Cone Photoreceptor Sensitivities and Unique Hue Chromatic Responses: Correlation and Causation Imply the Physiological Basis of Unique Hues

This paper relates major functions at the start and end of the color vision process. The process starts with three cone photoreceptors transducing light into electrical responses. Cone sensitivities were once expected to be Red Green Blue color matching functions (to mix colors) but microspectrometry proved otherwise: they instead peak in yellowish, greenish, and blueish hues. These physiological functions are an enigma, unmatched with any set of psychophysical (behavioral) functions. The end-result of the visual process is color sensation, whose essential percepts are unique (or pure) hues red, yellow, green, blue. Unique hues cannot be described by other hues, but can describe all other hues, e.g., that hue is reddish-blue. They are carried by four opponent chromatic response curves but the literature does not specify whether each curve represents a range of hues or only one hue (a unique) over its wavelength range. Here the latter is demonstrated, confirming that opponent chromatic responses define, and may be termed, unique hue chromatic responses. These psychophysical functions also are an enigma, unmatched with any physiological functions or basis. Here both enigmas are solved by demonstrating the three cone sensitivity curves and the three spectral chromatic response curves are almost identical sets (Pearson correlation coefficients r from 0.95–1.0) in peak wavelengths, curve shapes, math functions, and curve crossover wavelengths, though previously unrecognized due to presentation of curves in different formats, e.g., log, linear. (Red chromatic response curve is largely nonspectral and thus derives from two cones.) Close correlation combined with deterministic causation implies cones are the physiological basis of unique hues. This match of three physiological and three psychophysical functions is unique in color vision.     

A Full Color System for Quantitative Assessment of Histochemical and Immunohistochemical Staining Patterns

Quantitative measures of staining distributions are important to compare the presence and patterns of cells or macromolecules. Typically, achromatic thresholding systems are used to compare staining distributions. Achromatic video signals, however, lack sufficient resolution to identify and compare chromatic changes. The purpose of this study is to describe a full color system for analysis of chromatic staining distributions. The hardware system includes a Leitz Diaplan microscope, video camera, GVP videoboard and Amiga 3000 computer. Software was developed in “C” to partition the video signal into hue (H), saturation (S) and value (V). Also, percentage of stained area was determined. Kodak color filters were used to assess the accuracy and precision of the system. Craniofacial tissues were stained with varying concentrations of toluidine blue and primary anti-Brdll antibodies. HSV and the percentage of stained areas were determined and displayed low coefficients of error. HSV values also performed as expected for standard filters as well as cellular staining concentrations. This system is easily implemented and should be useful for comparing chromatic changes with any color resulting from histochemical or immunohistochemical procedures.     

Plumage colour in nestling blue tits: sexual dichromatism,condition dependence and genetic effects

Sexual-selection theory assumes that there are costs associated with ornamental plumage coloration. While pigment-based ornaments have repeatedly been shown to be condition dependent, this has been more difficult to demonstrate for structural colours. We present evidence for condition dependence of both types of plumage colour in nestling blue tits (Parus caeruleus). Using reflectance spectrometry, we show that blue tit nestlings are sexually dichromatic, with males having more chromatic (more 'saturated') and ultraviolet (UV)-shifted tail coloration and more chromatic yellow breast coloration. The sexual dimorphism in nestling tail coloration is qualitatively similar to that of chick-feeding adults from the same population. By contrast, the breast plumage of adult birds is not sexually dichromatic in terms of chroma. In nestlings, the chroma of both tail and breast feathers is positively associated with condition (body mass on day 14). The UV/blue hue of the tail feathers is influenced by paternally inherited genes, as indicated by a maternal half-sibling comparison. We conclude that the expression of both carotenoid-based and structural coloration seems to be condition dependent in blue tit nestlings, and that there are additional genetic effects on the hue of the UV/blue tail feathers. The signalling or other functions of sexual dichromatism in nestlings remain obscure. Our study shows that nestling blue tits are suitable model organisms for the study of ontogenetic costs and heritability of both carotenoid-based and structural colour in birds.     

