Extensive cone-dependent spectral opponency within a discrete zone of the lateral geniculate nucleus supporting mouse color vision.

Color vision, originating with opponent processing of spectrally distinct photoreceptor signals, plays important roles in animal behavior.1-4 Surprisingly, however, comparatively little is understood about color processing in the brain, including in widely used laboratory mammals such as mice. The retinal gradient in S- and M-cone opsin (co-)expression has traditionally been considered an impediment to mouse color vision.5-8 However, recent data indicate that mice exhibit robust chromatic discrimination within the central-upper visual field.9 Retinal color opponency has been reported to emerge from superimposing inhibitory surround receptive fields on the cone opsin expression gradient, and by introducing opponent rod signals in retinal regions with sparse M-cone opsin expression.10-13 The relative importance of these proposed mechanisms in determining the properties of neurons at higher visual processing stages remains unknown. We address these questions using multielectrode recordings from the lateral geniculate nucleus (LGN) in mice with altered M-cone spectral sensitivity (Opn1mwR) and multispectral stimuli that allow selective modulation of signaling by individual opsin classes. Remarkably, we find many (∼25%) LGN cells are color opponent, that such cells are localized to a distinct medial LGN zone and that their properties cannot simply be explained by the proposed retinal opponent mechanisms. Opponent responses in LGN can be driven solely by cones, independent of cone-opsin expression gradients and rod input, with many cells exhibiting spatially congruent antagonistic receptive fields. Our data therefore suggest previously unidentified mechanisms may support extensive and sophisticated color processing in the mouse LGN.

Variations in photoreceptor throughput to mouse visual cortex and the unique effects on tuning.

Visual input to primary visual cortex (V1) depends on highly adaptive filtering in the retina. In turn, isolation of V1 computations requires experimental control of retinal adaptation to infer its spatio-temporal-chromatic output. Here, we measure the balance of input to mouse V1, in the anesthetized setup, from the three main photoreceptor opsins-M-opsin, S-opsin, and rhodopsin-as a function of two stimulus dimensions. The first dimension is the level of light adaptation within the mesopic range, which governs the balance of rod and cone inputs to cortex. The second stimulus dimension is retinotopic position, which governs the balance of S- and M-cone opsin input due to the opsin expression gradient in the retina. The fitted model predicts opsin input under arbitrary lighting environments, which provides a much-needed handle on in-vivo studies of the mouse visual system. We use it here to reveal that V1 is rod-mediated in common laboratory settings yet cone-mediated in natural daylight. Next, we compare functional properties of V1 under rod and cone-mediated inputs. The results show that cone-mediated V1 responds to 2.5-fold higher temporal frequencies than rod-mediated V1. Furthermore, cone-mediated V1 has smaller receptive fields, yet similar spatial frequency tuning. V1 responses in rod-deficient (Gnat1-/-) mice confirm that the effects are due to differences in photoreceptor opsin contribution.

Summation of Temporal L-Cone- and M-Cone-Contrast in the Magno- and Parvocellular Retino-Geniculate Systems in Glaucoma.

