccentricity,” J. Opt. Soc. Am. A 10, 201–212 (1993).
[CrossRef] [PubMed]

1992 (6)

R. C. Reid, R. M. Shapley, “Spatial structure of cone inputs to receptive fields in primate lateral geniculate nucleus,” Nature 356, 716–718 (1992).
[CrossRef] [PubMed]

S. L. Merbs, J. Nathans, “Absorption spectra of human cone pigments,” Nature 356, 433–435 (1992).
[CrossRef] [PubMed]

D. M. Dacey, M. R. Petersen, “Dendritic field size and morphology of midget and parasol ganglion cells of the human retina,” Proc. Natl. Acad. Sci. USA 89, 9666–9670 (1992).
[CrossRef] [PubMed]

H. Hibino, “Red–green and yellow–blue opponent-color responses as a function of retinal eccentricity,” Vision Res. 32, 1955–1964 (1992).
[CrossRef] [PubMed]

J. D. Mollon, J. K. Bowmaker, “The spatial arrangement of cones in the primate fovea,” Nature 360, 677–679 (1992).
[CrossRef] [PubMed]

J. L. Nerger, C. M. Cicerone, “The ratio of L to M cones in the human parafoveal retina,” Vision Res. 32, 879–888 (1992).
[CrossRef] [PubMed]

1991 (2)

C. A. Curcio, K. A. Allen, K. R. Sloan, C. L. Lerea, J. B. Hurley, I. B. Klock, A. H. Milam, “Distribution and morphology of human cone photoreceptors stained with anti-blue,” J. Comp. Neurol. 312, 610–624 (1991).
[CrossRef] [PubMed]

I. Abramov, J. Gordon, H. Chan, “Color appearance in the peripheral retina: effect of stimulus size,” J. Opt. Soc. Am. A 8, 404–414 (1991).
[CrossRef] [PubMed]

1990 (2)

C. A. Curcio, K. R. Sloan, O. Packer, R. E. Kalina, A. E. Hendrickson, “Human photoreceptor topography,” J. Comp. Neurol. 292, 497–523 (1990).
[CrossRef] [PubMed]

J. Neitz, G. H. Jacobs, “Polymorphism in normal human color vision and its mechanisms,” Vision Res. 30, 621–636 (1990).
[CrossRef]

1989 (3)

H. Wässle, B. B. Boycott, J. Rohrenbeck, “Horizontal cells in the monkey retina: cone connections and dendritic network,” Eur. J. Neurosci. 1, 421–435 (1989).
[CrossRef] [PubMed]

R. L. P. Vimal, J. Pokorny, V. C. Smith, S. K. Shevell, “Foveal cone thresholds,” Vision Res. 29, 61–78 (1989).
[CrossRef] [PubMed]

C. M. Cicerone, J. L. Nerger, “The relative number of long-wavelength-sensitive to middle-wavelength-sensitive cones in the human fovea centralis,” Vision Res. 29, 115–128 (1989).
[CrossRef]

1987 (3)

P. K. Ahnelt, H. Kolb, R. Pflug, “Identification of a subtype of cone photoreceptor, likely to be blue-sensitive, in the human retina,” J. Comp. Neurol. 255, 18–34 (1987).
[CrossRef] [PubMed]

C. M. Cicerone, “Constraints placed on color vision models by the relative number of different cone classes in human fovea centralis,” Farbe 34, 59–66 (1987).

J. L. Schnapf, T. Kraft, D. A. Baylor, “Spectral sensitivities of human cone photoreceptors,” Nature 325, 439–441 (1987).
[CrossRef] [PubMed]

1985 (2)

D. G. Pelli, “Uncertainty explains many aspects of visual contrast detection and discrimination,” J. Opt. Soc. Am. A 2, 1508–1532 (1985).
[CrossRef] [PubMed]

F. M. de Monasterio, E. P. McCrane, J. K. Newlander, S. J. Schein, “Density profile of blue-sensitive cones along the horizontal meridian of macaque retina,” Invest. Ophthalmol. Visual Sci. 26, 289–302 (1985).