Anisotropy in the chromatic channel: a horizontal-vertical effect

We compared the chromatic contrast thresholds of drifting (2Hz) red-green sine-wave gratings of horizontal, vertical, and two oblique orientations at three spatial frequencies (2, 4, 8 cpd). Luminance contrast thresholds for yellow-black gratings were also obtained. The classic oblique effect was found for high spatial frequency luminance and chromatic stimuli. For chromatic thresholds, a significant difference was found between the horizontal and vertical thresholds of all observers. One observer was retested with her head tilted 45 deg and demonstrated that the anisotropy was specific to retinal coordinates. These results give evidence for orientation selectivity in the chromatic channel which is at least partially independent of that in the luminance channel. We estimated the degree of lateral chromatic aberration in our observers' eyes and discuss the possible contribution of this aberration to the horizontal-vertical difference in the chromatic channel.     

Optimum probabilistic processing in colour perception. II. Colour vision as template matching.

A statistical approach to account for psychophysical phenomena in human colour vision is presented. The central visual processor is viewed as an optimum recognizer of stochastic patterns supplied by the periphery. The processor makes an optimum estimate of the spectral parameters of the stimulus, given the wavelength filter characteristics of the periphery, the stochastic nature of the information and an internal template to which the external stimulus is matched. The estimate is constrained in ways inferred from empirical phenomena. Subjective brightness of monochromatic stimuli and related constant brightness manifolds in the colour space constitute the constraint for brightness estimation. Results analogous and in accord with those of earlier line element theories are obtained. The Bezold-Brücke hue shift constitutes the basic constraint for hue estimation. The hue estimate involves interrelation between the fields in the experiment. Similarities and differences both in basic conceptions and results introduced by the template matching notions are discussed.     

紫茉莉开花过程中花色和花色素的变化规律

紫茉莉是我国广泛分布的庭院花卉之一,具有丰富的花色。但不同花色紫茉莉在开花过程中的花色变化规律及其呈色机制还不清楚。以紫红色、黄色和白色紫茉莉为研究对象,分别通过色差仪测定法和紫外-可见分光光度法测定了不同开花时期不同花色紫茉莉花色表型及各类色素含量,探讨了其花色和色素变化规律,揭示其呈色机制。结果表明,从花蕾期到盛开期,紫红色紫茉莉花冠由淡绿色转变为紫红色,明度L^*值和色相b^*值减小,而色相a^*值、色度C^*值和色度角h值增大,叶绿素含量逐渐下降,类胡萝卜素、花色素苷和总黄酮含量逐渐升高;黄色紫茉莉花冠由淡绿色转变为黄色,盛开期具有最高的色度C^*值、色相a^*值和b^*值,整个开花过程具有较稳定的叶绿素和总黄酮含量,同时具有较高的类胡萝卜素含量;白色紫茉莉花冠由淡绿色转变为白色,过渡期具有最高的明度L^*值、色度C^*值、色相a^*值和b^*值,整个开花过程花色素苷和总黄酮含量较低,但随着开花进程逐渐升高,而类胡萝卜素含量稳定,过渡期总叶绿素含量显著低于其他2个时期。可见,不同花色紫茉莉开花过程中花色变化规律存在差异,而其差异性与其相应的色素成分变化密切相关。     

Development of contrast sensitivity and acuity of the infant colour system

We have monitored the development of infant colour vision by measuring chromatic contrast sensitivity and acuity in eight young infants over a period of 6 months. Steady-state visual evoked potentials (VEPS) were recorded in response to both chromatic (red-green) and luminance (red-black or green-black) patterns that were reversed in contrast over time. For most infants, no response could be obtained to chromatic stimuli of any size or contrast before 5 weeks of age, although luminance stimuli of 20% contrast gave reliable responses at that age. When responses to chromatic stimuli first appeared, they could be obtained only with stimuli of very low spatial frequency, 20 times lower than the acuity for luminance stimuli. Both contrast sensitivity and acuity for chromatic stimuli increased steadily, more rapidly than for luminance stimuli. As the spectral selectivities of infant cones are similar to those of adults, the difference in rate of development of luminance and chromatic contrast sensitivity and acuity stimuli probably reflects neural development of the infant colour system.     

