The perception of an object's colour remains constant despite large variations in the chromaticity of the illumination—colour constancy. Hering suggested that memory colours, the typical colours of objects, could help in estimating the illuminant's colour and therefore be an important factor in establishing colour constancy. Here we test whether the presence of objects with diagnostical colours (fruits, vegetables, etc) within a scene influence colour constancy for unknown coloured objects in the scene. Subjects matched one of four Munsell papers placed in a scene illuminated under either a reddish or a greenish lamp with the Munsell book of colour illuminated by a neutral lamp. The Munsell papers were embedded in four different scenes—one scene containing diagnostically coloured objects, one scene containing incongruent coloured objects, a third scene with geometrical objects of the same colour as the diagnostically coloured objects, and one scene containing non-diagnostically coloured objects (eg, a yellow coffee mug). All objects were placed against a black background. Colour constancy was on average significantly higher for the scene containing the diagnostically coloured objects compared with the other scenes tested. We conclude that the colours of familiar objects help in obtaining colour constancy for unknown objects.
Colour constancy; memory colours; object colour; colour appearance
Many animals use the spectral distribution of light to guide behaviour, but whether they have colour vision has been debated for over a century. Our strong subjective experience of colour and the fact that human vision is the paradigm for colour science inevitably raises the question of how we compare with other species. This article outlines four grades of ‘colour vision’ that can be related to the behavioural uses of spectral information, and perhaps to the underlying mechanisms. In the first, even without an (image-forming) eye, simple organisms can compare photoreceptor signals to locate a desired light environment. At the next grade, chromatic mechanisms along with spatial vision guide innate preferences for objects such as food or mates; this is sometimes described as wavelength-specific behaviour. Here, we compare the capabilities of di- and trichromatic vision, and ask why some animals have more than three spectral types of receptors. Behaviours guided by innate preferences are then distinguished from a grade that allows learning, in part because the ability to learn an arbitrary colour is evidence for a neural representation of colour. The fourth grade concerns colour appearance rather than colour difference: for instance, the distinction between hue and saturation, and colour categorization. These higher-level phenomena are essential to human colour perception but poorly known in animals, and we suggest how they can be studied. Finally, we observe that awareness of colour and colour qualia cannot be easily tested in animals.
colour vision; phototaxis; colour preference; colour learning; colour categorization; chromaticity
We investigated the memory colour effect for colour diagnostic artificial objects. Since knowledge about these objects and their colours has been learned in everyday life, these stimuli allow the investigation of the influence of acquired object knowledge on colour appearance. These investigations are relevant for questions about how object and colour information in high-level vision interact as well as for research about the influence of learning and experience on perception in general. In order to identify suitable artificial objects, we developed a reaction time paradigm that measures (subjective) colour diagnosticity. In the main experiment, participants adjusted sixteen such objects to their typical colour as well as to grey. If the achromatic object appears in its typical colour, then participants should adjust it to the opponent colour in order to subjectively perceive it as grey. We found that knowledge about the typical colour influences the colour appearance of artificial objects. This effect was particularly strong along the daylight axis.
Memory Colours; Artificial Objects; Object Colours; Colour Diagnosticity; Colour Appearance; Daylight Variation; Past Experience; Prior Knowledge
Nocturnal animals are said to sacrifice colour vision in favour of increased absolute sensitivity. This is true for most vertebrates that possess a dual retina with a single type of rod for colour-blind night vision and multiple types of cone for diurnal colour vision. However, among the nocturnal vertebrates, geckos are unusual because they have no rods but three cone types. Here, we show that geckos use their cones for colour vision in dim light. Two specimens of the nocturnal helmet gecko Tarentola (formerly Geckonia) chazaliae were able to discriminate blue from grey patterns by colour alone. Experiments were performed at 0.002 cd m(-2), a light intensity similar to dim moonlight. We conclude that nocturnal geckos can use cone-based colour vision at very dim light levels when humans rely on colour-blind rod vision.
Arrhythmic mammals are active both during day and night if they are allowed. The arrhythmic horses are in possession of one of the largest terrestrial animal eyes and the purpose of this study is to reveal whether their eye is sensitive enough to see colours at night. During the day horses are known to have dichromatic colour vision. To disclose whether they can discriminate colours in dim light a behavioural dual choice experiment was performed. We started the training and testing at daylight intensities and the horses continued to choose correctly at a high frequency down to light intensities corresponding to moonlight. One Shetland pony mare, was able to discriminate colours at 0.08 cd/m2, while a half blood gelding, still discriminated colours at 0.02 cd/m2. For comparison, the colour vision limit for several human subjects tested in the very same experiment was also 0.02 cd/m2. Hence, the threshold of colour vision for the horse that performed best was similar to that of the humans. The behavioural results are in line with calculations of the sensitivity of cone vision where the horse eye and human eye again are similar. The advantage of the large eye of the horse lies not in colour vision at night, but probably instead in achromatic tasks where presumably signal summation enhances sensitivity.