Purpose: The purpose of this study was to characterize summation of temporal L- and M-cone contrasts in the parvo- (P-) and magnocellular (M-) pathways in glaucoma and the relationship between the respective temporal contrast sensitivities (tCS) and clinical parameters. Methods: Perifoveal tCS to isolated or combined L- and M-cone contrasts (with different contrast ratios, and therefore different luminance and chromatic components) were measured at different temporal frequencies (at 1 or 2 Hz and at 20 Hz) using triple silent substitution in 73 subjects (13 healthy, 25 with glaucoma, and 35 with perimetric glaucoma). A vector summation model was used to analyze whether perception was driven by the P-pathway, the M-pathway, or both. Using this model, L- and M-cone input strengths (AL, AM) and phase differences between L- and M-cone inputs were estimated. Results: Perception was always mediated by the P-pathway at low frequencies, as indicated by a median phase angle of 179.84 degrees (cone opponency) and a median AL/AM ratio of 1.04 (balanced L- and M-cone input strengths). In contrast, perception was exclusively mediated by the M-pathway at higher frequencies (input strength not balanced: AL/AM = 2.94, median phase angles = 130.17 degrees). Differences in phase were not significant between diagnosis groups (Kruskal-Wallis = 0.092 for P- and 0.35 for M-pathway). We found differences between groups only for the M-pathway (L-cone tCS deviations at 20 Hz were significantly lower in the patients with glaucoma P = 0.014, with a strong tendency in M-cones P = 0.049). L-cone driven tCS deviations at 20 Hz were linearly correlated with perimetric mean defect (MD) and quadratically correlated with retinal nerve fiber layer (RNFL) thickness. Conclusions: Unaltered phase angles between L- and M-cone inputs in glaucoma indicated intact temporal processing. Only in the M-pathway, contrast sensitivity deviations were closely related to diagnosis group, MD, and RNFL thickness, indicating M-pathway involvement.

Methods for determining equiluminance in terms of L/M cone ratios.

Heterochromatic flicker photometry (HFP), minimum motion (MM), and minimally distinct border (MDB) settings have often been used to determine equiluminance, a relative intensity setting for two chromaticities that, in theory, eliminates the responses of a luminance or achromatic psychophysical mechanism. These settings have been taken to reflect the relative contribution of the long (L) and medium (M) wavelength cones to luminance, which varies widely across individuals. The present study compares HFP, MM, and MDB using stimuli that do not modulate the short (S) wavelength cones, in both practiced and naïve observers. MDB was performed with both flashed and steadily viewed stimuli. Results are represented in the (∆L/L, ∆M/M) plane of cone contrast space. Considering both practiced and naïve observers, both MM and HFP had excellent within-subject precision and high test-retest reliability, whereas HFP also had low between-subject variability. The MDB tasks were less reliable and less precise. The mean L:M contrast ratios at equiluminance were lower for the two temporal tasks (HFP and MM) compared to the spatial tasks (MDB), perhaps consistent with the existence of multiple luminance mechanisms. Overall, the results suggest that the best method for determining equiluminance is HFP, with MM being a close second.

Gonadotropin-releasing hormone-independent effects of recombinant vertebrate ancient long opsin in the goldfish Carassius auratus reveal alternative reproduction pathways.

Vertebrate ancient long (VAL)-opsin is a green-sensitive photoreceptor that shows high sequence similarity to vertebrate ancient opsin, which is considered to play a role in sexual maturation via gonadotropin-releasing hormone (GnRH); however, the role of VAL-opsin in vertebrate sexual maturity remains unclear. Therefore, we investigated the possible role of VAL-opsin in reproduction in the goldfish Carassius auratus under a state of GnRH inhibition. Goldfish were injected with recombinant VAL-opsin protein (0.5 µg/g body mass) and/or the GnRH antagonist cetrorelix (0.5 µg/fish), and changes in the mRNA expression levels of genes associated with goldfish reproduction were measured by quantitative polymerase chain reaction, including those involved in the hypothalamus-pituitary-gonad (HPG) axis, VAL-opsin, GnRH, the gonadotropins (GTHs) luteinizing hormone and follicle-stimulating hormone, and estrogen receptor (ER). Moreover, the fish were irradiated with a green light-emitting diode (520 nm) to observe the synergistic effect on the HPG axis with VAL-opsin. Green LED exposure significantly and slightly increased the VAL-opsin and GnRH levels, respectively; however, these effects were blocked in groups injected with cetrorelix at all time points. Cetrorelix significantly decreased the mRNA levels of GTHs and ER, whereas these hormones recovered by co-treatment with VAL-opsin. These results indicate that green LED is an effective light source to promote the expression of sex hormones in fish. Moreover, VAL-opsin not only affects activity of the HPG axis but also appears to act on the pituitary gland directly to stimulate a new sexual maturation pathway that promotes the secretion of GTHs independent of GnRH.

