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The intrinsic melanopsin photoresponse may initiate visual signals that differ in spatiotemporal characteristics from the cone-opsin- and rhodopsin-mediated signals. Applying the CIE standard observer functions in silent-substitution methods can require individual differences in photoreceptor spectral sensitivities and pre-receptoral filtering to be corrected; failure to do so can lead to the intrusion of more sensitive cone processes with putative melanopsin-directed stimuli. Here we evaluate heterochromatic flicker photometry (HFP) and photoreceptor-directed temporal white noise as techniques to limit the effect of these individual differences. Individualized luminous efficiency functions ( V ( λ )) were compared to the CIE standard observer functions. We show that adapting chromaticities used in silent-substitution methods can deviate by up to 54% in luminance when estimated with the individual and standard observer functions. These deviations lead to inadvertent cone intrusions in the visual functions measured with melanopsin-directed stimuli. To eliminate the intrusions, individual HFP corrections are sufficient at low frequencies (∼1 H z ) but temporal white noise is also required at higher frequencies to desensitize penumbral cones. We therefore recommend the selective application of individualized observer calibration and/or temporal white noise in silent-substitution paradigms when studying melanopsin-directed photoresponses.

We established a protocol using a well-established LED stimulator to measure temporal contrast sensitivities driven by sine-wave modulation of L- and M-cones in the perifovea using triple silent substitution. The stimulus was presented in an annular field (2° inner diameter, 13° outer diameter). We validated this technique by studying the contrast sensitivity of three color normal observers at 10 different temporal frequencies (between 1 and 28 Hz) over a large range of retinal illuminances (between 0.07 and 587 phot Td), spanning the complete mesopic range. In one subject, sensitivities to counterphase modulation of L- and M-cones and in-phase modulation of L, M, and S-cones were additionally measured, which putatively reflected the parvo- and magnocellular retinogeniculate pathways, respectively. Furthermore, we performed measurements of temporal contrast sensitivities as a function of frequency at 294 phot Td in two protanopes, in two deuteranopes, and in one subject with S-cone monochromacy. Quality of isolation was satisfactory and we were able to reproduce known physiological patterns of temporal vision, such as the typical temporal contrast sensitivity functions of the L- and M-cone, the parvo- and magnocellular retinogeniculate pathways, as well as the light adaptation curves. These results will help determine optimal stimulus conditions in future studies. Results from the dichromats and the S-cone monochromat also support the quality of isolation of our protocol and underpin its potential clinical value.

We explored the color constancy mechanisms of color-deficient observers under red, green, blue, and yellow illuminations. The red and green illuminations were defined individually by the longer axis of the color discrimination ellipsoid measured by the Cambridge Colour Test. Four dichromats (3 protanopes and 1 deuteranope), two anomalous trichromats (2 deuteranomalous observers), and five color-normal observers were asked to complete the color constancy task by making a simultaneous paper match under asymmetrical illuminations in haploscopic view on a monitor. The von Kries adaptation model was applied to estimate the cone responses. The model fits showed that for all color-deficient observers under all illuminations, the adjustment of the S-cone response or blue-yellow chromatically opponent responses modeled with the simple assumption of cone deletion in a certain type (S-M, S-L or S-(L+M)) was consistent with the principle of the von Kries model. The degree of adaptation was similar to that of color-normal observers. The results indicate that the color constancy of color-deficient observers is mediated by the simplified blue-yellow color system with a von Kries-type adaptation effect, even in the case of brightness match, as well as by a possible cone-level adaptation to the S-cone when the illumination produces a strong S-cone stimulation, such as blue illumination.

The Rayleigh match is defined by the range of mixtures of red and green lights that appear the same as an intensity-adjustable monochromatic yellow light. The perceptual match indicates that the red-green mixture and the yellow light have evoked the same respective cone absorptions in the L- and M-cone pathways. Going beyond the existing models, the Poisson noise in cone absorptions is proposed to make the matching proportion of red-green mixtures span a finite range because any mixture in that range evokes cone absorptions that do not differ from those by a yellow light by more than the variations in the absorption noise. We derive a mathematical formula linking the match midpoint or match range with the sensitivities and numerosities of the two cones. The noise-free, exact, matching point, close to the midpoint of the matching range, depends only on the L- and M-cone sensitivities to each of the red, green, and yellow lights [these sensitivities, in turn, depend on the preferred wavelengths (λmax) and optical densities of the cone pigments and the properties of prereceptoral light filtering]. Meanwhile, the matching range depends on both these cone sensitivities and the relative numerosity of the L and M cones. The model predicts that, in normal trichromats, all other things being equal, the match range is smallest when the ratio r between L and M cone densities is r=R(-1/2) with R as the ratio between the sensitivities of the L and M cones to the yellow light, i.e., when L and M cones are similarly abundant in typical cases, and, as r departs from R(-1/2), the match range increases. For example, when one cone type is 10 times more numerous, the match range increases two- to threefold, depending on the sensitivities of the cones. Testing these model predictions requires either a large data set to identify the effect of one factor (e.g., cone numerosity) while averaging out the effects of the other factors (e.g., cone sensitivities) or for all factors to be known. A corollary of this prediction is that, because they are more likely than usual to have L:M cone ratios skewed, the matching ranges of normal female trichromats who are carriers of dichromacy (but not anomalous trichromacy) are likely to have a larger matching range than usual, particularly for the deutan carriers. In addition, the model predicts that, in strong tetrachromats (whose four dimensions of color are preserved post-receptorally), either the Rayleigh matching is impossible or the matching range is typically smaller than usual.