When Do Short-Wave Cones Signal Blue or Red? A Solution Introducing the Concept of Primary and Secondary Cone Outputs

A recent paper by Oh and Sakata investigates the “incompletely solved mystery” of how the three cone responses map onto perceived hue, and particularly the S cone’s well-known problematic contribution to blueness and redness. Citing previous workers, they argue the twentieth century traditional multistage model does not satisfactorily account for color appearance. In their experiment, increasing S cone excitation with shortening wavelength from about 480–460 nm increased perceived blueness up to the unique Blue point at 470 nm, when (a) it began decreasing and (b) redness perception began increasing. The authors asked, What mechanism can be responsible for such functions? I demonstrate a solution. First, it is shown the problem does not lie in the traditional opponent color chromatic responses yellow-blue, red-green (y-b, r-g, which accurately predict the above functions), but in the traditional multistage model of mapping cone responses to chromatic response functions. Arguably, this is due to the S cone’s hypothetically signaling both blueness and redness by the same mechanism rather than by different, independent, mechanisms. Hence a new distinction or mechanism is proposed for a more accurate model, that introduces the new terms primary and secondary cone outputs. However, this distinction requires that the cones S, M, L each directly produce one of the three spectral chromatic responses b, g, y. Such a model was recently published, based on extremely high correlation of SML cone responsivities with the three spectral (bgy) chromatic responses. This model encodes the former directly onto the latter one-to-one as cone primary outputs, whilst S and L cones have a further or secondary function where each produces one of the two spectral lobes of r chromatic response. The proposed distinction between primary and secondary cone outputs is a new concept and useful tool in detailing cone outputs to chromatic channels, and provides a solution to the above “incompletely solved mystery.” Thus the S cone has a primary output producing the total b chromatic response and a secondary output that shares with the L cone the production of r chromatic response, thus aligning with Oh and Sokata’s results. The model similarly maps L cone to yellowness as primary output and to redness as secondary output.     

The effects of commonly used anaesthetics on colour measurements across body regions in the poeciliid fish,Girardinus metallicus

The effects of common anaesthetics on the hue, saturation and brightness measurements of the poeciliid fish Girardinus metallicus were investigated in two experiments. For both experiments the coloration of four body regions was measured from digital images of the same males obtained under three conditions: (1) control (in a water-filled chamber); (2) anaesthetised with MS-222; and (3) anaesthetised with eugenol (clove oil). In experiment 1 anaesthetised fish were photographed out of water. In experiment 2 all photographs were taken in a water-filled chamber. Anaesthetics altered coloration in both experiments. In the more methodologically consistent experiment 2 we found significantly different hue, increased saturation and decreased brightness in anaesthetic v. control conditions, consistent with darkening caused by the anaesthetics. The body regions differed in coloration consistent with countershading but did not differentially change in response to anaesthesia. These findings suggest that photographing fish in a water-filled chamber without anaesthetic is preferable for obtaining digital images for colour analysis and that multiple body regions of fish should be measured when assessing coloration patterns meaningful in behavioural contexts, to account for the gradients caused by countershading. We are encouraged that some researchers employ such methods already and caution against using anaesthetics except when absolutely necessary for immobilisation.     

Alouatta Trichromatic Color Vision: Cone Spectra and Physiological Responses Studied with Microspectrophotometry and Single Unit Retinal Electrophysiology

The howler monkeys (Alouatta sp.) are the only New World primates to exhibit routine trichromacy. Both males and females have three cone photopigments. However, in contrast to Old World monkeys, Alouatta has a locus control region upstream of each opsin gene on the X-chromosome and this might influence the retinal organization underlying its color vision. Post-mortem microspectrophotometry (MSP) was performed on the retinae of two male Alouatta to obtain rod and cone spectral sensitivities. The MSP data were consistent with only a single opsin being expressed in each cone and electrophysiological data were consistent with this primate expressing full trichromacy. To study the physiological organization of the retina underlying Alouatta trichromacy, we recorded from retinal ganglion cells of the same animals used for MSP measurements with a variety of achromatic and chromatic stimulus protocols. We found MC cells and PC cells in the Alouatta retina with similar properties to those previously found in the retina of other trichromatic primates. MC cells showed strong phasic responses to luminance changes and little response to chromatic pulses. PC cells showed strong tonic response to chromatic changes and small tonic response to luminance changes. Responses to other stimulus protocols (flicker photometry; changing the relative phase of red and green modulated lights; temporal modulation transfer functions) were also similar to those recorded in other trichromatic primates. MC cells also showed a pronounced frequency double response to chromatic modulation, and with luminance modulation response saturation accompanied by a phase advance between 10–20 Hz, characteristic of a contrast gain mechanism. This indicates a very similar retinal organization to Old-World monkeys. Cone-specific opsin expression in the presence of a locus control region for each opsin may call into question the hypothesis that this region exclusively controls opsin expression.     