Cones Support Alignment to an Inconsistent World by Suppressing Mouse Circadian Responses to the Blue Colors Associated with Twilight.

In humans, short-wavelength light evokes larger circadian responses than longer wavelengths [1-3]. This reflects the fact that melanopsin, a key contributor to circadian assessments of light intensity, most efficiently captures photons around 480 nm [4-8] and gives rise to the popular view that "blue" light exerts the strongest effects on the clock. However, in the natural world, there is often no direct correlation between perceived color (as reported by the cone-based visual system) and melanopsin excitation. Accordingly, although the mammalian clock does receive cone-based chromatic signals [9], the influence of color on circadian responses to light remains unclear. Here, we define the nature and functional significance of chromatic influences on the mouse circadian system. Using polychromatic lighting and mice with altered cone spectral sensitivity (Opn1mwR), we generate conditions that differ in color (i.e., ratio of L- to S-cone opsin activation) while providing identical melanopsin and rod activation. When biased toward S-opsin activation (appearing "blue"), these stimuli reliably produce weaker circadian behavioral responses than those favoring L-opsin ("yellow"). This influence of color (which is absent in animals lacking cone phototransduction; Cnga3-/-) aligns with natural changes in spectral composition over twilight, where decreasing solar angle is accompanied by a strong blue shift [9-11]. Accordingly, we find that naturalistic color changes support circadian alignment when environmental conditions render diurnal variations in light intensity weak/ambiguous sources of timing information. Our data thus establish how color contributes to circadian entrainment in mammals and provide important new insight to inform the design of lighting environments that benefit health.

The retinal pigments of the whale shark (Rhincodon typus) and their role in visual foraging ecology.

The spectral tuning properties of the whale shark (Rhincodon typus) rod (rhodopsin or Rh1) and long-wavelength-sensitive (LWS) cone visual pigments were examined to determine whether these retinal pigments have adapted to the broadband light spectrum available for surface foraging or to the narrowband blue-shifted light spectrum available at depth. Recently published whale shark genomes have identified orthologous genes for both the whale shark Rh1 and LWS cone opsins suggesting a duplex retina. Here, the whale shark Rh1 and LWS cone opsin sequences were examined to identify amino acid residues critical for spectral tuning. Surprisingly, the predicted absorbance maximum (λmax) for both the whale shark Rh1 and LWS visual pigments is near 500 nm. Although Rh1 λmax values near 500 nm are typical of terrestrial vertebrates, as well as surface foraging fish, it is uncommon for a vertebrate LWS cone pigment to be so greatly blue-shifted. We propose that the spectral tuning properties of both the whale shark Rh1 and LWS cone pigments are most likely adaptations to the broadband light spectrum available at the surface. Whale shark melanopsin (Opn4) deactivation kinetics was examined to better understand the underlying molecular mechanisms of the pupillary light reflex. Results show that the deactivation rate of whale shark Opn4 is similar to the Opn4 deactivation rate from vertebrates possessing duplex retinae and is significantly faster than the Opn4 deactivation rate from an aquatic rod monochromat lacking functional cone photoreceptors. The rapid deactivation rate of whale shark Opn4 is consistent with a functional cone class and would provide the animal with an exponential increase in the number of photons required for photoreceptor signaling when transitioning from photopic to scotopic light conditions, as is the case when diving.

The Retinal Basis of Vertebrate Color Vision.