By adaptational and other mechanisms, the visual system can compensate for moderate changes in the colour of the illumination on a scene. Although the colours of most surfaces are perceived to be constant (“colour constancy”), some are not. The effect of these residual colour changes on the ability of observers to identify surfaces by their apparent colour was determined theoretically from high-resolution hyperspectral images of natural scenes under different daylights with correlated colour temperatures 4300 K, 6500 K, and 25000 K. Perceived differences between colours were estimated with an approximately uniform colour-distance measure. The information preserved under illuminant changes increased with the number of surfaces in the sample, but was limited to a relatively low asymptotic value, indicating the importance of physical factors in constraining identification by apparent colour.
Shannon information; spectral reflectance; colour constancy; colour matching; colour space
It is hypothesized that colour vision and opponent processing of colour signals in the visual system evolved as a means of overcoming the extremely unfavourable lighting conditions in the natural environment of early vertebrates. The significant flicker of illumination inherent in the shallow-water environment complicated the visual process in the achromatic case, in particular preventing early detection of enemies. The presence of two spectral classes of photoreceptors and opponent interaction of their signals at a subsequent retinal level allowed elimination of the flicker from the retinal image. This new visual function provided certain advantages concerning reaction times and favoured survival. This assumption explains why the building blocks for colour vision arose so early, i.e. just after the active predatory lifestyle was mastered. The principal functions of colour vision inherent in extant animals required a more complex neural machinery for colour processing and evolved later as the result of a change in visual function favouring colour vision.
Colour constancy assessed by asymmetric simultaneous colour matching usually reveals limited levels of performance in the unadapted eye. Yet observers can readily discriminate illuminant changes on a scene from changes in the spectral reflectances of the surfaces making up the scene. This ability is probably based on judgements of relational colour constancy, in turn based on the physical stability of spatial ratios of cone excitations under illuminant changes. Evidence is presented suggesting that the ability to detect violations in relational colour constancy depends on temporal transient cues. Because colour constancy and relational colour constancy are closely connected, it should be possible to improve estimates of colour constancy by introducing similar transient cues into the matching task. To test this hypothesis, an experiment was performed in which observers made surface-colour matches between patterns presented in the same position in an alternating sequence with period 2 s or, as a control, presented simultaneously, side-by-side. The degree of constancy was significantly higher for sequential presentation, reaching 87% for matches averaged over 20 observers. Temporal cues may offer a useful source of information for making colour-constancy judgements.
colour constancy; relational colour constancy; temporal transients; surface colour; spatial cone-excitation ratios
The dramatic colours of biological communication signals raise questions about how animals perceive suprathreshold colour differences, and there are long-standing questions about colour preferences and colour categorization by non-human species. This study investigates preferences of foraging poultry chicks (Gallus gallus) as they peck at coloured objects. Work on colour recognition often deals with responses to monochromatic lights and how animals divide the spectrum. We used complementary colours, where the intermediate is grey, and related the chicks' choices to three models of the factors that may affect the attractiveness. Two models assume that attractiveness is determined by a metric based on the colour discrimination threshold either (i) by chromatic contrast against the background or (ii) relative to an internal standard. An alternative third model is that categorization is important. We tested newly hatched and 9-day-old chicks with four pairs of (avian) complementary colours, which were orange, blue, red and green for humans. Chromatic contrast was more relevant to newly hatched chicks than to 9-day-old birds, but in neither case could contrast alone account for preferences; especially for orange over blue. For older chicks, there is evidence for categorization of complementary colours, with a boundary at grey.
colour; vision; behaviour; Gallus gallus; chicks
Colour vision enables animals to detect and discriminate differences in chromatic cues independent of brightness. How the bee visual system manages this task is of interest for understanding information processing in miniaturized systems, as well as the relationship between bee pollinators and flowering plants. Bees can quickly discriminate dissimilar colours, but can also slowly learn to discriminate very similar colours, raising the question as to how the visual system can support this, or whether it is simply a learning and memory operation. We discuss the detailed neuroanatomical layout of the brain, identify probable brain areas for colour processing, and suggest that there may be multiple systems in the bee brain that mediate either coarse or fine colour discrimination ability in a manner dependent upon individual experience. These multiple colour pathways have been identified along both functional and anatomical lines in the bee brain, providing us with some insights into how the brain may operate to support complex colour discrimination behaviours.
