The mechanism of human color vision and potential implanted devices for artificial color vision.

Vision plays a major role in perceiving external stimuli and information in our daily lives. The neural mechanism of color vision is complicated, involving the co-ordinated functions of a variety of cells, such as retinal cells and lateral geniculate nucleus cells, as well as multiple levels of the visual cortex. In this work, we reviewed the history of experimental and theoretical studies on this issue, from the fundamental functions of the individual cells of the visual system to the coding in the transmission of neural signals and sophisticated brain processes at different levels. We discuss various hypotheses, models, and theories related to the color vision mechanism and present some suggestions for developing novel implanted devices that may help restore color vision in visually impaired people or introduce artificial color vision to those who need it.

Evaluation of contrast sensitivity in patients with congenital red-green color vision deficiency.

PURPOSE: To evaluate mesopic and photopic contrast sensitivity in patients with congenital red-green color vision deficiency regarding with and without glare conditions and to compare these findings with age- and gender-matched healthy controls with normal color vision. METHODS: Patients with congenital red-green color vision deficiency and age- and gender-matched healthy controls were included in this cross-sectional comparative study. Contrast sensitivity measurements were taken from all subjects in 4 different conditions; binocular mesopic-without glare, mesopic-with glare, photopic-without glare, photopic-with glare, and the results were compared. RESULTS: Twenty one patients with color vision deficiency (13 deuteranopic, 8 protanopic) and 22 age- and gender-matched healthy controls were included in the study. The mean age was 35.2 ± 13.5 years in the protan group, 30.6 ± 7.7 years in the deutan group, 32.0 ± 8.8 years in the control group, and there was no significant difference in age between the groups (P > 0.05). The mean mesopic and photopic contrast sensitivity values of the groups at all spatial frequencies (1.5, 3, 6, 12, 18 cpd) were not statistically significant when evaluated by the multifactor repeated measures test of ANOVA to evaluate the effect of light conditions (with and without glare) (P > .05). CONCLUSION: Mesopic and photopic contrast sensitivity values of patients with congenital red-green color vision deficiency were similar to healthy controls regarding with and without glare conditions.

Visual evoked potentials in patients with congenital color vision deficiency.

BACKGROUND/AIM: Congenital color vision deficiency (CCVD) is an eye disease characterized by abnormalities in the cone cells in the photoreceptor layer. Visual evoked potentials (VEPs) are electrophysiological tests that physiologically examine the optic nerve, other visual pathways, and the visual cortex. The aim of this research was to determine whether there are VEP abnormalities in CCVD patients. METHODS: Patients with CCVD and healthy individuals were included in this prospective case-control study. Participants with eye disease or neurodegenerative disease were excluded from the study. Pattern reversal VEP (PVEP), flash VEP (FVEP), and optical coherence tomography were performed on all participants. RESULTS: Twenty healthy individuals (15 male) and 21 patients with CCVD (18 male) were included in the study. The mean ages of healthy individuals and patients with CCVD were 29.8 ± 9.6 and 31.1 ± 10.9 years (p = 0.804). Retinal nerve fiber layer thickness and central macular thickness values did not differ between the two groups. In PVEP, Right P100, Left N75, P100, N135 values were delayed in CCVD patients compared to healthy individuals (p = 0.001, p = 0.032, p = 0.003, p = 0.032). At least one PVEP and FVEP abnormality was present in nine (42.9%) and six (28.6%) of the patients, respectively. PVEP or FVEP abnormalities were found in 13 (61.9%) of the patients. CONCLUSION: This study indicated that there may be PVEP and FVEP abnormalities in patients with CCVD.

How the orientation of the color gamut of natural scenes influences color discrimination in red-green dichromacy.

