Trichromatic Research Articles

Assessing the severity of colour vision loss – implications for occupational environments

AbstractColour is often used in safety‐critical environments to code and signal information. Colour is also used to enhance target ‘conspicuity’ and visual performance in ‘grouping’, segmentation and pop‐out tasks. Colour vision (CV) is not a key requirement in everyday life, but in some safety‐critical jobs, it can be a matter of life and death. The absence of normally functioning chromatic mechanisms leads to reduced chromatic sensitivity and also to changes in the perceived colour of objects. CV is needed in some occupations, but the majority of subjects with congenital colour deficiency (CCD) retain some red / green (RG) CV. When the latter is combined with normal, yellow / blue (YB) colour signals, many CCD subjects are able to carry out suprathreshold, colour‐related tasks as well as normal trichromats. Since severity of loss in congenital deficiency extends from almost normal to complete absence of CV, it is important to determine accurately severity of CV loss. Empirical studies carried out in some occupations established the degree of CV loss that can be classed as ‘safe’. This led to the establishment of minimum requirements that can be enforced. In addition to safety, the overall aim is to ensure that all subjects with anomalous trichromatic colour vision who can perform the safety‐critical, colour tasks in a given job as well as normal trichromats pass and do not end up being discriminated against unfairly on the basis of their CCD. Until very recently, it has not been possible to achieve this aim, simply because many existing CV tests lack sensitivity and / or specificity and also fail to quantify reliably the severity of CV loss. Improved understanding of CV and the recent development of sensitive and specific CV tests have transformed traditional colour assessment (Br Med Bull. 2017;122 (1):51–77). Results will be presented to show how a single colour assessment test can be used to establish the applicant’s class of CV and to quantify the severity of loss.

An arbitrary-spectrum spatial visual stimulator for vision research.

Visual neuroscientists require accurate control of visual stimulation. However, few stimulator solutions simultaneously offer high spatio-temporal resolution and free control over the spectra of the light sources, because they rely on off-the-shelf technology developed for human trichromatic vision. Importantly, consumer displays fail to drive UV-shifted short wavelength-sensitive photoreceptors, which strongly contribute to visual behaviour in many animals, including mice, zebrafish and fruit flies. Moreover, many non-mammalian species feature more than three spectral photoreceptor types. Here, we present a flexible, spatial visual stimulator with up to six arbitrary spectrum chromatic channels. It combines a standard digital light processing engine with open source hard- and software that can be easily adapted to the experimentalist's needs. We demonstrate the capability of this general visual stimulator experimentally in the in vitro mouse retinal whole-mount and the in vivo zebrafish. With this work, we intend to start a community effort of sharing and developing a common stimulator design for vision research.

Empirical Formulas for Estimating Backscattering and Absorption Coefficients in Complex Waters from Remote-Sensing Reflectance Spectra and Examples of Their Application.

Many standard methods used for the remote sensing of ocean colour have been developed, though mainly for clean, open ocean waters. This means that they may not always be effective in complex waters potentially containing high concentrations of optically significant constituents. This paper presents new empirical formulas for estimating selected inherent optical properties of water from remote-sensing reflectance spectra Rrs(λ), derived, among other things, for waters with high concentrations of dissolved and suspended substances. These formulas include one for estimating the backscattering coefficient bb(620) directly from the magnitude of Rrs in the red part of the spectrum, and another for estimating the absorption coefficient a(440) from the hue angle α. The latter quantity represents the water’s colour as it might be perceived by the human eye (trichromatic colour vision); it is easily calculated from the shape of the Rrs spectrum. These new formulas are based on a combined dataset. Most of the data were obtained in the specific, optically complex environment of the Baltic Sea. Additional data, taken from the NASA bio-Optical Marine Algorithm Dataset (NOMAD) and representing various regions of the global oceans, were used to widen the potential applicability of the new formulas. We indicate the reasons why these simple empirical relationships can be derived and compare them with the results of straightforward modelling; possible applications are also described. We present, among other things, an example of a simple semi-analytical algorithm using both new empirical formulas. This algorithm is a modified version of the well-known quasi-analytical algorithm (QAA), and it can improve the results obtained in optically complex waters. This algorithm allows one to estimate the full spectra of the backscattering and absorption coefficients, without the need for any additional a priori assumptions regarding the spectral shape of absorption by dissolved and suspended seawater constituents.