brain; conditioning; insect learning; task switching
Some theories of surface-colour perception assume that observers estimate the illuminant on a scene so that its effects can be discounted. A critical test of this interpretation of colour constancy is whether surface-colour matching is worse when the number of surfaces in a scene is so small that any illuminant estimate is unreliable. In the experiment reported here, observers made asymmetric colour matches between pairs of simultaneously presented Mondrian-like patterns under different daylights. The patterns had either 49 surfaces or a minimal 2 surfaces. No significant effect of number was found, suggesting that illuminant estimates are unnecessary for surface-colour matching.
The colour of animals' skin, fur, feathers or cuticula has been estimated in a large number of studies. The methods used to do so are diverse, with some being costly and not available to all researchers. In a study to measure plumage colour in a bird species, a new method of creating a colour chart was developed. While colour-charts have their own limitations, these can be minimised when they have the following properties: 1) being readily available to the majority of biologists, 2) containing a large array of colours to allow accurate recording and differentiation of subtle colour differences, 3) low cost, 4) adhering to a world-wide standard, and 5) being available in both hard-copy and digital formats to allow for various analytical methods. The method described below satisfies all of these requirements.
Colour charts estimated to fit the range of the species' plumage colours were created on the computer screen using web software that allowed for HTML-coding (in this case Dreamweaver™). The charts were adjusted using feathers from dead specimens until a satisfying range of darker and lighter colours were found. The resulting chart was printed out and was successfully used in the field to determine the plumage colour of hand-held birds.
Access to a computer and printer, and the software to enable the creation of a chart, is within the reach of the vast majority of biologists. The numbers of colours that can be generated should suit most studies, with the advantage of the method being that the chart can be individually tailored to the species under study. HTML colour coding is a worldwide standard, thus the colours used in studies can be described in the methods section of journal articles using the six-digit alphanumeric code. We believe this method is very useful as a low-tech method for future estimation of individual colour.
A Lantern for lesting Colour-Vision is arranged to show test colours in pairs as in the Board of Trade Lantern. It is adapted to use electric light, and is standardized by stringent testing. The paper discusses the experiments and considerations which led to the formulation of the allowable tolerances in the transmission and colour co-ordinate specifications of the filters, the colour temperature of the lamps and so on. The results of tests on normal and colour-defective subjects are described.
Colour changes in animals may be triggered by a variety of social and environmental factors and may occur over a matter of seconds or months. Crustaceans, like fiddler crabs (genus Uca), are particularly adept at changing their colour and have been the focus of numerous studies. However, few of these studies have attempted to quantitatively describe the individual variation in colour and pattern or their adaptive significance. This paper quantitatively describes the colour patterns of the fiddler crab Uca capricornis and their ability to change on a socially significant timescale. The most dramatic changes in colour pattern are associated with moulting. These ontogenetic changes result in a general reduction of the colour pattern with increasing size, although females are more colourful and variable than similarly-sized males. Uca capricornis are also capable of rapid colour changes in response to stress, but show no endogenous rhythms associated with the semilunar and tidal cycles commonly reported in other fiddler crabs. The extreme colour polymorphism and the relative stability of the colour patterns in Uca capricornis are consistent with their use in visually mediated mate recognition.
A group of doctors with congenital colour vision deficiency (CCVD) were compared with a group of controls in their assessment of colour blocks in the colour range of a widely available blood glucose testing stick. The majority of doctors with CCVD agreed with controls on colour matching. However, subjects with severe CCVD tended to match test blocks to a wider range of options than either those with a less severe defect or controls. This paper discusses the implications of these findings.
In animals, iridescence is generated by the interaction of light with biological tissues that are nanostructured to produce thin films or diffraction gratings. Uniquely among animal visual signals, the study of iridescent coloration contributes to biological and physical sciences by enhancing our understanding of the evolution of communication strategies, and by providing insights into physical optics and inspiring biomimetic technologies useful to humans. Iridescent colours are found in a broad diversity of animal taxa ranging from diminutive marine copepods to terrestrial insects and birds. Iridescent coloration has received a surge of research interest of late, and studies have focused on both characterizing the nanostructures responsible for producing iridescence and identifying the behavioural functions of iridescent colours. In this paper, we begin with a brief description of colour production mechanisms in animals and provide a general overview of the taxonomic distribution of iridescent colours. We then highlight unique properties of iridescent signals and review the proposed functions of iridescent coloration, focusing, in particular, on the ways in which iridescent colours allow animals to communicate with conspecifics and avoid predators. We conclude with a brief overview of non-communicative functions of iridescence in animals. Despite the vast amount of recent work on animal iridescence, our review reveals that many proposed functions of iridescent coloration remain virtually unexplored, and this area is clearly ripe for future research.