The jawless fish that were ancestral to all living vertebrates had four spectral cone types that were probably served by chromatic-opponent retinal circuits. Subsequent evolution of photoreceptor spectral sensitivities is documented for many vertebrate lineages, giving insight into the ecological adaptation of color vision. Beyond the photoreceptors, retinal color processing is best understood in mammals, especially the blueON system, which opposes short- against long-wavelength receptor responses. For other vertebrates that often have three or four types of cone pigment, new findings from zebrafish are extending older work on teleost fish and reptiles to reveal rich color circuitry. Here, horizontal cells establish diverse and complex spectral responses even in photoreceptor outputs. Cone-selective connections to bipolar cells then set up color-opponent synaptic layers in the inner retina, which lead to a large variety of color-opponent channels for transmission to the brain via retinal ganglion cells.

Electrodiagnosis of dichromacy.

Retinal and cortical signals initiated by a single cone type can be recorded using the spectral compensation (or silent substitution) paradigm. Moreover, responses to instantaneous excitation increments combined with gradual excitation decreases are dominated by the response to the excitation increment. Similarly, the response to a sudden excitation decrement dominates the overall response when combined with a gradual excitation increase. Here ERGs and VEPs were recorded from 34 volunteers [25.9 ±â€¯10.4 years old (mean ±â€¯1 SD); 25 males, 9 females] to sawtooth flicker (4 Hz) stimuli that elicited L- or M-cone responses using triple silent substitution. The mean luminance (284 cd/m2) and the mean chromaticity (x = 0.5686, y = 0.3716; CIE 1931 color space) remained constant and thus the state of adaptation was the same in all conditions. Color discrimination thresholds along protan, deutan, and tritan axes were obtained from all participants. Dichromatic subjects were genetically characterized by molecular analysis of their opsin genes. ERG responses to L-cone stimuli were absent in protanopes whereas ERG responses to M-cone stimuli were strongly reduced in deuteranopes. Dichromats showed generally reduced VEP amplitudes. Responses to cone-specific stimuli obtained with standard electrophysiological methods may give the same classification as that obtained with the Cambridge Colour Test and in some cases with the genetic analysis of the L- and M-opsin genes. Therefore, cone-specific ERGs and VEPs may be reliable methods to detect cone dysfunction. The present data confirm and emphasize the potential use of cone-specific stimulation, combined with standard visual electrodiagnostic protocols.

Spatial visual function in anomalous trichromats: Is less more?

Color deficiency is a common inherited disorder affecting 8% of Caucasian males with anomalous trichromacy (AT); it is the most common type of inherited color vision deficiency. Anomalous trichromacy is caused by alteration of one of the three cone-opsins' spectral sensitivity; it is usually considered to impose marked limitations for daily life as well as for choice of occupation. Nevertheless, we show here that anomalous trichromat subjects have superior basic visual functions such as visual acuity (VA), contrast sensitivity (CS), and stereo acuity, compared with participants with normal color vision. Both contrast sensitivity and stereo acuity performance were correlated with the severity of color deficiency. We further show that subjects with anomalous trichromacy exhibit a better ability to detect objects camouflaged in natural gray scale figures. The advantages of color-deficient subjects in spatial vision performance could explain the relatively high prevalence of color-vision polymorphism in humans.

Sensitivity to S-Cone Stimuli and the Development of Myopia.

Purpose: Longitudinal chromatic aberration (LCA) is a color signal available to the emmetropization process that causes greater myopic defocus of short wavelengths than long wavelengths. We measured individual differences in chromatic sensitivity to explore the role LCA may play in the development of refractive error. Methods: Forty-four observers were tested psychophysically after passing color screening tests and a questionnaire for visual defects. Refraction was measured and only subjects with myopia or hyperopia without severe astigmatism participated. Psychophysical detection thresholds for 3 cyc/deg achromatic, L-, M-, and S-cone-isolating Gabor patches and low-frequency S-cone increment (S+) and decrement (S-) blobs were measured. Parametric Pearson correlations for refractive error versus threshold were calculated and nonparametric bootstrap 95% percentage confidence intervals (BCIs) for r were computed. Results: S-cone Gabor detection thresholds were higher than achromatic, L-, and M-cone Gabors. S-cone Gabor thresholds were higher than either S+ or S- blobs. These results are consistent with studies using smaller samples of practiced observers. None of the thresholds for the Gabor stimuli were correlated with refractive error (RE). A negative correlation with RE was observed for both S+ (r = -0.28; P = 0.06; BCI: r = -0.5, -0.04) and S- (r = -0.23; P = 0.13; BCI = -0.46, 0.01) blobs, although this relationship did not reach conventional statistical significance. Conclusions: Thresholds for S+ and S- stimuli were negatively related to RE, indicating that myopes may have reduced sensitivity to low spatial frequency S-cone stimuli. This reduced S-cone sensitivity might have played a role in their failure to emmetropize normally.