Diverse Cell Types, Circuits, and Mechanisms for Color Vision in the Vertebrate Retina.

Synaptic interactions to extract information about wavelength, and thus color, begin in the vertebrate retina with three classes of light-sensitive cells: rod photoreceptors at low light levels, multiple types of cone photoreceptors that vary in spectral sensitivity, and intrinsically photosensitive ganglion cells that contain the photopigment melanopsin. When isolated from its neighbors, a photoreceptor confounds photon flux with wavelength and so by itself provides no information about color. The retina has evolved elaborate color opponent circuitry for extracting wavelength information by comparing the activities of different photoreceptor types broadly tuned to different parts of the visible spectrum. We review studies concerning the circuit mechanisms mediating opponent interactions in a range of species, from tetrachromatic fish with diverse color opponent cell types to common dichromatic mammals where cone opponency is restricted to a subset of specialized circuits. Distinct among mammals, primates have reinvented trichromatic color vision using novel strategies to incorporate evolution of an additional photopigment gene into the foveal structure and circuitry that supports high-resolution vision. Color vision is absent at scotopic light levels when only rods are active, but rods interact with cone signals to influence color perception at mesopic light levels. Recent evidence suggests melanopsin-mediated signals, which have been identified as a substrate for setting circadian rhythms, may also influence color perception. We consider circuits that may mediate these interactions. While cone opponency is a relatively simple neural computation, it has been implemented in vertebrates by diverse neural mechanisms that are not yet fully understood.

Melanopsin and Cone Photoreceptor Inputs to the Afferent Pupil Light Response.

Background: Retinal photoreceptors provide the main stage in the mammalian eye for regulating the retinal illumination through changes in pupil diameter, with a small population of melanopsin-expressing intrinsically photosensitive retinal ganglion cells (ipRGCs) forming the primary afferent pathway for this response. The purpose of this study is to determine how melanopsin interacts with the three cone photoreceptor classes in the human eye to modulate the light-adapted pupil response.Methods: We investigated the independent and combined contributions of the inner and outer retinal photoreceptor inputs to the afferent pupil pathway in participants with trichromatic color vision using a method to independently control the excitations of ipRGCs, cones and rods in the retina.Results: We show that melanopsin-directed stimuli cause a transient pupil constriction generated by cones in the shadow of retinal blood vessels; desensitizing these penumbral cone signals uncovers a signature melanopsin pupil response that includes a longer latency (292 ms) and slower time (4.1x) and velocity (7.7x) to constriction than for cone-directed stimuli, and which remains sustained post-stimulus offset. Compared to melanopsin-mediated pupil responses, the cone photoreceptor-initiated pupil responses are more transient with faster constriction latencies, higher velocities and a secondary constriction at light offset. The combined pupil responses reveal that melanopsin signals are additive with the cone signals.Conclusions: The visual system uses the L–, M–, and S–cone photoreceptor inputs to the afferent pupil pathway to accomplish the tonic modulations of pupil size to changes in image contrast. The inner retinal melanopsin-expressing ipRGCs mediate the longer-term, sustained pupil constriction to set the light-adapted pupil diameter during extended light exposures.

Melanopsin driven enhancement of cone-mediated visual processing

The precise control of visual sensitivity to variations in external lighting is critical for optimising human visual function in a changing environment. In photopic illumination, the cone photoreceptors and their post-receptoral pathways have a primary role in regulating these adjustments; it is not fully understood how a small population of melanopsin-expressing, intrinsically photosensitive retinal ganglion cells (ipRGCs) can interact with the three cone photoreceptor classes to modulate human visual function. Here we investigated interactions between these inner and outer retinal photoreceptor classes in participants with trichromatic colour vision under conditions that independently controlled the excitations of ipRGCs, cones and rods in the retina. In the peripheral retina, we show that interaction between melanopsin- and cone-directed signals affect conscious, image-forming vision. The interaction patterns are inconsistent with any potential effects arising from artefacts due to open-field and penumbral-cone contrasts, or rod intrusion. For cone signals mediated via each of the three primary visual pathways, melanopsin activation enhances contrast sensitivity. The contrast response functions indicate this enhancement is a more generalised facilitation effect that onsets at ∼9% melanopsin contrast. The implication for human vision is that the contrast sensitivity of cone-mediated visual processes can be modulated by the level of melanopsin excitation in the stimulus light.

