Camouflage is the primary defence of many animals and includes multiple strategies that interfere with figure-ground segmentation and object recognition. While matching background colours and textures is widespread and conceptually straightforward, less well explored are the optical ‘tricks’, collectively called disruptive colouration, that exploit perceptual grouping mechanisms. Adjacent high contrast colours create false edges, but this is not sufficient for an object’s shape to be broken up; some colours must blend with the background. We test the novel hypothesis that this will be particularly effective when the colour patches on the animal appear to belong to, not merely different background colours, but different background objects. We used computer-based experiments where human participants had to find cryptic targets on artificial backgrounds. Creating what appeared to be bi-coloured foreground objects on bi-coloured backgrounds, we generated colour boundaries that had identical local contrast but either lay within or between (illusory) objects. As predicted, error rates for targets matching what appeared to be different background objects were higher than for targets which had otherwise identical local contrast to the background but appeared to belong to single background objects. This provides evidence for disruptive colouration interfering with higher-level feature integration in addition to previously demonstrated low-level effects involving contour detection. In addition, detection was impeded in treatments where targets were on or in close proximity to multiple background colour or tone boundaries. This is consistent with other studies which show a deleterious influence of visual ‘clutter’ or background complexity on search.
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
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.
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
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
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.
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.
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
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
The colours of birds are diverse but limited relative to the colours they can perceive. This mismatch may be partially caused by the properties of their colour-production mechanisms. Aside from pigments, several classes of highly ordered nanostructures (thin films, amorphous three-dimensional arrays) can produce a range of colours. However, the variability of any single nanostructural class has rarely been explored. Dabbling ducks are a speciose clade with substantial interspecific variation in the iridescent coloration of their wing patches (specula). Here, we use electron microscopy, spectrophotometry, polarization and refractive index-matching experiments, and optical modelling to examine these colours. We show that, in all species examined, speculum colour is produced by a photonic heterostructure consisting of both a single thin-film of keratin and a two-dimensional hexagonal lattice of melanosomes in feather barbules. Although the range of possible variations of this heterostructure is theoretically broad, only relatively close-packed, energetically stable variants producing more saturated colours were observed, suggesting that ducks are either physically constrained to these configurations or are under selection for the colours that they produce. These data thus reveal a previously undescribed biophotonic structure and suggest that both physical variability and constraints within single nanostructural classes may help explain the broader patterns of colour across Aves.
photonic crystal; Anatidae; iridescence; thin films; sexual selection; nanophotonics
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
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
A combination of structural and pigmentary components is responsible for many of the colour displays of animals. Despite the ubiquity of this type of coloration, neither the relative contribution of structures and pigments to variation in such colour displays nor the relative effects of extrinsic factors on the structural and pigment-based components of such colour has been determined. Understanding the sources of colour variation is important because structures and pigments may convey different information to conspecifics. In an experiment on captive American goldfinches Carduelis tristis, we manipulated two parameters, carotenoid availability and food availability, known to affect the expression of carotenoid pigments in a full-factorial design. Yellow feathers from these birds were then analysed in two ways. First, we used full-spectrum spectrometry and high-performance liquid chromatography to examine the extent to which variation in white structural colour and total carotenoid content was associated with variation in colour properties of feathers. The carotenoid content of yellow feathers predicted two colour parameters (principal component 1—representing high values of ultraviolet and yellow chroma and low values of violet–blue chroma—and hue). Two different colour parameters (violet–blue and yellow chroma) from white de-pigmented feathers, as well as carotenoid content, predicted reflectance measurements from yellow feathers. Second, we determined the relative effects of our experimental manipulations on white structural colour and yellow colour. Carotenoid availability directly affected yellow colour, while food availability affected it only in combination with carotenoid availability. None of our manipulations had significant effects on the expression of white structural colour. Our results suggest that the contribution of microstructures to variation in the expression of yellow coloration is less than the contribution of carotenoid content, and that carotenoid deposition is more dependent on extrinsic variability than is the production of white structural colour.
American goldfinch; Carduelis tristis; carotenoid pigmentation; diet; honest signalling; sexual selection
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.
This paper deals with the generation of accurate, dense and coloured 3D models of outdoor scenarios from scanners. This is a challenging research field in which several problems still remain unsolved. In particular, the process of 3D model creation in outdoor scenes may be inefficient if the scene is digitalized under unsuitable technical (specific scanner on-board camera) and environmental (rain, dampness, changing illumination) conditions. We address our research towards the integration of images and range data to produce photorealistic models. Our proposal is based on decoupling the colour integration and geometry reconstruction stages, making them independent and controlled processes. This issue is approached from two different viewpoints. On the one hand, given a complete model (geometry plus texture), we propose a method to modify the original texture provided by the scanner on-board camera with the colour information extracted from external images taken at given moments and under specific environmental conditions. On the other hand, we propose an algorithm to directly assign external images onto the complete geometric model, thus avoiding tedious on-line calibration processes. We present the work conducted on two large Roman archaeological sites dating from the first century A.D., namely, the Theatre of Segobriga and the Fori Porticus of Emerita Augusta, both in Spain. The results obtained demonstrate that our approach could be useful in the digitalization and 3D modelling fields.