Cone Vision Changes in the Enhanced S-Cone Syndrome Caused by NR2E3 Gene Mutations.

Purpose: To determine the progression of cone vision loss in patients with recessive disease from NR2E3 gene mutations. Methods: Patients with NR2E3 mutations (n = 37) were studied as a retrospective observational case series clinically and with chromatic static perimetry. Patients were investigated cross-sectionally, and a subset was followed longitudinally. Results: Patients showed a range of visual acuities; there was no clear relationship to age. With kinetic perimetry (V4e target), a full field could be retained over many years. Other patients showed progression from a full field, with or without pericentral scotomas, to a small central island. Three patterns of S-cone function were defined, based on percentage of hypersensitive S-cone loci in the field. From occupying most of the visual field, hyperfunctioning S-cone loci could diminish in percent, remaining largely in the periphery. Normal S-cone functioning then dominates, followed by the appearance of an annular region of abnormal S-cone loci approximately 10° to 40° from the fovea. Overall, S-cone sensitivity declined 2.6 times faster than L/M-cone sensitivity. Conclusions: Murine proof-of-concept studies suggest that clinical trials of patients with NR2E3 mutations may be forthcoming. Patterns of S-cone hyperfunction across the field would serve as a means to categorize patients as entry criteria or cohort selection in clinical trials. S-cone perimetry can be measured in the clinic and would be the logical efficacy monitor for therapeutic strategies. Given further understanding of the natural history of the disease, targeting the annular region of S-cone dysfunction for a focal therapy or for monitoring in a retina-wide intervention warrants consideration.

Are red, yellow, green, and blue perceptual categories?

This study investigated categorical perception for unique hues in order to establish a relationship between color appearance, color discrimination, and low-level (second-stage) mechanisms. We tested whether pure red, yellow, green, and blue, (unique hues) coincide with troughs, and their transitions (binary hues) with peaks of sensitivity in DKL-space. Results partially confirmed this idea: JNDs demarcated perceptual categories at the binary hues around green, blue and less clearly around yellow, when colors were isoluminant with the background and when accounting for the overall variation of sensitivity by fitting an ellipse. The categorical JND pattern for those three categories was in line with the effect of the second-stage mechanisms. In contrast, the results for unique red, binary red-yellow, and the JNDs for dark colors clearly contradicted categorical perception. There was a JND maximum around the center of red and JNDs strongly decreased away from the center. Although this observation alone would also be in line with categorical perception; unique red was shifted away from the center towards yellow so that unique red was close to the minimum instead of the maximum JND, hence contradicting categorical perception. In addition, we also showed that observers do not adjust unique hues more consistently than binary hues, confirming a previous study. Taken together, our findings suggest that some of the unique hues could be inherent in the early stages of color processing. At the same time, they also raise questions about complex effects of lightness, chroma and instructions on the measurements of JNDs and unique hues.

M to L cone ratios determine eye sizes and baseline refractions in chickens.

