A framework for analysing colour pattern geometry: adjacent colours

The analysis of colour pattern geometry is not as well advanced as the analysis of colour, although this reflects a lack of an analytical framework. The present study proposes an approach based on a consideration of which colours are adjacent to each other. Both vertebrate and invertebrate eyes do not take static images of the world but move across the field of view. As a consequence, the eye takes transects across the field of view responding to the colours and luminances within patches and to the colour and/or luminance transitions between patches. The framework and methods suggested here are based upon transects across colour patterns and make it possible to estimate colour pattern parameters that capture not only the relative areas of each patch class, but also the relative frequencies of colour/luminance transitions or adjacency. This allows tests of new hypotheses about colour patterns at the same time as including colour, pattern, and texture. Eleven groups of predictions are made with respect to the often conflicting needs of communication with conspecifics, avoiding predation, and finding food. New phenomena may be discovered as a result of these methods and predictions. For example, certain colour transitions may be used for species recognition even though the same colours are used by all species. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ??, ??–??.     

Colour Vision Impairment in Young Alcohol Consumers

Alcohol consumption among young adults is widely accepted in modern society and may be the starting point for abusive use of alcohol at later stages of life. Chronic alcohol exposure can lead to visual function impairment. In the present study, we investigated the spatial luminance contrast sensitivity, colour arrangement ability, and colour discrimination thresholds on young adults that weekly consume alcoholic beverages without clinical concerns. Twenty-four young adults were evaluated by an ophthalmologist and performed three psychophysical tests to evaluate their vision functions. We estimated the spatial luminance contrast sensitivity function at 11 spatial frequencies ranging from 0.1 to 30 cycles/degree. No difference in contrast sensitivity was observed comparing alcohol consumers and control subjects. For the evaluation of colour vision, we used the Farnsworth-Munsell 100 hue test (FM 100 test) to test subject’s ability to perform a colour arrangement task and the Mollon-Reffin test (MR test) to measure subject’s colour discrimination thresholds. Alcohol consumers made more mistakes than controls in the FM100 test, and their mistakes were diffusely distributed in the FM colour space without any colour axis preference. Alcohol consumers also performed worse than controls in the MR test and had higher colour discrimination thresholds compared to controls around three different reference points of a perceptually homogeneous colour space, the CIE 1976 chromaticity diagram. There was no colour axis preference in the threshold elevation observed among alcoholic subjects. Young adult weekly alcohol consumers showed subclinical colour vision losses with preservation of spatial luminance contrast sensitivity. Adolescence and young adult age are periods of important neurological development and alcohol exposure during this period of life might be responsible for deficits in visual functions, especially colour vision that is very sensitive to neurotoxicants.     

Rockpool Gobies Change Colour for Camouflage

Camouflage is found in a wide range of species living in numerous habitat types, offering protection from visually guided predators. This includes many species from the intertidal zone, which must cope with background types diverse in appearance and with multiple predator groups foraging at high and low tide. Many animals are capable of either relatively slow (hours, days, weeks) or rapid (seconds and minutes) colour change in order to better resemble the background against which they are found, but most work has been restricted to a few species or taxa. It is often suggested that many small intertidal fish are capable of colour change for camouflage, yet little experimental work has addressed this. Here, we test rock gobies (Gobius paganellus) for colour change abilities, and whether they can tune their appearance to match the background. In two experiments, we place gobies on backgrounds of different brightness (black or white), and of different colours (red and blue) and use digital image analysis and modelling of predator (avian) vision to quantify colour and luminance (perceived lightness) changes and camouflage. We find that gobies are capable of rapid colour change (occurring within one minute), and that they can change their luminance on lighter or darker backgrounds. When presented on backgrounds of different colours, gobies also change their colour (hue and saturation) while keeping luminance the same. These changes lead to predicted improvements in camouflage match to the background. Our study shows that small rockpool fish are capable of rapid visual change for concealment, and that this may be an important mechanism in many species to avoid predation, especially in complex heterogeneous environments.     

On the measurement and classification of colour in studies of animal colour patterns

In studies of animal colouration it is no longer necessary to rely on subjective assessments of colour and conspicuousness, nor on methods which rely upon human vision. This is important because animals vary greatly in colour vision and colour is context-dependent. New methods make it practical to measure the colour spectrum of pattern elements (patches) of animals and their visual backgrounds for the conditions under which patch spectra reach the conspecific's, predator's or prey's eyes. These methods can be used in both terrestrial and aquatic habitats. A patch's colour is dependent not only upon its reflectance spectrum, but also upon the ambient light spectrum, the transmission properties of air or water, and the veiling light spectrum. These factors change with time of day, weather, season and microhabitat, so colours must be measured under the conditions prevalent when colour patterns are normally used. Methods of measuring, classifying and comparing colours are presented, as well as techniques for assessing the conspicuousness of colour patterns as a whole. Some implications of the effect of environmental light and vision are also discussed.     