3D modelling; texture fusion; 3D digitalization; range data
Despite major differences between human and avian colour vision, previous studies of cuckoo egg mimicry have used human colour vision (or standards based thereon) to assess colour matching. Using ultraviolet-visible reflectance spectrophotometry (300-700 nm), we measured museum collections of eggs of the red-chested cuckoo and its hosts. The first three principal components explained more than 99% of the variance in spectra, and measures of cuckoo host egg similarity derived from these transformations were compared with measures of cuckoo host egg similarity estimated by human observers unaware of the hypotheses we were testing. Monte Carlo methods were used to simulate laying of cuckoo eggs at random in nests. Results showed that host and cuckoo eggs were very highly matched for an ultraviolet versus greenness component, which was not detected by humans. Furthermore, whereas cuckoo and host were dissimilar in achromatic brightness, humans did not detect this difference. Our study thus reveals aspects of cuckoo-host egg colour matching which have hitherto not been described. These results suggest subtleties and complexities in the evolution of host-cuckoo egg mimicry that were not previously suspected. Our results also have the potential to explain the longstanding paradox that some host species accept cuckoo eggs that are non-mimetic to the human eye.
Human perception of plant leaf and flower colour can influence species management. Colour and colour contrast may influence the detectability of invasive or rare species during surveys. Quantitative, repeatable measures of plant colour are required for comparison across studies and generalisation across species. We present a standard method for measuring plant leaf and flower colour traits using images taken with digital cameras. We demonstrate the method by quantifying the colour of and colour difference between the flowers of eleven grassland species near Falls Creek, Australia, as part of an invasive species detection experiment. The reliability of the method was tested by measuring the leaf colour of five residential garden shrub species in Ballarat, Australia using five different types of digital camera. Flowers and leaves had overlapping but distinct colour distributions. Calculated colour differences corresponded well with qualitative comparisons. Estimates of proportional cover of yellow flowers identified using colour measurements correlated well with estimates obtained by measuring and counting individual flowers. Digital SLR and mirrorless cameras were superior to phone cameras and point-and-shoot cameras for producing reliable measurements, particularly under variable lighting conditions. The analysis of digital images taken with digital cameras is a practicable method for quantifying plant flower and leaf colour in the field or lab. Quantitative, repeatable measurements allow for comparisons between species and generalisations across species and studies. This allows plant colour to be related to human perception and preferences and, ultimately, species management.
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.
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
Many birds see in the ultraviolet (300–400 nm), but there is limited evidence for colour communication (signalling by spectral shape independently of brightness) in this 'hidden' waveband. Such data are critical for the understanding of extravagant plumage colours, some of which show considerable UV reflectance. We investigated UV colour vision in female social responses to the male UV/violet ornament in bluethroats, Luscinia s. svecica. In an outdoor aviary at the breeding grounds, 16 females were each presented with a unique pair of males of equal age. In UVR (UV reduction) males, sunblock chemicals reduced only the UV reflectance and thereby the spectral shape (colour) of the throat ornament. In NR (neutral reduction) males, an achromatic pigment in the same base solvent (preen gland fat) was used for a corresponding but uniform brightness reduction. Both colour and brightness effects were invisible to human eyes, and were monitored by spectrometry. In 13 of the 16 trials, the female associated most with the NR male, a preference that implies that UV colour vision is used in mate choice by female bluethroats. Reflectance differences between one-year-old and older males were significant only in UV, suggestive of a UV colour cue in age-related mate preferences.
Colour Vision Reflectance Ultraviolet Mate Choice Sexual Selection Luscinia Svecica
In this paper we describe a low-cost spectrometric detector that can be easily assembled in a laboratory for less than €80 with a minimal number of optical components and which has proved sensitive and flexible enough for real-life applications. The starting point for the idea to construct this small, compact low-cost spectrometric detector was the decision to use a tri-colour light-emitting diode (LED) of the red-green-blue (RGB) type as a light source with the objective of achieving some flexibility in the selection of the wavelength (430 nm, 565 nm, 625 nm) but avoiding the use of optical fibres. Due to the dislocation of the emitters of the different coloured light, the tri-colour LED-based detector required an optical geometry that differs from those that are described in literature. The proposed novel geometry, with a coil-type glass flow-through cell with up to four ascending turns, proved useful and fit for the purpose. The simplicity of the device means it requires a minimal number of optical components, i.e., only a tri-colour LED and a photoresistor. In order to make a flow-injection analysis (FIA) with the spectrometric detector even more accessible for those with a limited budget, we additionally describe a low-cost simplified syringe-pump-based FIA set-up (€625), the assembling of which requires no more than basic technical facilities. We used such a set-up to test the performance of the proposed spectrometric detector for flow-injection analyses. The tests proved its suitability for real-life applications. The design procedures are also described.
Spectrometric Detector; Tri-Colour Light-Emitting Diode (LED); Flow-Injection Analysis (FIA); Syringe Pump; Low-Budget Instrumentation
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.
Based on partially-coherent digital in-line holography, we report a field-portable microscope that can render lensfree colour images over a wide field-of-view of e.g., >20 mm2. This computational holographic microscope weighs less than 145 grams with dimensions smaller than 17×6×5 cm, making it especially suitable for field settings and point-of-care use. In this lensfree imaging design, we merged a colorization algorithm with a source shifting based multi-height pixel super-resolution technique to mitigate ‘rainbow’ like colour artefacts that are typical in holographic imaging. This image processing scheme is based on transforming the colour components of an RGB image into YUV colour space, which separates colour information from brightness component of an image. The resolution of our super-resolution colour microscope was characterized using a USAF test chart to confirm sub-micron spatial resolution, even for reconstructions that employ multi-height phase recovery to handle dense and connected objects. To further demonstrate the performance of this colour microscope Papanicolaou (Pap) smears were also successfully imaged. This field-portable and wide-field computational colour microscope could be useful for tele-medicine applications in resource poor settings.
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.