Color constancy in goldfish: the limits

Color constancy was investigated in behavioral training experiments on colors ranging from blue to yellow, located in the color space close to Planck's locus representing the main changes in natural skylight. Two individual goldfish were trained to peck at a test field of medium hue out of a series of 13-15 yellowish and bluish test fields presented simultaneously on a black background. During training the tank in which the fish were swimming freely was illuminated with white light. Correct choices were rewarded with food. During the tests differently saturated yellow or blue illumination was used. The degree of color constancy was inferred from the choice behavior under these illuminations. Perfect color constancy was found up to a certain degree of saturation of the colored light. Beyond this level test fields other than the training test field were chosen, indicating imperfect color constancy. Color constancy was quantified by applying color metrics on the basis of the goldfish cone sensitivity functions.     

Physiological mechanisms of color constancy

Color constancy is the term given to the ability to recognize the color of objects correctly under different conditions of illumination. For this purpose the visual system must determine the character of the illumination, introduce a correction for it into the spectal composition of the light received from the object, and hence recreate the true color of its surface. Behavioral experiments on fish showed that they possess constant color vision of objects. Electrophysiological experiments on ganglion cells of the color type showed that the simplest mechanisms of correction for illumination are found at the retinal level. An investigation of model algorithms providing for color constancy showed thatthe presence of color vision makes it much easier to recognize the three-dimensional form of objects. This fact compels a reexamination of established views regarding the place and role of color vision in functions of the animal visual system as a whole.Institute for Problems in Information Transmission, Academy of Sciences of the USSR, Moscow. Translated from Neirofiziologiya, Vol. 7, No. 1, pp. 21–26, January–February, 1975.     

Experience in early infancy is indispensable for color perception

Early visual experience is indispensable to shape the maturation of cortical circuits during development. Monocular deprivation in infancy, for instance, leads to an irreversible reduction of visually driven activity in the visual cortex through the deprived eye and a loss of binocular depth perception. It was tested whether or not early experience is also necessary for color perception. Infant monkeys were reared for nearly a year in a separate room where the illumination came from only monochromatic lights. After extensive training, they were able to perform color matching. But, their judgment of color similarity was quite different from that of normal animals. Furthermore, they had severe deficits in color constancy; their color vision was very much wavelength dominated, so they could not compensate for the changes in wavelength composition. These results indicate that early visual experience is also indispensable for normal color perception.     

Mathematical biophysics of color vision: III. Color constancy

A mechanism is presented which can account for certain aspects of the phenomena of color constancy. The mechanism involves interaction between a given region and the remaining field. Each region is represented by a color center having the structure previously introduced (Landahl, 1952,Bull. Math. Biophysics,14, 317–25) to account for a number of phenomena of color vision. The trichromatic, symmetric mechanism is introduced for simplicity. The interaction is such that collaterals from each of the primaries representing the background send elements to each of the centers corresponding to the primaries representing the spot. However, the collaterals impinging upon unlike centers are excitatory while the collaterals impinging on like centers, corresponding to the same primary colors, are inhibitory. With proper choice of coefficients, the result is that for small changes in illumination, the resulting apparent color is unchanged. However, for greater changes in the color of the illumination, there results a distortion of the apparent color. A number of examples are illustrated numerically. This research was supported in whole or in part by the U. S. Air Force under Contract AF 49(638)-414 monitored by the Air Force Office of Scientific Research.     

Rethinking Colour Constancy

Colour constancy needs to be reconsidered in light of the limits imposed by metamer mismatching. Metamer mismatching refers to the fact that two objects reflecting metameric light under one illumination may reflect non-metameric light under a second; so two objects appearing as having the same colour under one illuminant can appear as having different colours under a second. Yet since Helmholtz, object colour has generally been believed to remain relatively constant. The deviations from colour constancy registered in experiments are usually thought to be small enough that they do not contradict the notion of colour constancy. However, it is important to determine how the deviations from colour constancy relate to the limits metamer mismatching imposes on constancy. Hence, we calculated metamer mismatching’s effect for the 20 Munsell papers and 8 pairs of illuminants employed in the colour constancy study by Logvinenko and Tokunaga and found it to be so extensive that the two notions—metamer mismatching and colour constancy—must be mutually exclusive. In particular, the notion of colour constancy leads to some paradoxical phenomena such as the possibility of 20 objects having the same colour under chromatic light dispersing into a hue circle of colours under neutral light. Thus, colour constancy refers to a phenomenon, which because of metamer mismatching, simply cannot exist. Moreover, it obscures the really important visual phenomenon; namely, the alteration of object colours induced by illumination change. We show that colour is not an independent, intrinsic attribute of an object, but rather an attribute of an object/light pair, and then define a concept of material colour in terms of equivalence classes of such object/light pairs. We suggest that studying the shift in material colour under a change in illuminant will be more fruitful than pursuing colour constancy’s false premise that colour is an intrinsic attribute of an object.     