We investigated the brightness (i.e., perceived luminance) of isolated L- and M-cone pulses to seek a perceptual correlate of our previous reports that M-on electroretinograms resemble L-off responses, implying the operation of post-receptoral opponent processing. Using triple silent substitutions, cone increments were generated in a 4-primary ganzfeld, masked by random positive or negative luminance bias. The results show that M-cone increments decrease in brightness, while L-cone increments increase. These differences became smaller as field size reduced; this was not eccentricity or area dependent. We speculate about early retinal input into brightness perception.

An important goal in characterizing human color vision is to order color percepts in a way that captures their similarities and differences. This has resulted in the continuing evolution of "uniform color spaces," in which the distances within the space represent the perceptual differences between the stimuli. While these metrics are now very successful in predicting how color percepts are scaled, they do so in largely empirical, ad hoc ways, with limited reference to actual mechanisms of color vision. In this article our aim is to instead begin with general and plausible assumptions about color coding, and then develop a model of color appearance that explicitly incorporates them. We show that many of the features of empirically defined color order systems (those of Munsell, Pantone, NCS, and others) as well as many of the basic phenomena of color perception, emerge naturally from fairly simple principles of color information encoding in the visual system and how it can be optimized for the spectral characteristics of the environment.

There is theoretical and empirical support for long-term adaptation of human vision to chromatic regularities in the environment. The current study investigates whether relationships of luminance and chromaticity in the natural environment could drive chromatic adaptation independently and differently for bright and dark colors. This is motivated by psychophysical evidence of systematic difference shifts in red-green chromatic sensitivities between contextually bright- versus dark-colored stimuli. For some broad classes of scene content, consistent shifts in chromaticity are found between high and low light levels within images. Especially in those images in which sky and terrain are juxtaposed, this shift has direction and magnitude consistent with the observed psychophysical shifts in the red-green balance between bright and dark colors. Taken together, these findings suggest that relative weighting of M- and L-cone signals could be adapted, in a luminance-dependent fashion, to regularities in the natural environment.

Equilibrium (unique) red, green, blue, and yellow stimuli look bright in a black surround, but they look dark in a bright white surround, and yellow changes to brown. We investigated differences in equilibrium-hue chromaticity between bright and dark hues to reveal changes in weighting of cone and rod signals. The largest, most consistent shifts were found between yellow and brown, with equilibrium-brown chromaticity shifted toward red compared to equilibrium yellow at both photopic and mesopic levels. Also, at mesopic levels, rod influence reversed for most observers from a green bias for yellow to a red bias for brown. Bright/dark differences for blue, green, and red were much smaller and/or less consistent. Thus, shifts of cone and rod hue biases between bright and dark hues are most prominent in L-M-cone pathways, especially those activated by yellow and brown stimuli.

This study investigated the mechanisms of flicker detection suppression by measuring mesopic rod and cone critical flicker frequencies (CFFs) at different center and surround illuminance levels. Stimuli were generated with a four-primary photostimulator that provided independent control of rod and cone excitations. The results showed that dim surrounds ≤0.2 Td suppressed cone-mediated CFFs at ≥20 Td but not rod-mediated CFFs. These results can be understood in terms of peak amplitudes of photoreceptor impulse response functions under different stimulation conditions., (© 2012 Optical Society of America)

To model the light-guiding properties of a hexagonal array of dielectric cylinders, we have numerically solved Maxwell's equations with the finite-difference time-domain technique. The sizes and refractive indices of the cylinders are representative of those of the outer segments of the cone photoreceptors in the human central retina. In the array, light propagates predominantly as a "slow" mode, with a noticeable contribution of a "fast" mode, with the optical field localized in the intra- and inter-cylinder spaces, respectively. Interference between these modes leads to substantial (up to approximately 60%) axial oscillations in optical power within the cylinders. Our numerical model offered approximate dependence of the optical intensity distribution within the cylinders on their radii and separations.