The Blue Coloration of the Common Surgeonfish, Paracanthurus hepatus-II. Color Revelation and Color Changes

Measurements of spectral reflectance from the sky-blue portion of skin from the common surgeonfish, Paracanthurus hepatus, showed a relatively steep peak at around 490 nm. We consider that a multilayer thin-film interference phenomenon of the non-ideal type, which occurs in stacks of very thin light-reflecting platelets in iridophores of that region, is primarily responsible for the revelation of that hue. The structural organization of the iridophore closely resembles that of bluish damselfish species, although one difference is the presence of iridophores in a monolayer in the damselfish compared to the double layer of iridophores in the uppermost part of the dermis of surgeonfish. If compared with the vivid cobalt blue tone of the damselfish, the purity of the blue hue of the surgeonfish is rather low. This may be ascribable mainly to the double layer of iridophores in the latter since incident lightrays are complicatedly reflected and scattered in the strata. The dark-blue hue of the characteristic scissors-shaped pattern on the trunk of surgeonfish is mainly due to the dense population of melanophores, because iridophores are only present there in a scattered fashion. Photographic and spectral reflectance studies in vivo, as well as photomicrographic, photo-electric, and spectrometric examinations of the state of chromatophores in skin specimens in vitro, indicate that both melanophores and iridophores are motile. Physiological analyses disclosed that melanophores are under the control of the sympathetic nervous and the endocrine systems, while iridophores are regulated mainly by nerves. The body color of surgeonfish shows circadian changes, and becomes paler at night; this effect may be mediated by the pineal hormone, melatonin, which aggregates pigment in melanophores.     

Separate colour-opponent mechanisms underlie the detection and discrimination of moving chromatic targets.

Current opinion holds that human colour vision is mediated primarily via a colour-opponent pathway that carries information about both wavelength and luminance contrast (type I). However, some authors argue that chromatic sensitivity may be limited by a different geniculostriate pathway, which carries information about wavelength alone (type II). We provide psychophysical evidence that both pathways may contribute to the perception of moving, chromatic targets in humans, depending on the nature of the visual discrimination. In experiment 1, we show that adaptation to drifting, red-green stimuli causes reductions in contrast sensitivity for both the detection and direction discrimination of moving chromatic targets. Importantly, the effects of adaptation are not directionally specific. In experiment 2, we show that adaptation to luminance gratings results in reduced sensitivity for the direction discrimination, but not the detection of moving chromatic targets. We suggest that sensitivity for the direction discrimination of chromatic targets is limited by a colour-opponent pathway that also conveys luminance-contrast information, whereas the detection of such targets is limited by a pathway with access to colour information alone. The properties of these pathways are consistent with the known properties of type-I and type-II neurons of the primate parvocellular lateral geniculate nucleus and their cortical projections. These findings may explain the known differences between detection and direction discrimination thresholds for chromatic targets moving at low to moderate velocities.     