In natural scenes, visual discrimination of colored surfaces by individuals with X-linked dichromacy is known to be only a little poorer than in normal trichromacy. This surprising result may be related to the properties of the colors of these scenes, like the shape and orientation of the color gamut, uneven frequency, and a considerable variation in lightness. It is unclear, however, how much each of these factors contributes to the small impairment in discrimination, in particular, what is the contribution of the orientation of the gamut. We measured the discrimination of colors from natural scenes by six normal trichromats and six dichromats. Colors were drawn either from the original color gamut of the scenes or from gamut-rotated versions of the scenes. Pairs of colors were randomly drawn from hyperspectral images of one rural and one urban environment and presented on a screen. As expected, dichromats were only a little poorer than normal trichromats at discrimination but the disadvantage varied systematically with the orientation of the color gamut by a factor of about three with a minimum around a yellow-green axis. Dichromats also took longer to respond, and the response times were modulated with the orientation of the color gamut in a similar way as the loss in discrimination. For the scenes tested here, these results imply an important impact of the orientation of the gamut on discrimination. They also indicate that the predominantly yellow-blue orientation of the gamut of natural scene might not be optimal for discrimination in dichromacy.

The genomic evolution of visual opsin genes in amphibians.

Among tetrapod (terrestrial) vertebrates, amphibians remain more closely tied to an amphibious lifestyle than amniotes, and their visual opsin genes may be adapted to this lifestyle. Previous studies have discussed physiological, morphological, and molecular changes in the evolution of amphibian vision. We predicted the locations of the visual opsin genes, their neighboring genes, and the tuning sites of the visual opsins, in 39 amphibian genomes. We found that all of the examined genomes lacked the Rh2 gene. The caecilian genomes have further lost the SWS1 and SWS2 genes; only the Rh1 and LWS genes were retained. The loss of the SWS1 and SWS2 genes in caecilians may be correlated with their cryptic lifestyles. The opsin gene syntenies were predicted to be highly similar to those of other bony vertebrates. Moreover, dual syntenies were identified in allotetraploid Xenopus laevis and X. borealis. Tuning site analysis showed that only some Caudata species might have UV vision. In addition, the S164A that occurred several times in LWS evolution might either functionally compensate for the Rh2 gene loss or fine-tuning visual adaptation. Our study provides the first genomic evidence for a caecilian LWS gene and a genomic viewpoint of visual opsin genes by reviewing the gains and losses of visual opsin genes, the rearrangement of syntenies, and the alteration of spectral tuning in the course of amphibians' evolution.

Horizontal-cell like Dm9 neurons in Drosophila modulate photoreceptor output to supply multiple functions in early visual processing.

Dm9 neurons in Drosophila have been proposed as functional homologs of horizontal cells in the outer retina of vertebrates. Here we combine genetic dissection of neuronal circuit function, two-photon calcium imaging in Dm9 and inner photoreceptors, and immunohistochemical analysis to reveal novel insights into the functional role of Dm9 in early visual processing. Our experiments show that Dm9 receive input from all four types of inner photoreceptor R7p, R7y, R8p, and R8y. Histamine released from all types R7/R8 directly inhibits Dm9 via the histamine receptor Ort, and outweighs simultaneous histamine-independent excitation of Dm9 by UV-sensitive R7. Dm9 in turn provides inhibitory feedback to all R7/R8, which is sufficient for color-opponent processing in R7 but not R8. Color opponent processing in R8 requires additional synaptic inhibition by R7 of the same ommatidium via axo-axonal synapses and the second Drosophila histamine receptor HisCl1. Notably, optogenetic inhibition of Dm9 prohibits color opponent processing in all types of R7/R8 and decreases intracellular calcium in photoreceptor terminals. The latter likely results from reduced release of excitatory glutamate from Dm9 and shifts overall photoreceptor sensitivity toward higher light intensities. In summary, our results underscore a key role of Dm9 in color opponent processing in Drosophila and suggest a second role of Dm9 in regulating light adaptation in inner photoreceptors. These novel findings on Dm9 are indeed reminiscent of the versatile functions of horizontal cells in the vertebrate retina.

A Bayesian observer model reveals a prior for natural daylights in hue perception.

Incorporating statistical characteristics of stimuli in perceptual processing can be highly beneficial for reliable estimation from noisy sensory measurements but may generate perceptual bias. According to Bayesian inference, perceptual biases arise from the integration of internal priors with noisy sensory inputs. In this study, we used a Bayesian observer model to derive biases and priors in hue perception based on discrimination data for hue ensembles with varying levels of chromatic noise. Our results showed that discrimination thresholds for isoluminant stimuli with hue defined by azimuth angle in cone-opponent color space exhibited a bimodal pattern, with lowest thresholds near a non-cardinal blue-yellow axis that aligns closely with the variation of natural daylights. Perceptual biases showed zero crossings around this axis, indicating repulsion away from yellow and attraction towards blue. These biases could be explained by the Bayesian observer model through a non-uniform prior with a preference for blue. Our findings suggest that visual processing takes advantage of knowledge of the distribution of colors in natural environments for hue perception.