Social Perception of Facial Color Appearance for Human Trichromatic Versus Dichromatic Color Vision.

Typical human color vision is trichromatic, on the basis that we have three distinct classes of photoreceptors. A recent evolutionary account posits that trichromacy facilitates detecting subtle skin color changes to better distinguish important social states related to proceptivity, health, and emotion in others. Across two experiments, we manipulated the facial color appearance of images consistent with a skin blood perfusion response and asked participants to evaluate the perceived attractiveness, health, and anger of the face (trichromatic condition). We additionally simulated what these faces would look like for three dichromatic conditions (protanopia, deuteranopia, tritanopia). The results demonstrated that flushed (relative to baseline) faces were perceived as more attractive, healthy, and angry in the trichromatic and tritanopia conditions, but not in the protanopia and deuteranopia conditions. The results provide empirical support for the social perception account of trichromatic color vision evolution and lead to systematic predictions of social perception based on ecological social perception theory.

Less is more: lemurs (Eulemur spp.) may benefit from loss of trichromatic vision

Vertebrate color vision is an ideal system for studying the gains and losses of genetic variation across lineages and impacts on behavior. Among placental mammals, trichromatic vision is unique to primates and is argued to be adaptive for foraging on reddish food. However, trichromacy is variably present in lemurs, including species within the cathemeral genus Eulemur, due to inter- and intra-specific variation in X-linked opsin genes. Although this variation could result from genetic drift, it could also reflect ecological adaptation. To understand ecological contributions to color vision variation, we examined cone opsin genes of 11 Eulemur species. We found that only E. flavifrons and E. macaco have polymorphic trichromacy. Most dichromatic species have an “M” (green-shifted) opsin; uniquely, one species (E. rubriventer) has dichromacy based on an “L” (red-shifted) opsin. This latter result appears to represent loss of polymorphic trichromacy from a dichromatic (M opsin) or polymorphic Eulemur ancestor. To address potential ecological explanations for opsin variation, we studied the dietary behavior of wild E. rubriventer and collected reflectance spectra from plant species consumed. Visual models suggest that trichromacy should provide an advantage for detecting reddish foods; however, luminance contrasts were greatest for dichromats with the L opsin. As E. rubriventer are often active in low-light rainforest conditions, luminance cues may be relatively important, which could favor the L opsin, while also leading to relaxed selection on, or selection against, trichromacy. The presence of different opsin alleles across Eulemur species could represent adaptations related to diet, activity pattern, or habitat. Loss of genetic variation, often thought to be maladaptive, can occur through natural selection. Among primates, some species have trichromatic color vision, the ability to distinguish reddish and greenish hues; others are red-green colorblind (dichromatic). We examined adaptive explanations for color vision differences by studying cone opsin genes and behavior in wild lemurs (Eulemur)—a genus that is active both day and night. We found that color vision is variable in Eulemur species, and full trichromatic vision was likely lost in at least one lineage. Foraging ecology of dichromatic Eulemur rubriventer indicates that trichromatic vision should be advantageous for foraging on reddish foods, but brightness cues are more salient to this species’ vision. We suggest brightness may be more important than color to this species, particularly at night, and loss of trichromacy could be adaptive in some lemurs.

A β-carotene-binding protein carrying a red pigment regulates body-color transition between green and black in locusts

Changes of body color have important effects for animals in adapting to variable environments. The migratory locust exhibits body color polyphenism between solitary and gregarious individuals, with the former displaying a uniform green coloration and the latter having a prominent pattern of black dorsal and brown ventral surface. However, the molecular mechanism underlying the density-dependent body color changes of conspecific locusts remain largely unknown. Here, we found that upregulation of β-carotene-binding protein promotes the accumulation of red pigment, which added to the green color palette present in solitary locusts changes it from green to black, and that downregulation of this protein led to the reverse, changing the color of gregarious locusts from black to green. Our results provide insight that color changes of locusts are dependent on variation in the red β-carotene pigment binding to βCBP. This finding of animal coloration corresponds with trichromatic theory of color vision.