Colour constancy in insects

Colour constancy is the perceptual phenomenon that the colour of an object appears largely unchanged, even if the spectral composition of the illuminating light changes. Colour constancy has been found in all insect species so far tested. Especially the pollinating insects offer a remarkable opportunity to study the ecological significance of colour constancy since they spend much of their adult lives identifying and choosing between colour targets (flowers) under continuously changing ambient lighting conditions. In bees, whose colour vision is best studied among the insects, the compensation provided by colour constancy is only partial and its efficiency depends on the area of colour space. There is no evidence for complete ‘discounting’ of the illuminant in bees, and the spectral composition of the light can itself be used as adaptive information. In patchy illumination, bees adjust their spatial foraging to minimise transitions between variously illuminated zones. Modelling allows the quantification of the adaptive benefits of various colour constancy mechanisms in the economy of nature. We also discuss the neural mechanisms and cognitive operations that might underpin colour constancy in insects.     

Color and polarization vision in foraging Papilio

This paper gives an overview of behavioral studies on the color and polarization vision of the Japanese yellow swallowtail butterfly, Papilio xuthus. We focus on indoor experiments on foraging individuals. Butterflies trained to visit a disk of certain color correctly select that color among various other colors and/or shades of gray. Correct selection persists under colored illumination, but is systematically shifted by background colors, indicating color constancy and simultaneous color contrast. While their eyes contain six classes of spectral receptors, their wavelength discrimination performance indicates that their color vision is tetrachromatic. P. xuthus innately prefers brighter targets, but can be trained to select dimmer ones under certain conditions. Butterflies trained to a dark red stimulus select an orange disk presented on a bright gray background over one on dark gray. The former probably appears darker to them, indicating brightness contrast. P. xuthus has a strong innate preference for vertically polarized light, but the selection of polarized light changes depending on the intensity of simultaneously presented unpolarized light. Discrimination of polarization also depends on background intensity. Similarities between brightness and polarization vision suggest that P. xuthus perceive polarization angle as brightness, such that vertical polarization appears brighter than horizontal polarization.     

Local and relational judgements of surface colour: constancy indices and discrimination performance

Colour constancy is generally assumed to arise from a combination of perceptual constancy mechanisms operating to partially discount illumination changes and relational mechanisms involved in judging the colour relationships between object surfaces. Here we examined the characteristics of these mechanisms using a 'yes/no' task. Subjects judged whether a target colour patch embedded in an array of coloured patches (a) stayed the same across a simulated temporal illuminant change (local colour judgement), or (b) changed in a manner consistent with the illuminant change (relational colour judgement). The colour of the target patch remained constant in one-third of the trials, changed in accord with the illuminant shift in another third, and shifted partially with the illuminant change in the remaining third. We found that perceptual constancy was relatively weak and relational constancy strong, as assessed using a modified colour constancy index. Randomising the spatial positions of coloured patches across the illuminant change did not affect subjects' constancy indices. Application of signal detection analysis revealed some otherwise hidden effects. In the case of relational judgements, subjects adopted more conservative criteria (fewer true and false positives) with randomisation, maintaining a constant level of discrimination performance (d'). For local judgements, randomisation led to small increases in performance but no changes in criteria. We conclude that signal detection theory provides a useful tool to supplement conventional approaches to understanding colour constancy.     

Sensory, computational and cognitive components of human colour constancy

When the illumination on a scene changes, so do the visual signals elicited by that scene. In spite of these changes, the objects within a scene tend to remain constant in their apparent colour. We start this review by discussing the psychophysical procedures that have been used to quantify colour constancy. The transformation imposed on the visual signals by a change in illumination dictates what the visual system must 'undo' to achieve constancy. The problem is mathematically underdetermined, and can be solved only by exploiting regularities of the visual world. The last decade has seen a substantial increase in our knowledge of such regularities as technical advances have made it possible to make empirical measurements of large numbers of environmental scenes and illuminants. This review provides a taxonomy of models of human colour constancy based first on the assumptions they make about how the inverse transformation might be simplified, and second, on how the parameters of the inverse transformation might be set by elements of a complex scene. Candidate algorithms for human colour constancy are represented graphically and pictorially, and the availability and utility of an accurate estimate of the illuminant is discussed. Throughout this review, we consider both the information that is, in principle, available and empirical assessments of what information the visual system actually uses. In the final section we discuss where in our visual systems these computations might be implemented.     