Clinical Comparative Study of Shade Measurement Using Two Methods: Dental Guides and Spectrophotometry.

Tooth color is a determining factor in the fabrication of dental prostheses. The aim of the present study is to compare two measurement methods used in the field of dentistry: dental guides and spectrophotometry. A total of 2768 natural teeth were measured using the Vita Classical and Vita 3D-Master dental guides (Vita-Zahnfabrik, Bad Säckingen, Germany), as well as a Vita Easyshade Compact spectrophotometer (Vita-Zahnfabrik). The measurements were carried out by one operator under suitable illumination conditions at 5500 degrees Kelvin. The obtained results show that the measurements obtained with the Vita Classical dental guide classifies teeth into the A-B categories, while the spectrophotometer preferentially classifies teeth into the B-C categories. The correlation coefficients obtained with the dental guides ranged from -0.32 to -0.39 (p < 0.01), while those for the spectrophotometer ranged from -0.35 to -0.55 (p < 0.01). Therefore, we can conclude that the spectrophotometer is more reliable and reproducible in its measurements than the dental guides.

Color discrimination in fixed saturation level of patients with acute traumatic injury.

Introduction: Traumatic brain injury (TBI) is an important public health concern and that may lead to severe neural sequels, such as color vision deficits. Methods: We evaluated the color vision of 10 TBI patients with normal cognitive function using a color discrimination test in a fixed saturation level. We also analyzed computerized tomography scans to identify the local of the brain damages. Results: Four TBI patients that had lesions in brain areas of the ventral visual streams, five TBI patients had lesions inferred in brain areas of the dorsal visual stream, and one TBI patient had lesion in the occipital area. All the patients had cognitive and color vision screened and they had characterized the chromatic discrimination at high and low saturation. All participants had no significant cognitive impairment in the moment of the color vision test. Additionally, they had perfect performance for discrimination of chromatic stimulus at high saturation and similar to controls (n = 37 age-matched participants). Three of four TBI patients with lesions in the ventral brain and one patient with lesion in the occipital area had impairment of the chromatic discrimination at low saturation. All TBI patients with lesions in the dorsal brain had performance similar or slightly worse than the controls. Conclusion: Chromatic discrimination at low saturation was associated to visual damage in the ventral region of the brain and is a potential tool for functional evaluation of brain damage in TBI patients.

Secondary not subordinate: Opsin localization suggests possibility for color sensitivity in salticid secondary eyes.

The principal eyes of jumping spiders (Salticidae) integrate a dual-lens system, a tiered retinal matrix with multiple photoreceptor classes and muscular control of retinal movements to form high resolution images, extract color information, and dynamically evaluate visual scenes. While much work has been done to characterize these more complex principal anterior eyes, little work has investigated the three other pairs of simpler secondary eyes: the anterior lateral eye pair and two posterior (lateral and median) pairs of eyes. We investigated the opsin protein component of visual pigments in the eyes of three species of salticid using transcriptomics and immunohistochemistry. Based on characterization and localization of a set of three conserved opsins (Rh1 - green sensitive, Rh2 - blue sensitive, and Rh3 - ultraviolet sensitive) we have identified potential photoreceptors for blue light detection in the eyes of two out of three species: Menemerus bivittatus (Chrysillini) and Habrocestum africanum (Hasarinii). Additionally, the photoreceptor diversity of the secondary eyes exhibits more variation than previous estimates, particularly for the small, posterior median eyes previously considered vestigial in some species. In all three species investigated the lateral eyes were dominated by green-sensitive visual pigments (RH1 opsins), while the posterior median retinas were dominated by opsins forming short-wavelength sensitive visual pigments (e.g. RH2 and/or RH3/RH4). There was also variation among secondary eye types and among species in the distribution of opsins in retinal photoreceptors, particularly for the putatively blue-sensitive visual pigment formed from RH2. Our findings suggest secondary eyes have the potential for color vision, with observed differences between species likely associated with different ecologies and visual tasks.