Following a hypothesis raised by M. and J. Neitz, Seattle, we have tested whether the abundance and the ratio of Long wavelength-sensitive (L) to Middle wavelength-sensitive (M) cones may affect eye size and development of myopia in the chicken. Fourteen chickens were treated with frosted plastic diffusers in front of one eye on day 10 post-hatching for a period of 7 days to induce deprivation myopia. Ocular dimensions were measured by A-scan ultrasonography at the beginning and at the end of the treatment and development of refractive state was tracked using infrared photorefraction. At the end of the treatment period, L and M cone densities and ratios were analyzed in retinal flat mounts of both myopic and control eyes, using the red and yellow oil droplets as markers. Because large numbers of cones were counted (>10000), software was written in Visual C++ for automated cone detection and density analysis. (1) On average, 9.7 ± 1.7D of deprivation myopia was induced in 7 days (range from 6.8D to 13.7D) with an average increase in axial length by 0.65 ± 0.20 mm (range 0.42 mm-1.00 mm), (2) the increase in vitreous chamber depth was correlated with the increase in myopic refractive error, (3) average central M cone densities were 10,498 cells/mm2, and L cone densities 9574 cells/mm2. In the periphery, M cone densities were 6343 cells/mm2 and L cones 5735 cells/mm2 (4) M to L cone ratios were highly correlated in both eyes of each animal (p < 0.01 in all cases), (5) the most striking finding was that ratios of M to L cones were significantly correlated with vitreous chamber depths and refractive states in the control eyes with normal vision, both in the central and peripheral retinas (p < 0.05 to p < 0.01), (6) M to L cone ratios did however not predict the amount of deprivation myopia that could be induced. M and L cone ratios are most likely genetically determined in each animal. The more L cones, the deeper the vitreous chambers and the more myopic were the refractions in eyes. M to L cone ratios may determine the set point of emmetropization and thereby ultimately the probability of becoming myopic. Deprivation myopia was not determined by M to L cone ratios.

Paradoxical pupil responses to isolated M-cone increments.

M-cone onsets appear dimmer than the background and elicit electroretinograms (ERGs) resembling the light offset response. We sought a corresponding anomalous pupillary light reflex (PLR) using a 4-primary ganzfeld as stimulator and pupillometer. Increments and decrements of white light were compared with M- and L-cone onsets and offsets using silent substitution. Luminance bias (LB) could be added to or subtracted from the cone-isolating stimuli. There was a normal PLR to L-cone increments, but the pupil constricted mainly to M-cone decrements. Changing LB produced a neutral point where on and off responses were balanced. The results reflect ERG and psychophysical studies. This observation may be linked to the antagonistic nature of the M-cone input to cone opponent mechanisms.

Photoreceptor-specific light adaptation of critical flicker frequency in trichromat and dichromat observers.

The silent substitution paradigm offers possibilities to investigate and compare the temporal properties of mechanisms driven by single photoreceptor types, including the critical flicker frequency (CFF), in which the state of adaptation can be kept as invariant. We have (1) measured CFFs using triple silent substitutions to isolate L-, M-, and S-cone as well as rod-driven pathways under identical mean luminances and chromaticities; (2) repeated the CFF measurements at different mean luminances in order to validate the Ferry-Porter law (stating that the relationship between CFF and the log retinal illuminance-log I-is linear); and (3) compared these CFF versus log I functions for L-, M-, S-cone-, and rod-isolating stimuli for five trichromats and four X-linked dichromats (two protanopes, two deuteranopes). We show that the effects of luminance on the CFFs with silent substitution are comparable to those measured previously with chromatic stimuli. We found that M-cone-driven CFFs are smaller in trichromats than in protanopes. Furthermore, the slopes of the M-cone-driven CFF versus log I functions are smaller in trichromats. Possibly, the lacking L-cones are replaced by M-cones in these two protanopes and the CFF depends on cone density. Furthermore, we found that in trichromats, the slopes of the CFF-log I functions are smaller for M-cone- than for L-cone-isolating stimuli. This contradicts the current interpretation of the CFF-log I functions for chromatic stimuli, which states that CFF is mediated by the most strongly modulated photoreceptor type. Thus, the larger slopes that were previously found with medium-wavelength chromatic stimuli compared with long-wavelength chromatic stimuli seem to be the result of an addition of signals from different photoreceptors and do not necessarily result from M-cones being inherently faster.