No Effect of Red on Personality Trait Self-Ratings: Testing for Effects of Font Color

Unlike most other mammals, humans are trichromats and have the ability to perceive the color red. An explanation for the evolution of humans’ trichromatic color vision is that it offers humans the advantage to detect ripe fruit. Apart from this explanation, psychological theories have proposed that color, especially the color red, conveys information that affects psychological functioning, but results have been mixed. Whereas studies have extensively tested effects of red on performance measures, it is unclear whether this effect generalizes to self-ratings, one of the most frequently used methods in psychological research. In line with theory and empirical evidence, we argue that exposure to red can lead to distorted responses in self-ratings on the basis of the same underlying mechanism that affects results on performance measures. We varied the font color (hue values) of self-ratings in two online studies. In a first exploratory study, we found an effect of font color on personality trait self-ratings (N = 145). We attempted to rigorously replicate this finding in a larger sample (N = 1,007) but did not detect any effect. The findings underline the importance of rigorous research on effects of color on psychological functioning and call into question the proposition that red has ubiquitous effects.

Color statistics of objects, and color tuning of object cortex in macaque monkey

We hypothesized that the parts of scenes identified by human observers as “objects” show distinct color properties from backgrounds, and that the brain uses this information towards object recognition. To test this hypothesis, we examined the color statistics of naturally and artificially colored objects and backgrounds in a database of over 20,000 images annotated with object labels. Objects tended to be warmer colored (L-cone response > M-cone response) and more saturated compared to backgrounds. That the distinguishing chromatic property of objects was defined mostly by the L-M post-receptoral mechanism, rather than the S mechanism, is consistent with the idea that trichromatic color vision evolved in response to a selective pressure to identify objects. We also show that classifiers trained using only color information could distinguish animate versus inanimate objects, and at a performance level that was comparable to classification using shape features. Animate/inanimate is considered a fundamental superordinate category distinction, previously thought to be computed by the brain using only shape information. Our results show that color could contribute to animate/inanimate, and likely other, object-category assignments. Finally, color-tuning measured in two macaque monkeys with functional magnetic resonance imaging (fMRI), and confirmed by fMRI-guided microelectrode recording, supports the idea that responsiveness to color reflects the global functional organization of inferior temporal cortex, the brain region implicated in object vision. More strongly in IT than in V1, colors associated with objects elicited higher responses than colors less often associated with objects.

Luminescence properties and energy transfer in Ce3+/Tb3+co–doped Y5Si3O12N oxynitride phosphors

A series of Y5Si3O12N:Ce3+ single-doped and Ce3+/Tb3+ co–doped phosphors, as potential candidates for application in tri–chromatic white LEDs, were successfully synthesized by a high temperature solid state reaction method. The addition of Li2CO3 flux (5 wt%) improved the purity of the resultant phosphors and increased their emission intensity. An intense, broad excitation band in the range of 200–400 nm with a peak at 358 nm was recorded, which suits the needs of UV LED chips. Under this excitation wavelength, intense blue emission, which peaked at 439 nm, was recorded in the Ce3+ single-doped phosphors, while two distinct luminescence bands were observed in the Ce3+/Tb3+ co–doped Y5Si3O12N phosphors: a blue one, which peaked at 439 nm and is ascribed to Ce3+ ions, and an efficient green one, which peaked at 549 nm and is assigned to 5D4→7F5 transitions in Tb3+ ions. A high luminescence quantum yield, of 85%, was achieved. The experimental results illustrate that efficient energy transfer can take place from the sensitizer Ce3+ ions to the activator Tb3+ ions in the Ce3+/Tb3+ co–doped Y5Si3O12N phosphors, through predominantly dipole–dipole interactions.