Are ipRGCs involved in human color vision? Hints from physiology, psychophysics, and natural image statistics.

Human photoreceptors consist of cones, rods, and melanopsin-expressing intrinsically photosensitive retinal ganglion cells (ipRGCs). First studied in circadian regulation and pupillary control, ipRGCs project to a variety of brain centers suggesting a broader involvement beyond non-visual functions. IpRGC responses are stable, long-lasting, and with a particular codification of photoreceptor signals. In comparison with the transient and adaptive nature of cone and rod signals, ipRGCs' signaling might provide an ecological advantage to different attributes of color vision. Previous studies have indicated melanopsin's influence on visual responses yet its contribution to color perception in humans remains debated. We summarized evidence and hypotheses (from physiology, psychophysics, and natural image statistics) about direct and indirect involvement of ipRGCs in human color vision, by first briefly assessing the current knowledge about the role of melanopsin and ipRGCs in vision and codification of spectral signals. We then approached the question about melanopsin activation eliciting a color percept, discussing studies using the silent substitution method. Finally, we explore various avenues through which ipRGCs might impact color perception indirectly, such as through involvement in peripheral color matching, post-receptoral pathways, color constancy, long-term chromatic adaptation, and chromatic induction. While there is consensus about the role of ipRGCs in brightness perception, confirming its direct contribution to human color perception requires further investigation. We proposed potential approaches for future research, emphasizing the need for empirical validation and methodological thoroughness to elucidate the exact role of ipRGCs in human color vision.

Predicted effectiveness of EnChroma multi-notch filters for enhancing color perception in anomalous trichromats.

EnChroma filters are aids designed to improve color vision for anomalous trichromats. Their use is controversial because the results of lab-based assessments of their effectiveness have so far largely failed to agree with positive anecdotal reports. However, the effectiveness of EnChroma filters will vary depending on the conditions of viewing, including whether the stimuli are broadband reflective surfaces or colors presented on RGB displays, whether illumination spectra are broadband or narrowband, the transmission spectra of particular filters, and the cone spectral sensitivity functions of the observer. We created a model of anomalous trichromatic color vision to predict the effects of EnChroma filters on the color signals impaired in anomalous trichromacy. Using the model we varied illumination, filter type and observer cone sensitivity functions, and tested the effect of presenting colors as broadband reflective surfaces or on RGB displays. We also used hyperspectral images to assess the impact of the filters on anomalous trichromats' color vision for natural scenes. Model results predicted that the filters should be broadly effective at enhancing anomalous trichromats' equivalent to L/(L + M) chromatic contrasts under a range of viewing conditions, but are substantially more effective for deuteranomals than for protanomals. The filters are predicted to be more effective for broadband reflective surfaces presented under broadband illuminants than for surfaces presented under narrowband illuminants or for colors presented on RGB displays. Since the potential impacts of contrast adaptation and perceptual learning are not considered in the model, it needs to be empirically validated. Results of empirical tests of the effects of EnChroma filters on deuteranomalous color vision in comparison with model predictions are presented in an accompanying paper (Somers et al., in prep.).

Empirical tests of the effectiveness of EnChroma multi-notch filters for enhancing color vision in deuteranomaly.

Manufacturers of notch filter-based aids for color vision claim that their products can enhance color perception for people with anomalous trichromacy, a form of color vision deficiency (CVD). Anecdotal reports imply that people with CVD can have radically enhanced color vision when using the filters. However, existing empirical research largely focussed on the effect of notch filters on performance on diagnostic tests for CVD has not found that they have any substantial effect. Informed by a model of anomalous trichromatic color vision, we selected stimuli predicted to reveal the effects of EnChroma filters. Using these stimuli, we tested the ability of EnChroma filters to enhance color vision for 10 deuteranomalous trichromats in three experiments: 1. asymmetric color matching between test and control filter conditions, 2. color discrimination measured using four alternative forced-choice, and 3. color appearance measured using dissimilarity ratings to reconstruct subjective color spaces using multidimensional scaling. To investigate potential effects of long-term adaptation or perceptual learning, participants completed all three experiments at two time points, on first exposure to the filters, and after a week of regular use. We found a significant effect of the filters on color matches in the direction predicted by the model at both time points, implying that the filters can enhance the anomalous trichromatic color gamut. However, we found minimal effect of the filters on color discrimination at threshold. We found a significant effect of the filters in enhancing the appearance of colors along the red-green axis at the first time point, and a trend in the same direction at the second time point. Our results provide the first quantitative experimental evidence that notch filters can enhance color perception for anomalous trichromats.