1984 (2)

1983 (3)

C. Noorlander, J. J. Koenderink, R. J. Den Ouden, B. W. Edens, “Sensitivity to spatiotemporal colour contrast in the peripheral visual field,” Vision Res. 23, 1–11 (1983).
[CrossRef] [PubMed]

W. Paulus, A. Kroger-Paulus, “A new concept of retinal colour coding,” Vision Res. 23, 529–540 (1983).
[CrossRef] [PubMed]

J. E. Thornton, E. N. Pugh, “Red/green opponency at detection threshold,” Science 219, 191–193 (1983).
[CrossRef] [PubMed]

1982 (4)

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

U. Stabell, B. Stabell, “Color-vision in the peripheral retina under photopic conditions,” Vision Res. 22, 839–844 (1982).
[CrossRef]

H. Uchikawa, P. K. Kaiser, K. Uchikawa, “Color-discrimination perimetry,” Color Res. Appl. 7, 264–272 (1982).
[CrossRef]

J. A. Castano, H. G. Sperling, “Sensitivity of the blue-sensitive cones across the central retina,” Vision Res. 22, 661–673 (1982).
[CrossRef] [PubMed]

1981 (3)

D. R. Williams, D. I. A. MacLeod, M. M. Hayhoe, “Punctuate sensitivity of the blue-sensitive mechanism,” Vision Res. 21, 1357–1375 (1981).
[CrossRef]

J. Nachmias, “On the psychometric function for contrast detection,” Vision Res. 21, 215–223 (1981).
[CrossRef] [PubMed]

J. Larimer, “Red/green opponent colors equilibria measured on chromatic adapting fields: evidence for gain changes and restoring forces,” Vision Res. 21, 501–512 (1981).
[CrossRef] [PubMed]

1980 (2)

J. K. Bowmaker, H. J. A. Dartnall, “Visual pigments of rods and cones in a human retina,” J. Physiol. (London) 298, 501–511 (1980).

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

1979 (1)

1977 (5)

A. L. Nagy, J. L. Zacks, “The effects of psychophysical procedure and stimulus duration in the measurement of Bezold–Brücke hue shifts,” Vision Res. 17, 193–200 (1977).
[CrossRef] [PubMed]

M. Alpern, E. N. Pugh, “Variation in the action spectrum of erythrolabe among deuteranopes,” J. Physiol. (London) 266, 613–646 (1977).

M. Alpern, T. Wake, “Cone pigments in human deutan colour vision defects,” J. Physiol. (London) 266, 595–612 (1977).

J. Pokorny, V. C. Smith, “Evaluation of single pigment shift model of anomalous trichromacy,” J. Opt. Soc. Am. 67, 1196–1209 (1977).
[CrossRef] [PubMed]

R. E. Marc, H. G. Sperling, “Chromatic organization of primate cones,” Science 196, 454–456 (1977).
[CrossRef] [PubMed]

1975 (2)

D. H. Krantz, “Color measurement and color theory. Opponent-colors theory,” J. Math. Psychol. 12, 304–327 (1975).
[CrossRef]

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)

J. Larimer, D. H. Krantz, C. M. Cicerone, “Opponent-process additivity. I. Red/green equilibria,” Vision Res. 14, 1127–1140 (1974).
[CrossRef] [PubMed]

1973 (1)

B. R. Wooten, G. Wald, “Color-vision mechanisms in the peripheral retinas of normal and dichromatic observers,” J. Gen. Physiol. 61, 125–145 (1973).
[CrossRef] [PubMed]

1972 (1)

B. Sakitt, “Counting every quantum,” J. Physiol. (London) 223, 513–529 (1972).

1971 (2)

M. Alpern, “Rhodopsin kinetics in the human eye,” J. Physiol. (London) 217, 447–471 (1971).

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

1966 (2)

F. W. Campbell, R. W. Gubisch, “Optical quality of the human eye,” J. Physiol. (London) 186, 558–578 (1966).

T. N. Wiesel, D. H. Hubel, “Spatial and chromatic interactions in the lateral geniculate body of the rhesus monkey,” J. Neurophysiol. 29, 1115–1156 (1966).
[PubMed]

1964 (2)

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1962 (1)

1959 (1)

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1955 (1)

1948 (1)

H. L. De Vries, “The heredity of the relative number of red and green receptors in the human eye,” Genetica (The Hague) 24, 199–212 (1948).