Why are fruits colorful? The relative importance of achromatic and chromatic contrasts for detection by birds

The colors of fruits and flowers are traditionally viewed as an adaptation to increase the detectability of plant organs to animal vectors. The detectability of visual signals increases with increasing contrasts between target and background. Contrasts consist of a chromatic aspect (color) and an achromatic aspect (light intensity), which are perceived separately by animals. To evaluate the relative importance of fruits’ chromatic and achromatic contrasts for the detection by avian fruit consumers we conducted an experiment with artificial fruits of four different colors in a tropical forest. We displayed the fruits against two different backgrounds, an artificial background and a natural one, because they differed in achromatic properties. We found no effect of the type of background on fruit detection rates. Detection rates differed for the four fruit colors. The probability of detection was explained by the chromatic contrast between fruits and their background, not by the achromatic contrasts. We suggest that birds attend primarily to chromatic contrast probably because these are more reliably detected under variable light conditions. Consistent with this hypothesis, we found habitat-specific differences in the conspicuousness of natural fruit colors in the study area. Fruits of understory species that are subjected to the variable light conditions within a forest displayed higher chromatic contrasts than species growing in the open restinga forest with constant bright illumination. There was no such difference for achromatic contrasts. In sum, we suggest that fruit colors differ between habitats because fruit colors that have strong chromatic contrasts against background can increase plants’ reproductive success, particularly under variable light conditions.     

Texture segregation in chromatic element-arrangement patterns.

An element-arrangement pattern is composed of two types of elements arranged differently in different regions of a pattern. Rapid texture segregation depends on spontaneously discriminating the difference in the arrangement of the elements. Five experiments investigated the perceived segregation of patterns composed of two types of squares arranged in vertical stripes in the top and bottom regions and in a checkerboard arrangement in the middle region. The squares were either equal in luminance and different in hue or equal in hue and different in luminance. The rated similarities of the two hues in a pattern failed to predict perceived segregation. For a given background luminance, the perceived segregation was predicted by the square-root of the sum of the squares of the differences in the outputs of the L - M + S and L + M - S opponent channels, where L, M, and S were the cone contrasts of the long-, medium-, and short-wavelength receptors. The perceived similarity of the two hues in a pattern was not affected by the background luminance but was a function of cone excitations instead. For patterns differing in hue and equal in luminance, perceived segregation was an inverse function of the background luminance. A white background decreased the perceived segregation, but a black background did not. The effect of background luminance was not on the discrimination of the individual hues. The two hues making up a texture pattern were clearly distinguishable on a white background. A white background interfered with the discrimination of the vertical and diagonal columns of squares that distinguished the texture regions. For patterns differing in luminance and equal in hue, black and white backgrounds decreased the perceived segregation. The results indicate that adapting to an achromatic luminance distant from the luminance of the squares increased the Weber threshold for discriminating luminance differences, but did not increase the Weber threshold for discriminating hue differences. The experiments also revealed that luminance was the primary factor affecting perceived segregation and that perceived brightness is secondary. The results are consistent with the hypothesis that perceived segregation in element-arrangement patterns is primarily a function of the differences in the outputs of relatively early filtering mechanisms that encode pattern differences prior to the specification of the element shapes and their properties.     

Chromatophore activity during natural pattern expression by the squid Sepioteuthis lessoniana: contributions of miniature oscillation

Squid can rapidly change the chromatic patterns on their body. The patterns are created by the expansion and retraction of chromatophores. The chromatophore consists of a central pigment-containing cell surrounded by radial muscles that are controlled by motor neurons located in the central nervous system (CNS). In this study we used semi-intact squid (Sepioteuthis lessoniana) displaying centrally controlled natural patterns to analyze spatial and temporal activities of chromatophores located on the dorsal mantle skin. We found that chromatophores oscillated with miniature expansions/retractions at various frequencies, even when the chromatic patterns appear macroscopically stable. The frequencies of this miniature oscillation differed between "feature" and "background" areas of chromatic patterns. Higher frequencies occurred in feature areas, whereas lower frequencies were detected in background areas. We also observed synchronization of the oscillation during chromatic pattern expression. The expansion size of chromatophores oscillating at high frequency correlated with the number of synchronized chromatophores but not the oscillation frequency. Miniature oscillations were not observed in denervated chromatophores. These results suggest that miniature oscillations of chromatophores are driven by motor neuronal activities in the CNS and that frequency and synchrony of this oscillation determine the chromatic pattern and the expansion size, respectively.     