Design considerations for the enhancement of human color vision by breaking binocular redundancy

To see color, the human visual system combines the response of three types of cone cells in the retina—a compressive process that discards a significant amount of spectral information. Here, we present designs based on thin-film optical filters with the goal of enhancing human color vision by breaking its inherent binocular redundancy, providing different spectral content to each eye. We fabricated a set of optical filters that “splits” the response of the short-wavelength cone between the two eyes in individuals with typical trichromatic vision, simulating the presence of approximately four distinct cone types. Such an increase in the number of effective cone types can reduce the prevalence of metamers—pairs of distinct spectra that resolve to the same tristimulus values. This technique may result in an enhancement of spectral perception, with applications ranging from camouflage detection and anti-counterfeiting to new types of artwork and data visualization.

Understanding innate preferences of wild bee species: responses to wavelength-dependent selective excitation of blue and green photoreceptor types.

Bees have a trichromatic vision with ultraviolet, blue, and green photoreceptors in their compound eyes. While the three photoreceptor types comprise the 'color space' at the perceptual level, preferential excitation of one or two of the photoreceptor types has been shown to play an important role in innate color preferences of bumble bees. Bees have been shown to exhibit strong attraction to fluorescence emission exclusively in the blue spectral region. It is not known if emission exclusively in the green spectral region produces similar attraction. Here, we examined responses of wild bees to traps designed to selectively stimulate either the blue or the green photoreceptor using sunlight-induced fluorescence in the 420-480 or 510-540nm region, respectively. Additionally, we probed how subtle changes in the spectral characteristics of the traps affect the bee captures once a highly selective excitation of the blue photoreceptor is achieved. It was established that selective excitation of the green photoreceptor type was not attractive, in contrast to that of the blue photoreceptor type. However, once a highly selective excitation of the blue photoreceptor type (at ~ 400-480nm) was achieved, the wild bees favored strong excitation at 430-480nm over that in the 400-420nm region.

Spectral sensitivities of ants – a review

AbstractAnts constitute one of the most intriguing animal groups with their advanced social lifes, different life histories and sensory modalities, one of which is vision. Chemosensation dominates all other modalities in the accomplishment of different vital tasks, but vision, varying from total blindness in some species to a relatively well-developed vision providing ants the basis for visually-guided behaviors, is also of importance. Although studies on ant vision mainly focused on recognition of and guidance by landmark cues in artificial and/or natural conditions, spectral sensitivities of their compound eyes and ocelli were also disclosed, but to a lesser extent. In this review, we have tried to present current data on the spectral sensitivities of the different ant species tested so far and the different methodological approaches. The results, as well as the similarities and/or discrepancies of the methodologies applied, were compared. General tendencies in ants’ spectral sensitivities are presented in a comparative manner and the role of opsins and ant ocelli in their spectral sensitivity is discussed in addition to the sensitivity of ants to long wavelengths. Extraocular sensitivity was also shown in some ant species. The advantages and/or disadvantages of a dichromatic and trichromatic color vision system are discussed from an ecological perspective.

A Temporal White Noise Analysis for Extracting the Impulse Response Function of the Human Electroretinogram.

PurposeWe introduce a method for determining the impulse response function (IRF) of the ERG derived from responses to temporal white noise (TWN) stimuli.MethodsThis white noise ERG (wnERG) was recorded in participants with normal trichromatic vision to full-field (Ganzfeld) and 39.3° diameter focal stimuli at mesopic and photopic mean luminances and at different TWN contrasts. The IRF was obtained by cross-correlating the TWN stimulus with the wnERG.ResultsWe show that wnERG recordings are highly repeatable, with good signal-to-noise ratio, and do not lead to blink artifacts. The wnERG resembles a flash ERG waveform with an initial negativity (N1) followed by a positivity (P1), with amplitudes that are linearly related to stimulus contrast. These N1 and N1-P1 components showed commonalties in implicit times with the a- and b-waves of flash ERGs. There was a clear transition from rod- to cone-driven wnERGs at ∼1 photopic cd.m−2. We infer that oscillatory potentials found with the flash ERG, but not the wnERG, may reflect retinal nonlinearities due to the compression of energy into a short time period during a stimulus flash.ConclusionThe wnERG provides a new approach to study the physiology of the retina using a stimulation method with adaptation and contrast conditions similar to natural scenes to allow for independent variation of stimulus strength and mean luminance, which is not possible with the conventional flash ERG.Translational RelevanceThe white noise ERG methodology will be of benefit for clinical studies and animal models in the evaluation of hypotheses related to cellular redundancy to understand the effects of disease on specific visual pathways.