Opsin mutants alter host plant selection by color vision in the nocturnal invasive pest Tuta absoluta.

In insects, vision is crucial in finding host plants, but its role in nocturnal insects is largely unknown. Vision involves responses to specific spectra of photon wavelengths and opsins plays an important role in this process. Long-wavelength sensitive opsin (LW opsin) and blue-sensitive opsin (BL opsin) are main visual opsin proteins and play important in behavior regulation.We used CRISPR/Cas9 technology to mutate the long-wavelength-sensitive and blue wavelength-sensitive genes and explored the role of vision in the nocturnal invasive pest Tuta absoluta. Light wave experiments revealed that LW2(-/-) and BL(-/-) mutants showed abnormal wavelength tropism. Both LW2 and BL mutations affected the preference of T. absoluta for the green environment. Mutations in LW2 and BL are necessary to inhibit visual attraction. The elimination of LW2 and BL affected the preference of leaf moths for green plants, and mutations in both induced a preference in moths for white plants. Behavioral changes resulting from LW2(-/-) and BL(-/-) mutants were not affected by sense of smell, further supporting the regulatory role of vision in insect behavior. To the best of our knowledge, this is the first study to reveal that vision, not smell, plays an important role in the host-seeking behavior of nocturnal insects at night, of which LW2 and BL opsins are key regulatory factors. These study findings will drive the development of the "vision-ecology" theory.

Cone-Opponent Ganglion Cells in the Primate Fovea Tuned to Noncardinal Color Directions.

A long-standing question in vision science is how the three cone photoreceptor types-long (L), medium (M), and short (S) wavelength sensitive-combine to generate our perception of color. Hue perception can be described along two opponent axes: red-green and blue-yellow. Psychophysical measurements of color appearance indicate that the cone inputs to the red-green and blue-yellow opponent axes are M vs. L + S and L vs. M + S, respectively. However, the "cardinal directions of color space" revealed by psychophysical measurements of color detection thresholds following adaptation are L vs. M and S vs. L + M. These cardinal directions match the most common cone-opponent retinal ganglion cells (RGCs) in the primate retina. Accordingly, the cone opponency necessary for color appearance is thought to be established in the cortex. While neurons with the appropriate M vs. L + S and L vs. M + S opponency have been reported in the retina and lateral geniculate nucleus, their existence continues to be debated. Resolving this long-standing debate is necessary because a complete account of the cone opponency in the retinal output is critical for understanding how downstream neural circuits process color. Here, we performed adaptive optics calcium imaging to noninvasively measure foveal RGC light responses in the living Macaca fascicularis eye. We confirm the presence of L vs. M + S and M vs. L + S neurons with noncardinal cone opponency and demonstrate that cone-opponent signals in the retinal output are more diverse than classically thought.

Spaco: A comprehensive tool for coloring spatial data at single-cell resolution.

Understanding tissue architecture and niche-specific microenvironments in spatially resolved transcriptomics (SRT) requires in situ annotation and labeling of cells. Effective spatial visualization of these data demands appropriate colorization of numerous cell types. However, current colorization frameworks often inadequately account for the spatial relationships between cell types. This results in perceptual ambiguity in neighboring cells of biological distinct types, particularly in complex environments such as brain or tumor. To address this, we introduce Spaco, a potent tool for spatially aware colorization. Spaco utilizes the Degree of Interlacement metric to construct a weighted graph that evaluates the spatial relationships among different cell types, refining color assignments. Furthermore, Spaco incorporates an adaptive palette selection approach to amplify chromatic distinctions. When benchmarked on four diverse datasets, Spaco outperforms existing solutions, capturing complex spatial relationships and boosting visual clarity. Spaco ensures broad accessibility by accommodating color vision deficiency and offering open-accessible code in both Python and R.