Trichromatic colour vision in the hummingbird hawkmoth, Macroglossum stellatarum L.

Hymenopterans have long been shown to choose colours by means of the spectral distribution and independently of the intensity (true colour vision). The same ability has only very recently been proven for two butterfly species. We present evidence for the existence of true colour vision in the European hummingbird hawkmoth, Macroglossum stellatarum. Moths were trained in dual-choice situations to spectral lights of a rewarding and an unrewarding wavelength. After training, unrewarded tests were performed during which the intensities of the lights were changed. The results confirm that the species has three spectral receptor types and uses true colour vision when learning the colour of a food source. If colour vision is not possible since only one receptor type is receiving input from both stimuli, the moths learn to associate some achromatic cue correlated to the receptor quantum catch, with the reward. The moths learn spectral cues rapidly and choose correctly after one to several rewarded visits even when trained to different colours in sequence. Accepted: 24 February 1999     

Communication and camouflage with the same 'bright' colours in reef fishes

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.     

The contribution of single and double cones to spectral sensitivity in budgerigars during changing light conditions

Bird colour vision is mediated by single cones, while double cones and rods mediate luminance vision in bright and dim light, respectively. In daylight conditions, birds use colour vision to discriminate large objects such as fruit and plumage patches, and luminance vision to detect fine spatial detail and motion. However, decreasing light intensity favours achromatic mechanisms and eventually, in dim light, luminance vision outperforms colour vision in all visual tasks. We have used behavioural tests in budgerigars (Melopsittacus undulatus) to investigate how single cones, double cones and rods contribute to spectral sensitivity for large (3.4°) static monochromatic stimuli at light intensities ranging from 0.08 to 63.5 cd/m². We found no influences of rods at any intensity level. Single cones dominate the spectral sensitivity function at intensities above 1.1 cd/m², as predicted by a receptor noise-limited colour discrimination model. Below 1.1 cd/m², spectral sensitivity is lower than expected at all wavelengths except 575 nm, which corresponds to double cone function. We suggest that luminance vision mediated by double cones restores visual sensitivity when single cone sensitivity quickly decreases at light intensities close to the absolute threshold of colour vision.     

Generalised Chromaticity Diagrams for Animals with <Emphasis Type="Italic">n</Emphasis>-chromatic Colour Vision

Chromaticity diagrams for tri- and tetrachromatic animals (with three and four cone classes in their retina, respectively, contributing to colour perception) are widely used in studies of animal colour vision. These diagrams not only allow the graphical representation of perceived colours, but the coordinates of colours plotted within these diagrams can be used to extract colour metrics, such as hue and chroma, or can be used directly in statistical analyses, and therefore aid our understanding of vision-mediated behaviour. However, many invertebrate species have more than four cone classes in their retina, and may therefore have pentachromatic or hexachromatic (or greater) vision. This paper describes an extension to the triangular and tetrahedral chromaticity diagrams commonly used for tri- and tetrachromats, respectively, that allows colour coordinates (and hence colour metrics) to be calculated for animals with more than four cone classes. Because the resulting chromaticity diagrams have more than three dimensions, meaningful ways to visualise the spatial position of plotted colours are discussed.     

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.     

Double cones are used for colour discrimination in the reef fish,Rhinecanthus aculeatus

Double cones (DCs) are the most common cone types in fish, reptiles and birds. It has been suggested that DCs are used for achromatic tasks such as luminance, motion and polarization vision. Here we show that a reef fish Rhinecanthus aculeatus can discriminate colours on the basis of the difference between the signals of individual members of DCs. This is the first direct evidence that individual members of DCs are used in colour vision as independent spectral channels.     