The butterfly Papilio xuthus detects visual motion using chromatic contrast

Many insects’ motion vision is achromatic and thus dependent on brightness rather than on colour contrast. We investigate whether this is true of the butterfly Papilio xuthus, an animal noted for its complex retinal organization, by measuring head movements of restrained animals in response to moving two-colour patterns. Responses were never eliminated across a range of relative colour intensities, indicating that motion can be detected through chromatic contrast in the absence of luminance contrast. Furthermore, we identify an interaction between colour and contrast polarity in sensitivity to achromatic patterns, suggesting that ON and OFF contrasts are processed by two channels with different spectral sensitivities. We propose a model of the motion detection process in the retina/lamina based on these observations.     

Discrimination of coloured patterns by honeybees through chromatic and achromatic cues

We investigated pattern discrimination by worker honeybees, Apis mellifera, focusing on the roles of spectral cues and the angular size of patterns. Free-flying bees were trained to discriminate concentric patterns in a Y-maze. The rewarded pattern could be composed of either a cyan and a yellow colour, which presented both different chromatic and achromatic L-receptor contrast, or an orange and a blue colour, which presented different chromatic cues, but the same L-receptor contrast. The non-rewarded alternative was either a single-coloured disc with the colour of the central disc or the surrounding ring of the pattern, a checkerboard pattern with non-resolvable squares, the reversed pattern, or the elements of the training pattern (disc or ring alone). Bees resolved and learned both colour elements in the rewarded patterns and their spatial properties. When the patterns subtended large visual angles, this discrimination used chromatic cues only. Patterns with yellow or orange central discs were generalised toward the yellow and orange colours, respectively. When the patterns subtended a visual angle close to the detection limit and L-receptor contrast was mediating discrimination, pattern perception was reduced: bees perceived only the pattern element with higher contrast.     

Discrimination of coloured stimuli by honeybees: alternative use of achromatic and chromatic signals

Using a Y-maze experimental set-up, honeybees Apis mellifera were trained to a coloured disc presented against an achromatic background. In subsequent tests they were given a choice between the trained disc and an alternative disc that differed either in its chromatic properties, or in the amount of achromatic green contrast that it produced against the background. Tests were conducted in two experimental situations: one in which discs subtended a visual angle of 30° (as viewed by the bee at the decision point in the Y-maze), and another in which the angle was 6.5° or 5° (depending on the experiment). At the visual angle of 30°, the bees' choice behaviour was governed by the differences in chromatic properties, and not by the differences in the amount of green contrast. With the 6.5°- and 5°-discs, on the other hand, it was governed by the differences in the amount of green contrast, and not by the differences in chromatic properties. Consequently, in the present discrimination task, bees use either chromatic or achromatic cues, depending on the visual angle subtended by the stimuli at the eye. Results of a further experiment, in which the trained disc was tested against discs that produced various amounts of green contrast, confirm the above conclusion and show, in addition, that bees learn the green-contrast difference between a trained and a non-rewarded alternative. Accepted: 25 October 1996     

Chromatic Patterns of the Hermit Crab Calcinus tubularis Related to the Occupied Shell

Crustacea are known to develop different chromatic patterns due to many factors. Regarding decapods, chromatism was mainly studied in crabs, while very little is known about chromatic patterns in hermit crabs. Calcinus tubularis is a typical infralittoral rocky bottom hermit crab, studied for different aspects of its biology except chromatic variations. This paper aims at describing the different colour morphologies of C. tubularis, discussing hypothesis of why they develop, and testing if in nature the crab prefers a shell with a chromatic pattern similar to that of its body. One hundred and forty crabs were observed and filmed in the laboratory. They were subdivided into two groups, according to their chromatic pattern: 1) light and 2) dark crabs; the shells they occupied were also subdivided into the two groups of 1) light and 2) dark shells on the basis of the epibionts encrusting them. Observations of 129 crabs suggest that the colour depends neither on depth nor on size, intermoult period, diet, reproductive period but it might be connected to genetic factors and might help crab to camouflage. Camouflage is suggested by the fact that 79.3% of the total examined specimens occupy shells with a chromatic pattern resembling that of the crabs themselves. This phenomenon is significantly more recurrent in females than in males and could help the crabs to be cryptic, first with the occupied shell and secondly with the habitat (rocks encrusted by photophylous algae).