Rethinking the Origin of Primates by Reconstructing Their Diel Activity Patterns Using Genetics and Morphology

Phylogenetic inference typically invokes nocturnality as ancestral in primates; however, some recent studies posit that diurnality is. Here, through adaptive evolutionary analyses of phototransduction genes by using a variety of approaches (restricted branch/branch-site models and unrestricted branch-site-based models (BS-REL, BUSTED and RELAX)), our results consistently showed that ancestral primates were subjected to enhanced positive selection for bright-light vision and relatively weak selection for dim-light vision. These results suggest that ancestral primates were mainly diurnal with some crepuscularity and support diurnality as plesiomorphic from Euarchontoglires. Our analyses show relaxed selection on motion detection in ancestral primates, suggesting that ancestral primates decreased their emphasis on mobile prey (e.g., insects). However, within primates, the results show that ancestral Haplorrhini were likely nocturnal, suggesting that evolution of the retinal fovea occurred within ancestral primates rather than within haplorrhines as was previously hypothesized. Our findings offer a reassessment of the visual adaptation of ancestral primates. The evolution of the retinal fovea, trichromatic vision and orbital convergence in ancestral primates may have helped them to efficiently discriminate, target, and obtain edible fruits and/or leaves from a green foliage background instead of relying on mobile insect prey.

Possibility of trichromatic colouration of glass

Possibility of trichromatic colouration of glass

Experimental evidence that primate trichromacy is well suited for detecting primate social colour signals.

Primate trichromatic colour vision has been hypothesized to be well tuned for detecting variation in facial coloration, which could be due to selection on either signal wavelengths or the sensitivities of the photoreceptors themselves. We provide one of the first empirical tests of this idea by asking whether, when compared with other visual systems, the information obtained through primate trichromatic vision confers an improved ability to detect the changes in facial colour that female macaque monkeys exhibit when they are proceptive. We presented pairs of digital images of faces of the same monkey to human observers and asked them to select the proceptive face. We tested images that simulated what would be seen by common catarrhine trichromatic vision, two additional trichromatic conditions and three dichromatic conditions. Performance under conditions of common catarrhine trichromacy, and trichromacy with narrowly separated LM cone pigments (common in female platyrrhines), was better than for evenly spaced trichromacy or for any of the dichromatic conditions. These results suggest that primate trichromatic colour vision confers excellent ability to detect meaningful variation in primate face colour. This is consistent with the hypothesis that social information detection has acted on either primate signal spectral reflectance or photoreceptor spectral tuning, or both.

Colour vision requirements in visually demanding occupations.

Normal trichromatic colour vision (CV) is often required as a condition for employment in visually demanding occupations. If this requirement could be enforced using current, colour assessment tests, a significant percentage of subjects with anomalous, congenital trichromacy who can perform the suprathreshold, colour-related tasks encountered in many occupations with the same accuracy as normal trichromats would fail. These applicants would therefore be discriminated against unfairly. One solution to this problem is to produce minimum, justifiable CV requirements that are specific to each occupation. This has been done successfully for commercial aviation (i.e. the flight crew) and for Transport for London train drivers. An alternative approach is to make use of new findings and the statistical outcomes of past practices to produce graded, justifiable CV categories that can be enforced. To achieve this aim, we analysed colour assessment outcomes and quantified severity of CV loss in 1363 subjects. The severity of CV loss was measured in each subject and statistical, pass/fail outcomes established for each of the most commonly used, conventional colour assessment tests and protocols. This evidence and new findings that relate severity of loss to the effective use of colour signals in a number of tasks provide the basis for a new colour grading system based on six categories. A single colour assessment test is needed to establish the applicant's CV category which can range from 'supernormal', for the most stringent, colour-demanding tasks, to 'severe colour deficiency', when red/green CV is either absent or extremely weak.