We present evidence for the wavelength dependence of the directionality of light reflected from cone receptor cells (optical Stiles-Crawford effect): Blue light is more directional than red. According to the waveguide-scattering model of Marcos et al. [J. Opt. Soc. Am. A 15, 2012 (1998)], directionality is the sum of a waveguide component and a scattering component. The latter is proportional to 1 over wavelength squared, and it is related to the row-to-row spacing of the cone lattice. Our results allow a firm confirmation of Marcos et al.'s theory. For a 1.9-deg foveal area, group mean (n = 18) cone spacing was 3.42 microm, in good agreement with anatomical data. Group mean waveguide directionality was 0.077 mm(-2).

In the primate retina, H1 horizontal cells form an electrically coupled network and receive convergent input from long- (L-) and middle- (M-) wavelength-sensitive cones. Using an in vitro preparation of the intact retina to record the light-evoked voltage responses of H1 cells, we systematically varied the L- and M-cone stimulus contrast and measured the relative L- and M-cone input strength for 137 cells across 33 retinas from three Old World species (Macaca nemestrina, M. fascicularis, and Papio anubis). We found that the L- and the M-cone inputs were summed by the H1 cell in proportion to the stimulus cone contrast, which yielded a measure of what we term L- and M-cone contrast gain. The proportion of L-cone contrast gain was highly variable, ranging from 25% to 90% [mean +/- standard deviation, (60 +/- 14)%]. This variability was accounted for by retinal location within an individual, with the temporal retina showing a consistently higher percentage of L-cone gain, and by large overall variation across individuals, with the mean percentage of L-cone gain ranging from 32% to 80%. We hypothesize that the relative L- and M-cone contrast gain is determined simply by the relative number of L and M cones in the H1 cell's receptive field and that the variability in L- and M-cone contrast gain reflects a corresponding variability in the mosaic of L and M cones.

A primary goal of this study was to establish whether the magnitude of the short-wavelength-sensitive- (S-) cone signal into the yellow/blue (Y/B) mechanism was influenced by the absolute or the relative numbers of S cones. This was assessed by measuring the locus of unique green for various test sizes at four eccentric locations chosen to exploit differences in the underlying mosaic of S cones. In general, the locus of unique green was unaffected by test size, retinal quadrant, or rod input but was influenced by retinal eccentricity. The locus of unique green shifted to shorter wavelengths as retinal eccentricity increased from 1 degrees to 8 degrees. The data do not support a model whereby the S-cone signal is determined by the absolute number of S cones, but a model based on the relative number of S cones cannot be eliminated.

Flicker photometric measurements yield spectral sensitivity curves that are well fitted by sums of the spectral sensitivity curves of long-wavelength-sensitive (L) cones and middle-wavelength-sensitive (M) cones if the L cones are given twice the weight of the M cones. This result has been interpreted as implying that L cones are more numerous than M cones but is also consistent with a different numerical ratio, say, 1:1, and with the assignment of greater weight to the L cone input than to the M cone input by the mechanism subserving flicker photometry. Measurements of temporal sensitivity are presented for lights that modulate the inputs of either only the L cones or only the M cones. Sensitivity to modulation of the L cones is approximately twice that of modulation of the M cones at approximately 30 Hz, but that advantage disappears at approximately 2 Hz. Thus flicker sensitivity is equivocal with regard to cone numerosity. Electrophysiological, anatomical, and psychophysical evidence is reviewed, with particular weight placed on the statistics of color appearance of small, brief, monochromatic lights and on increment thresholds measured on the same observers. It is concluded that, in the central fovea, the ratio of L:M cone numbers is close to unity and may not be so variable as is usually supposed.

First measures of sensitivity and the need for a model to interpret them are addressed. Then modeling in the Fourier domain is promoted by a demonstration of how much an approach explains spatial sensitization and its dependence on luminance. Then the retinal illuminance and receptor absorptions produced by various stimuli are derived to foster interpretation of the neural mechanisms underlying various psychophysical phenomena. Finally, the sequence and the anatomical loci of the processes controlling visual sensitivity are addressed. It is concluded that multiplicative adaptation often has effects identical to response compression followed by subtractive adaptation and that, perhaps as a consequence, there is no evidence of retinal gain changes in human cone vision until light levels are well above those available in natural scenes and in most contemporary psychophysical experiments; that contrast gain control fine tunes sensitivity to patterns at all luminances; and that response compression, modulated by subtractive adaptation, predominates in the control of sensitivity in human cone vision.