iridescence; behaviour; function; communication; sexual selection; predation
Animals change their body coloration for a variety of purposes including communication, thermoregulation and crypsis. The cues that trigger adaptive colour change are often unclear, and the role of colour vision remains largely untested. Here, we investigated the bluestriped fangblenny (Plagiotremus rhinorhynchos), an aggressive mimic that changes its body coloration to impersonate a variety of coral reef fishes. In this field, we determined the fish species that the fangblenny associated with and measured the spectral reflectance of mimics and their models. We measured the spectral absorbance characteristics of the retinal photoreceptor visual pigments in the bluestriped fangblenny using microspectrophotometry and found it to have rod photoreceptors (λmax 498 nm), single cones (449 nm) and double cones (561 nm principal member; 520 nm accessory member). Using theoretical vision models, fangblennies could discriminate between the colours they adopted and the colours of the fish they associated with. Potential signal receivers (Abudefduf abdominalis and Ctenochaetus strigosus) perceived colours of most mimics to closely resemble fishes they associated with. However, fishes with ultraviolet-sensitive visual pigments were better at discriminating between mimics and models. Therefore, colour vision could be used by fangblennies when initiating colour change enabling them to accurately resemble fishes they associate with and to avoid detection by signal receivers.
facultative mimicry; microspectrophotometry; colour vision; colour change; spectral reflectance; coral reef fish
The aim of this paper is to classify the land covered with oat crops, and the quantification of frost damage on oats, while plants are still in the flowering stage. The images are taken by a digital colour camera CCD-based sensor. Unsupervised classification methods are applied because the plants present different spectral signatures, depending on two main factors: illumination and the affected state. The colour space used in this application is CIELab, based on the decomposition of the colour in three channels, because it is the closest to human colour perception. The histogram of each channel is successively split into regions by thresholding. The best threshold to be applied is automatically obtained as a combination of three thresholding strategies: (a) Otsu’s method, (b) Isodata algorithm, and (c) Fuzzy thresholding. The fusion of these automatic thresholding techniques and the design of the classification strategy are some of the main findings of the paper, which allows an estimation of the damages and a prediction of the oat production.
digital image sensor; agricultural images; unsupervised classification; automatic thresholding; CIELab colour space; fuzzy error matrix; oat frost damage
Many animals use concealing markings to reduce the risk of predation. These include background pattern matching (crypsis), where the coloration matches a random sample of the background and disruptive patterns, whose effectiveness has been hypothesized to lie in breaking up the body into a series of apparently unrelated objects. We have previously established the effectiveness of disruptive coloration against avian predators, using artificial moth-like stimuli with colours designed to match natural backgrounds as perceived by birds. Here, we investigate the mechanism by which disruptive patterns reduce detectability, using a computational vision model of edge detection applied to photographs of our experimental stimuli, calibrated for bird colour vision. We show that, disruptive coloration is effective by exploiting edge detection algorithms that we use to model early visual processing. Thus, ‘false’ edges are detected within the body rather than at its periphery, so inhibiting successful detection of the animal's body outline.
disruptive coloration; background matching; edge detection; vision; camouflage
The eyes of most diurnal reptiles and birds contain coloured retinal filters-oil droplets. Although these filters are widespread, their adaptive advantage remains uncertain. To understand why coloured oil droplets appeared and were retained during evolution, I consider both the benefits and the costs of light filtering in the retina. Oil droplets decrease cone quantum catch and reduce the overlap in sensitivity between spectrally adjacent cones. The reduction of spectral overlap increases the volume occupied by object colours in a cone space, whereas the decrease in quantum catch increases noise, and thus reduces the discriminability of similar colours. The trade-off between these two effects determines the total benefit of oil droplets. Calculations show that coloured oil droplets increase the number of object colours that can be discriminated, and thus are beneficial for colour vision.