Simultaneous processing of spatial and chromatic components of patterned stimuli by the human visual system

We report measurements on discrimination of orientation and magnification made for elements differentiated in colour and/or luminance from their background. By performing measurements at a series of background luminances and for fixed luminance of the elements, we show that with colour contrast, discrimination for both spatial parameters is unimpaired when the background is at isoluminance with the elements. Under simple luminance contrast, however, these discriminations become poorer when the background luminance is within some +/- 5% of that of the elements, and are completely absent when the two values are the same. A deuteranomalous subject is unable to make the spatial discrimination around the isoluminance point for colour contrasts which are too small for him to distinguish, but for which subjects with normal colour vision maintain spatial discriminations at isoluminance. This observation establishes that the physiological mechanisms of normal colour vision, rather than stimulus artefacts, mediate the observed spatial discriminations. We conclude that the visual processing of colour and spatial parameters such as orientation and magnification are intrinsically related to each other.     

Visual ecology of flies with particular reference to colour vision and colour preferences

The visual ecology of flies is outstanding among insects due to a combination of specific attributes. Flies’ compound eyes possess an open rhabdom and thus separate rhabdomeres in each ommatidium assigned to two visual pathways. The highly sensitive, monovariant neural superposition system is based on the excitation of the peripheral rhabdomeres of the retinula cells R1–6 and controls optomotor reactions. The two forms of central rhabdomeres of R7/8 retinula cells in each ommatidium build up a system with four photoreceptors sensitive in different wavelength ranges and thought to account for colour vision. Evidence from wavelength discrimination tests suggests that all colour stimuli are assigned to one of just four colour categories, but cooperation of the two pathways is also evident. Flies use colour cues for various behavioural reactions such as flower visitation, proboscis extension, host finding, and egg deposition. Direct evidence for colour vision, the ability to discriminate colours according to spectral shape but independent of intensity, has been demonstrated for few fly species only. Indirect evidence for colour vision provided from electrophysiological recordings of the spectral sensitivity of photoreceptors and opsin genes indicates similar requisites in various flies; the flies’ responses to coloured targets, however, are much more diverse.     

Conjunctions of colour, luminance and orientation: the role of colour and luminance contrast on saliency and proximity grouping in texture segregation

To examine whether perceptual grouping on the basis of orientation can be performed simultaneously with or only subsequently to grouping according to colour or luminance, we tested whether subjects are able to segregate arrays of texture elements that differ from surrounding elements by conjunctions of either (i) colour and orientation, or (ii) luminance contrast and orientation, or (iii) luminance contrast polarity and orientation. Subjects were able to use conjunctions between luminance and orientation for segregation but not conjunctions between colour or contrast polarity and orientation. Our results suggest that (i) in agreement with earlier findings, there seem to exist no specific conjunction detectors for colour and orientation or contrast polarity and orientation, and (ii) when orientation defined textures are to be distinguished by virtue of differences in luminance, colour, or contrast polarity, luminance provides a much stronger cue than colour or contrast polarity for saliency-based orientation grouping.     

The uses of colour vision: behavioural and physiological distinctiveness of colour stimuli

Colour and greyscale (black and white) pictures look different to us, but it is not clear whether the difference in appearance is a consequence of the way our visual system uses colour signals or a by-product of our experience. In principle, colour images are qualitatively different from greyscale images because they make it possible to use different processing strategies. Colour signals provide important cues for segmenting the image into areas that represent different objects and for linking together areas that represent the same object. If this property of colour signals is exploited in visual processing we would expect colour stimuli to look different, as a class, from greyscale stimuli. We would also expect that adding colour signals to greyscale signals should change the way that those signals are processed. We have investigated these questions in behavioural and in physiological experiments. We find that male marmosets (all of which are dichromats) rapidly learn to distinguish between colour and greyscale copies of the same images. The discrimination transfers to new image pairs, to new colours and to image pairs in which the colour and greyscale images are spatially different. We find that, in a proportion of neurons recorded in the marmoset visual cortex, colour-shifts in opposite directions produce similar enhancements of the response to a luminance stimulus. We conclude that colour is, both behaviourally and physiologically, a distinctive property of images.     

SPATIAL FLOWER PARAMETERS AND INSECT SPATIAL VISION

The present article reviews recent and older literature on the spatial parameters that flowers display, as well as on the capacities of anthophilous insects to perceive and use these parameters for optimizing their foraging success. Although co-evolution of plants and pollinators has frequently been discussed with respect to floral colours and insect colour vision, it has rarely been assessed with respect to insect spatial vision and spatial floral cues, such as shape, pattern, size, contrast, symmetry, spatial frequency, contour density and orientation of contours. This review is an attempt to fill this gap. From experimental findings and observations on both flowers and insects, we arrive at the conclusion that all of the spatial and spatio-temporal parameters that flowers offer are relevant to the foraging task and are tuned to the insect's visual capacities and visually guided behaviour. We try, in addition, to indicate that temporal cues are closely related to spatial cues, and must therefore be included when flower–pollinator interactions are examined. We include results that show that colour vision and spatial vision have diverged over the course of evolution, particularly regarding the processing of spatio-temporal information, but that colour vision plays a role in the processing of spatial cues that are independent of temporal parameters. By presenting this review we hope to contribute to closer collaboration among scientists working in the vast fields of botany, ecology, evolution, ethology and sensory physiology.     