On the effect of scene motion on color constancy

A series of experiments with human subjects have shown that color constancy improves when an object moves. It has been hypothesized that this effect is due to some kind of influence of high-level motion processing. We have built a computational model for color perception which replicates the results qualitatively which have been obtained with human subjects. We show that input from high-level motion processing is not required. In our model, the dependence is an effect of eye movement in combination with neural processing. Depending on the type of stimulus used, the eye either tracks the object or the background. When the object moves but is tracked by the observer, the background appears to move when considering the stimulus with respect to eye coordinates. Hence, the retinal input is different for the two conditions leading to a difference in color constancy performance.     

Necessary and sufficient conditions for Von Kries chromatic adaptation to give color constancy

Necessary and sufficient spectral conditions are presented for Von Kries chromatic adaptation to give color constancy. Von-Kries-invariant reflectance spectra are computed for illuminant spectral power distributions that are arbitrary linear combinations of the first three daylight phases. Experiments are suggested to test models of color constancy using computed spectra (either exact or approximate) within the illuminant-invariant framework.     

Broad spectral sensitivities in the honeybee's photoreceptors limit colour constancy

Colour constancy allows for visual systems to be view stimuli independent of changes in spectral illumination. Chromatic adaptation is likely to be an important mechanism in colour constancy and can be explained by use of the von Kries coefficient law. Chromatic adaptation is compared for the honeybee and three hypothetical visual systems. It is shown that the spectral breadth and asymmetry of photoreceptors in the honeybee may limit colour constancy. In particular, it is demonstrated that the absorption of short-wavelength radiation by the cis-band of chromophore is responsible for a poorer correction for bee colours rich in ultraviolet reflectance. The results are discussed in relation to theoretical considerations of von Kries colour constancy and the physiology of eye design in some other species for which colour constancy has been demonstrated. Accepted: 14 August 1999     

Flower constancy in the hoverflies Episyrphus balteatus (Degeer) and Syrphus ribesii (L.) (Syrphidae)

The causes and consequences of flower constancy have been thefocus of many studies, but almost all have examined the foragingbehavior of bumblebees, honeybees, or butterflies. We test whetherconstancy occurs in an overlooked group of pollinators, thesyrphid flies. Foraging sequences of wild flies of two species,Episyrphus balteatus and Syrphus ribesii were examined whenvisiting flowers in seminatural plant communities and in artificialarrays of two color morphs of Lobularia maritima planted ata range of frequencies. Both species exhibited marked floralconstancy when foraging in the mixed-plant community. Becauseall groups of pollinating insect so far examined exhibit constancyat least under some circumstances, we suggest that this is thepredominant strategy used by pollinators and that there is probablya common explanation. Neither syrphid species exhibited constancyto different color morphs within a plant species, in contrastto previously published studies of Hymenoptera foraging amongpolymorphic flowers, which all describe positive frequency-dependentselection. Possible explanations for this discrepancy are discussed.We argue that constancy in these syrphids is unlikely to resultfrom learning constraints on handling ability, currently themost widely accepted explanation for flower constancy, becausethey forage primarily for pollen which is easily located inmostflowers they visit.     

Color categorization and color constancy in a neural network model of V4

We develop a neural network model that instantiates color constancy and color categorization in a single unified framework. Previous models achieve similar effects but ignore important biological constraints. Color constancy in this model is achieved by a new application of the double opponent cells found in the blobs of the visual cortex. Color categorization emerges naturally, as a consequence of processing chromatic stimuli as vectors in a four-dimensional color space. A computer simulation of this model is subjected to the classic psychophysical tests that first uncovered these phenomena, and its response matches psychophysical results very closely.     

Biological significance of distinguishing between similar colours in spectrally variable illumination: bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>) as a case study

Individual bumblebees were trained to choose between rewarded target flowers and non-rewarded distractor flowers in a controlled illumination laboratory. Bees learnt to discriminate similar colours, but with smaller colour distances the frequency of errors increased. This indicates that pollen transfer might occur between flowers with similar colours, even if these colours are distinguishable. The effect of similar colours on reducing foraging accuracy of bees is evident for colour distances high above discrimination threshold, which explains previous field observations showing that bees do not exhibit complete flower constancy unless flower colour between species is distinct. Bees tested in spectrally different illumination conditions experienced a significant decrease in their ability to discriminate between similar colours. The extent to which this happens differs in different areas of colour space, which is consistent with a von Kries-type model of colour constancy. We find that it would be beneficial for plant species to have highly distinctive colour signals to overcome limitations on the bees performance in reliably judging differences between similar colours. An exception to this finding was flowers that varied in shape, in which case bees used this cue to compensate for inaccuracies of colour vision.     