Contrast-dependent red-green balance shifts depend on S-cone activity.

Previous research from our lab has established that red-green-balanced yellow targets become greenish-brown as surround luminance increases, while red-green-balanced brown targets become reddish-yellow as surround luminance decreases. To help assess the generality and underlying processes of this contrast-dependent red-green hue shift, we investigated red-green hue shifts for target stimuli that appeared achromatic or blue as well as yellow/brown. Results confirmed that the red-green hue shift was largest for yellow/brown targets and was progressively reduced for achromatic and blue targets as target excitation of S cones increased. The magnitude of the hue shift could be predicted by the S/(L+M) excitation of the target when bright white surrounds are used. The hue shift also requires that the target and surround are presented to the same eye, consistent with processing in monocular pathways. Increased S-cone excitation by the surround was associated with red-green hue shifts for all targets equally. Thus, S-cone signals from bright white surrounds might play a role in the contrast-dependent red-green hue shift, but the source of the variation of the magnitude of the hue shift with variations in target S-cone excitation when presented on those surrounds is unknown.

Human S-cone electroretinograms obtained by silent substitution stimulation.

We used triple silent substitution stimuli to characterize human S-cone electroretinograms (ERGs) in normal trichromats. Short-wavelength-cone (S-cone) ERGs were found to have different morphological features and temporal frequency response characteristics compared to ERGs derived from L-cones, M-cones, and rod photoreceptors in normal participants. Furthermore, in two cases of retinal pathology, blue cone monochromatism (BCM) and enhanced S-cone syndrome (ESCS), S-cone ERGs elicited by our stimuli were preserved and enhanced, respectively. The results from both normal and pathological retinae demonstrate that triple silent substitution stimuli can be used to generate ERGs that provide an assay of human S-cone function.

Contrast-dependent red-green hue shift.

On bright surrounds, red-green-balanced yellow targets become greenish brown with decreased target luminance, and red-green-balanced brown targets become reddish yellow with increased target luminance. These effects imply luminance- and/or contrast-dependent weighting of M- and L-cone signals in post-receptoral pathways. We show psychophysically that luminance contrast between the surround and the target is the primary determinant of the magnitude of red-green hue shift, requiring surround luminance at least twice the target luminance and increasing with further increases of surround/target contrast. There is a much smaller effect of absolute stimulus luminance, with dimmer stimuli showing slightly larger hue shifts. To evaluate a possible retinal origin of the changes in cone-signal weightings underlying the hue shift, we recorded spike responses from both ON- and OFF-center midget ganglion cells in peripheral primate retina. We found no evidence that the relative strength of L- and M-cone post-receptoral responses changed systematically with change of surround irradiance. Nor was there any systematic difference between ON- and OFF-subtypes. This suggests that the change in cone signal weighting occurs later in the visual system.

Asymmetric high-contrast masking in S cone increment and decrement pathways.

Physiological, anatomical, and psychophysical evidence points to important differences between visual processing of short-wave cone increments and decrement (S+ and S-) stimuli. The present study uses the pedestal discrimination paradigm to investigate potential differences, using S+ and S- tests presented on (L)ong-wave, (M)edium-wave, S, L+M, L-M, and achromatic pedestals, of both contrast polarities. Results show that high contrast 'purplish' (S+ or -(L+M)) pedestals produce substantially more masking of both S+ and S- tests than 'yellowish' (S- or +(L+M)) pedestals do. The other pedestals produce no masking. These findings suggest greater nonlinearity - either a static nonlinearity or contrast gain control - in the mechanisms responsible for the 'purplish' polarity, likely the S ON pathway.