This review outlines how eyes of terrestrial vertebrates and insects meet the competing requirements of coding both spatial and spectral information. There is no unique solution to this problem. Thus, mammals and honeybees use their long-wavelength receptors for both achromatic (luminance) and colour vision, whereas flies and birds probably use separate sets of photoreceptors for the two purposes. In particular, we look at spectral tuning and diversification among ‘long-wavelength’ receptors (sensitivity maxima at greater than 500 nm), which play a primary role in luminance vision. Data on spectral sensitivities and phylogeny of visual photopigments can be incorporated into theoretical models to suggest how eyes are adapted to coding natural stimuli. Models indicate, for example, that animal colour vision—involving five or fewer broadly tuned receptors—is well matched to most natural spectra. We can also predict that the particular objects of interest and signal-to-noise ratios will affect the optimal eye design. Nonetheless, it remains difficult to account for the adaptive significance of features such as co-expression of photopigments in single receptors, variation in spectral sensitivities of mammalian L-cone pigments and the diversification of long-wavelength receptors that has occurred in several terrestrial lineages.
colour vision; evolution; ecology; retina; photoreceptor
Reef fishes present the observer with the most diverse and stunning assemblage of animal colours anywhere on earth. The functions of some of these colours and their combinations are examined using new non-subjective spectrophotometric measurements of the colours of fishes and their habitat. Conclusions reached are as follows: (i) the spectra of colours in high spatial frequency patterns are often well designed to be very conspicuous to a colour vision system at close range but well camouflaged at a distance; (ii) blue and yellow, the most frequently used colours in reef fishes, may be good for camouflage or communication depending on the background they are viewed against; and (iii) reef fishes use a combination of colour and behaviour to regulate their conspicuousness and crypsis.
This paper presents an improvement in the colour image segmentation in the Hue Saturation (HS) sub-space. The authors propose to inject (add) a colour vector in the Red Green Blue (RGB) space to increase the class separation in the HS plane. The goal of the work is the development of an algorithm to obtain the optimal colour vector for injection that maximizes the separation between the classes in the HS plane. The chromatic Chrominace-1 Chrominance-2 sub-space (of the Luminance Chrominace-1 Chrominance-2 (YC1C2) space) is used to obtain the optimal vector to add. The proposal is applied on each frame of a colour image sequence in real-time. It has been tested in applications with reduced contrast between the colours of the background and the object, and particularly when the size of the object is very small in comparison with the size of the captured scene. Numerous tests have confirmed that this proposal improves the segmentation process, considerably reducing the effects of the variation of the light intensity of the scene. Several tests have been made in skin segmentation in applications for sign language recognition via computer vision, where an accurate segmentation of hands and face is required.
pixel classification; colour clustering; colour segmentation; class separation; colour sub-spaces; colour injection
Perception of the body's outline and three-dimensional shape arises from visual cues such as shading, contour, perspective and texture. When a uniformly coloured prey animal is illuminated from above by sunlight, a shadow may be cast on the body, generating a brightness contrast between the dorsal and ventral surfaces. For animals such as caterpillars, which live among flat leaves, a difference in reflectance over the body surface may degrade the degree of background matching and provide cues to shape from shading. This may make otherwise cryptic prey more conspicuous to visually hunting predators. Cryptically coloured prey are expected to match their substrate in colour, pattern and texture (though disruptive patterning is an exception), but they may also abolish self-shadowing and therefore either reduce shape cues or maintain their degree of background matching through countershading: a gradation of pigment on the body of an animal so that the surface closest to illumination is darker. In this study, we report the results from a series of field experiments where artificial prey resembling lepidopteran larvae were presented on the upper surfaces of beech tree branches so that they could be viewed by free-living birds. We demonstrate that countershading is superior to uniform coloration in terms of reducing attack by free-living predators. This result persisted even when we fixed prey to the underside of branches, simulating the resting position of many tree-living caterpillars. Our experiments provide the first demonstration, in an ecologically valid visual context, that shadowing on bodies (such as lepidopteran larvae) provides cues that visually hunting predators use to detect potential prey species, and that countershading counterbalances shadowing to enhance cryptic protection.
countershading; crypsis; predation; animal coloration; defensive coloration
Colour vision allows animals to reliably distinguish differences in the distributions of spectral energies reaching the eye. Although not universal, a capacity for colour vision is sufficiently widespread across the animal kingdom to provide prima facie evidence of its importance as a tool for analysing and interpreting the visual environment. The basic biological mechanisms on which vertebrate colour vision ultimately rests, the cone opsin genes and the photopigments they specify, are highly conserved. Within that constraint, however, the utilization of these basic elements varies in striking ways in that they appear, disappear and emerge in altered form during the course of evolution. These changes, along with other alterations in the visual system, have led to profound variations in the nature and salience of colour vision among the vertebrates. This article concerns the evolution of colour vision among the mammals, viewing that process in the context of relevant biological mechanisms, of variations in mammalian colour vision, and of the utility of colour vision.
colour vision; opsin genes; photopigments; dichromacy; trichromacy; mammals