Feedback from Horizontal Cells to Cones Mediates Color Induction and May Facilitate Color Constancy in Rainbow Trout

Color vision is most beneficial when the visual system is color constant and can correct the excitations of photoreceptors for differences in environmental irradiance. A phenomenon related to color constancy is color induction, where the color of an object shifts away from the color of its surroundings. These two phenomena depend on chromatic spatial integration, which was suggested to originate at the feedback synapse from horizontal cells (HC) to cones. However, the exact retinal site was never determined. Using the electroretinogram and compound action potential recordings, we estimated the spectral sensitivity of the photoresponse of cones, the output of cones, and the optic nerve in rainbow trout. Recordings were performed before and following pharmacological inhibition of HC-cone feedback, and were repeated under two colored backgrounds to estimate the efficiency of color induction. No color induction could be detected in the photoresponse of cones. However, the efficiency of color induction in the cone output and optic nerve was substantial, with the efficiency in the optic nerve being significantly higher than in the cone output. We found that the efficiency of color induction in the cone output and optic nerve decreased significantly with the inhibition of HC-cone feedback. Therefore, our findings suggest not only that color induction originates as a result of HC-cone feedback, but also that this effect of HC-cone feedback is further amplified at downstream retinal elements, possibly through feedback mechanisms at the inner plexiform layer. This study provides evidence for an important role of HC-cone feedback in mediating color induction, and therefore, likely also in mediating color constancy.     

Color vision in the comb frequency domain

In 1982, Horace Barlow considered the question of human trichromacy in the context of information theory: according to the Sampling Theorem, three types of receptors covering the visible spectrum (400-700 nm) might be sufficient to reconstruct the color signal. Although Barlow was led to reject the direct application of the Sampling Theorem to explain color dimensionality, the theoretical framework offers a fresh point of view for analyzing the color system in conjunction with the physical characteristics of natural color signals. This review aims to illustrate that if the strict mathematical reconstruction (as implied by the Sampling Theorem) is replaced by a pragmatic approximation of color signals, then trichromacy, with its subsequent opponent-color process, could be regarded as an optimization of color constancy abilities in the spectral environment of primates. Higher dimension systems (tetrachromacy) found in other species can also serve the purpose of color constancy optimization in environments where color signals exhibit a finer spectral structure.     

Color perception and the limits of color constancy

Summary A summary of colorimetry is given and the limits of color constancy mechanism under changing illuminations are discussed.     

Pollinator response to flower color polymorphism and floral display in a plant with a single-locus floral color polymorphism: consequences for plant reproduction

Variation in flower color, particularly polymorphism, in which two or more different flower color phenotypes occur in the same population or species, may be affected or maintained by mechanisms that depend on pollinators. Furthermore, variation in floral display may affect pollinator response and plant reproductive success through changes in pollinator visitation and availability of compatible pollen. To asses if flower color polymorphism and floral display influences pollinator preferences and movements within and among plants and fitness-related variables we used the self-incompatible species Cosmos bipinnatus Cav. (Asteraceae), a model system with single-locus flower color polymorphism that comprises three morphs: white (recessive homozygous), pink (heterozygous co-dominate), and purple (dominant homozygous) flowers. We measured the preferences of pollinators for each morph and constancy index for each pollinator species, pollination visitation rate, floral traits, and female fitness measures. Flower color morphs differed in floral trait measures and seed production. Pollinators foraged nonrandomly with respect to flower color. The most frequent morph, the pink morph, was the most visited and pollinators exhibited the highest constancy for this morph. Moreover, this morph exhibited the highest female fitness. Pollinators responded strongly to floral display size, while probed more capitulums from plants with large total display sizes, they left a great proportion of them unvisited. Furthermore, total pollinator visitation showed a positive relation with female fitness. Results suggest that although pollinators preferred the heterozygous morph, they alternate indiscriminately among morphs making this polymorphism stable.     

Neural network approach for image chromatic adaptation for skin color detection

The goal of image chromatic adaptation is to remove the effect of illumination and to obtain color data that reflects precisely the physical contents of the scene. We present in this paper an approach to image chromatic adaptation using Neural Networks (NN) with application for detecting--adapting human skin color. The NN is trained on randomly chosen color images containing human subject under various illuminating conditions, thereby enabling the model to dynamically adapt to the changing illumination conditions. The proposed network predicts directly the illuminant estimate in the image so as to adapt to human skin color. The comparison of our method with Gray World, White Patch and NN on White Patch methods for skin color stabilization is presented. The skin regions in the NN stabilized images are successfully detected using a computationally inexpensive thresholding operation. We also present results on detecting skin regions on a data set of test images. The results are promising and suggest a new approach for adapting human skin color using neural networks.