Expression of red/green-cone opsin mutants K82E, P187S, M273K result in unique pathobiological perturbations to cone structure and function.

Long-and middle-wavelength cone photoreceptors, which are responsible for our visual acuity and color vision, comprise ~95% of our total cone population and are concentrated in the fovea of our retina. Previously, we characterized the disease mechanisms of the L/M-cone opsin missense mutations N94K, W177R, P307L, R330Q and G338E, all of which are associated with congenital blue cone monochromacy (BCM) or color-vision deficiency. Here, we used a similar viral vector-based gene delivery approach in M-opsin knockout mice to investigate the pathogenic consequences of the BCM or color-vision deficient associated L-cone opsin (OPN1LW) mutants K82E, P187S, and M273K. We investigated their subcellular localization, the pathogenic effects on cone structure, function, and cone viability. K82E mutants were detected predominately in cone outer segments, and its expression partially restored expression and correct localization of cone PDE6α' and cone transducin γ. As a result, K82E also demonstrated the ability to mediate cone light responses. In contrast, expression of P187S was minimally detected by either western blot or by immunohistochemistry, probably due to efficient degradation of the mutant protein. M273K cone opsin appeared to be misfolded as it was primarily localized to the cone inner segment and endoplasmic reticulum. Additionally, M273K did not restore the expression of cone PDE6α' and cone transducin γ in dorsal cone OS, presumably by its inability to bind 11-cis retinal. Consistent with the observed expression pattern, P187S and M273K cone opsin mutants were unable to mediate light responses. Moreover, expression of K82E, P187S, and M273K mutants reduced cone viability. Due to the distinct expression patterns and phenotypic differences of these mutants observed in vivo, we suggest that the pathobiological mechanisms of these mutants are distinct.

Validation of a New Digital and Automated Color Perception Test.

Although color vision deficiencies are very prevalent, there are no ideal methods for assessing color vision in all environments. We compared a new digital and automated method that quantifies color perception for the three protan, deutan, and tritan axes with two of the most commonly used color tests in daily practice: the Ishihara 38 plates test and the Farnsworth-Munsell 100-Hue test. One hundred patients underwent a triple examination composed of the new DIVE Color Test, the Ishihara test, and the Farnsworth-Munsell 100-Hue test. The DIVE Color Test was performed twice in forty participants to assess its repeatability. In the trichromatic group, the mean age stood at 20.57 ± 9.22 years compared with 25.99 ± 15.86 years in the dyschromatic group. The DIVE and Ishihara tests exhibited excellent agreement in identifying participants with color deficiency (Cohen's kappa = 1.00), while it was 0.81 when comparing DIVE and Farnsworth. The correlation between the global perception values of Farnsworth (TES) and DIVE (GCS) was 0.80. The repeatability of the DIVE Color Test was high according to Bland-Altman analysis with an intraclass correlation coefficient of 0.83. According to Ishihara, the DIVE Color Test proved to be an effective and reproducible tool for red-green color vision deficiency detection, capable of determining the severity of the defect in each of the three axes faster and more accurately than both Ishihara and Farnsworth.

Prevalence and genotypic frequency of color vision defects among primary schoolchildren in Adama Town, Eastern Ethiopia.

Color vision deficiency is a common X-linked genetic disorder affecting the day-to-day lives of individuals, in which school-aged children's academic performance can be negatively affected. The aim of this study was to evaluate the prevalence and genotypic frequency of congenital color vision defects (CVD), among primary schoolchildren in Adama, Ethiopia. A school-based cross-sectional study design was used. Students were purposively selected based on their ethnicity but were randomly selected from their sections, resulting in a final sample size estimated at 846 schoolchildren who had received informed consent from their families. Data was gathered using the Ishihara color vision test, 38-plate edition. The result of the study revealed that the total prevalence of CVD was much higher (5.6%) among the male children than that of the females, which was only about 1.79%. The prevalence rates of CVD among the targeted ethnic groups were found to be the highest among Amhara (7.45%) > Oromo (5.00%) > Gurage (2.13%) children, respectively, in descending order. 62.76% of the study subjects were homozygous dominant (AA), followed by those with a heterozygous genotype (Aa) (32.51%), and the remaining 4.73% had recessive (aa) genes.