Colour and brightness coding in the central nervous system: theoretical aspects and visual evoked potentials to homogeneous red and green stimuli

We designed visual evoked potentials experiments to study the differential aspects of colour and brightness coding in man. The substitution of equally bright red and green stimuli for a background yellow was investigated and compared with different luminance increments and decrements of red and green. A dominant N87 component was found for a colour change from yellow to brighter red colours, which was less pronounced for green and absent for yellow luminance changes. It is also absent for pure red luminance increments and green luminance changes, but reappears with red luminance decrements or red-offset. The data are discussed within the framework of a new concept of how the visual system fuses red-green information and black-white border information. Retinal X-cells can transmit colour and high spatial frequency achromatic information simultaneously by encoding only the presence of edges (a.c.) for the black-white stimuli and the presence of both edges (a.c.) and uniform areas of colour (d.c.) for red-green stimuli. Phylogenetically this kind of information transmission enables colour vision to be implemented in a retina such as the cat's by adding only a second class of cones. Barlow's economy principle will be violated for colour in the periphery, but restored early in the striate cortex where there is an early decoding of the combined chromatic and achromatic information by the concentric double opponent cells. The N87 behaviour correlates with the proposed discharge of peripheral X-type cells, but not with the discharge of cortical double opponent concentric or simple cells, which no longer respond to homogeneous colour stimuli. It is suggested that N87 may be generated by geniculate afferents in the dendritic arborization of cortical cells, reflecting the behaviour of peripheral units, and thus the violation of the economy principle, rather than the next step in cortical processing. The early cortical restoration of the economy principle is supported by the absence of any further dissociated behaviour for colour and brightness in later components.     

The Absolute Threshold of Colour Vision in the Horse

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.     

Comparing entire colour patterns as birds see them

Colour patterns and their visual backgrounds consist of a mosaic of patches that vary in colour, brightness, size, shape and position. Most studies of crypsis, aposematism, sexual selection, or other forms of signalling concentrate on one or two patch classes (colours), either ignoring the rest of the colour pattern, or analysing the patches separately. We summarize methods of comparing colour patterns making use of known properties of bird eyes. The methods are easily modifiable for other animal visual systems. We present a new statistical method to compare entire colour patterns rather than comparing multiple pairs of patches. Unlike previous methods, the new method detects differences in the relationships among the colours, not just differences in colours. We present tests of the method's ability to detect a variety of kinds of differences between natural colour patterns and provide suggestions for analysis.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 405–431.     

Colour preferences and colour vision in poultry chicks

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.     

How and why are uniformly polarization-sensitive retinae subject to polarization-related artefacts? Correction of some errors in the theory of polarization-induced false colours

If the photoreceptors of a colour vision system are polarization sensitive, the system detects polarization-induced false colours. Based on the functional similarities between polarization vision and colour vision, earlier it was believed that a uniformly polarization-sensitive (insect) retina (UPSR)-in which receptors of all spectral types have the same polarization sensitivity ratio and microvilli direction-cannot detect polarization-induced false colours. Here we show that, contrary to this belief, a colour vision based on a UPSR is subject to polarization-related artefacts, because both the degree and the angle of polarization of light reflected from natural surfaces depend on wavelength. Our second goal is to correct certain errors in the theory of polarizational false colours. The quantitative estimation of the influence of polarization sensitivity on colour vision was recently motivated by the suggestion that certain Papilio butterflies detect such false colours. The theoretical basis of this subject is to calculate the colour loci in the colour space of a visual system from the quantum catches of polarization-sensitive receptors of different spectral types. Horváth et al. (J. Exp. Biol. 205 (2002) 3281) gave the first exact mathematical and receptor-physiological derivation of formulae for these calculations. Here we prove that the two formulae given earlier by others are inappropriate or erroneous. This, however, does not influence the validity of the experimental data and the principal conclusions drawn about the colour vision and polarization sensitivity in Papilio butterflies.