1946 (1)

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1937 (1)

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Figures (10)

Fig. 1
Fig. 1

A, Observer’s view of foveal and eccentric presentations of the stimulus. For foveal measurements (top), the observer was asked to fixate the center of the square whose corners are marked by four dots. For measurements at 2 deg eccentricity (middle), the fixation pattern consisted of three dots in the configuration shown here or in its mirror image. The stimulus was presented midway between the two vertically placed dots. The observer was asked to fixate the third dot. For eccentricities of ⩾7 deg (bottom), the observer was asked to fixate a single fixation light delivered by a source external to the Maxwellian-view apparatus, as shown in this configuration or in its mirror image. B, Sequence of presentation of bleach, background, and test lights presented in the periphery. The bleaching light was first presented for 10 s, after which the L- or M-cone-favoring background was immediately presented. After 3 min of light adaptation, test trials were presented. Foveal measurements were identical, except that the rod bleach was not applied.

Fig. 2
Fig. 2

Example of incremental threshold measurements used to determine the intensity of the background required for raising threshold for the test by 0.5 log unit. The data are derived from the psychometric functions defined by ratings ⩾2 (see text). Open circles, 60% seen according to this criterion. Plus signs, Stiles’s theoretical threshold-versus-intensity function. The intensity of the background was chosen as that which elevated threshold 0.5 log unit above the absolute level, as indicated by the arrowhead placed on the abscissa.

Fig. 3
Fig. 3

Probability of detection functions for observer KL for nasal eccentricities 28, 7, and 2 deg, fovea, and temporal eccentricities 2, 7, 17, and 28 deg. Open symbols, L cone conditions; filled symbols, M cone conditions. The best-fitting theoretical functions as shown by the dotted curves (see text) were used to estimate the number of L cones (NL) and the number of M cones (NM) that contribute to detection at each location. The abscissa displays relative intensity in terms of a linear scale of relative numbers of quanta absorbed per cone. The nominal value of 1 displayed on the abscissa for the foveal results corresponds to an estimated value of 4.63×103 quanta per flash delivered at the cornea based on radiometric measurements.

Fig. 4
Fig. 4

Probability of detection functions for observer SL shown for nasal eccentricities 28 and 7 deg, fovea, and temporal eccentricities 7, 17, and 28 deg. Open symbols, L cone conditions; filled symbols, M cone conditions. The best-fitting theoretical functions as shown by the dotted curves (see text) were used to estimate the number of L cones (NL) and the number of M cones (NM) that contribute to detection at each location. The abscissa displays relative intensity in terms of a linear scale of relative numbers of quanta absorbed per cone. The nominal value of 1 displayed on the abscissa for the foveal results corresponds to an estimated value of 3.87×103 quanta per flash delivered at the cornea based on radiometric measurements.

Fig. 5
Fig. 5

Relative number of L and M cones plotted as a function of eccentricity. Squares, observer KL’s results; triangles, observer SL’s results. The slopes of the horizontal lines, drawn through each observer’s average value, are not significantly different from zero.

Fig. 6
Fig. 6

Estimated density of the total number of L and M cones derived for observers KL and SL, plotted as a function of eccentricity. Also shown are the histological estimates from studies by Østerberg1 and by Curcio et al.2 for total L, M, and S cone density.

Fig. 7
Fig. 7

Unique yellow wavelengths (±1 standard error of the mean) plotted as a function of eccentricity. Squares, observer KL’s results; triangles, observer SL’s results. The slopes of the horizontal lines drawn through each observer’s average value are not significantly different from zero.

Fig. 8
Fig. 8

Proportion of test lights of a given wavelength judged to appear green as a function of wavelength plotted for observers VC and MS. The smallest circles represent the results for a 3-arc-min test; the medium-sized circles, for a 10-arc-min test; and the largest circles, for a 30-arc-min test.

Fig. 9
Fig. 9

Unique yellow (the wavelength judged green 50% of the time and red 50% of the time) plotted as a function of test size ranging from 3 to 60 arc min for observers VC and MS.

Fig. 10
Fig. 10

Unique yellow plotted as a function of the relative number of L and M cones for foveal measurements in 11 observers [4 observers from this study (filled circles), and 7 from a previous study16]. The predictions are from the model described in the text [Eq. (2)]. A relative weight of center versus surround of 3.8 for the red–green opponent receptive fields best describes these results.

Equations (4)

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P(x)=1-Q(x)N.
Q(x)=k=1q-1(e-xxk/k!),
cM(λ)-[jL(λ)+kM(λ)],
(c-1)M(λY)-jL(λY)=0.

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