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.     

A simple method for measuring colour in wild animals: validation and use on chest patch colour in geladas (Theropithecus gelada)

Adaptive hypotheses about colour variation are widespread in behavioural ecology, and several methods of objective colour assessment have been proposed and validated for use in a wide variety of taxa. However, to date, the most objective and reliable methods of assessing colour are not readily applied to wild animals. In the present study, we present a simple method for assessing colour in unrestrained, wild subjects using digital photography. The method we describe uses a digital camera, a colour standard, and colour analysis software, and can be used to measure any part of the visible colour spectrum. We demonstrate that the method: (1) is accurate and precise across different light conditions; (2) satisfies previous criteria regarding linearity and red, green, and blue equality; and (3) can be independently validated visually. In contrast with previous digital methods, this method can be used under natural light conditions and can be readily applied to subjects in their natural habitat. To illustrate this, we use the method to measure chest colour in wild geladas ( Theropithecus gelada ). Unique among primates, geladas have a red patch of skin on their chest and neck, which, for males, is thought to be a sexually selected signal. Offering some support to this hypothesis, we found differences in chest 'redness' for males across different age groups, with males in their reproductive prime exhibiting the reddest chests.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 94 , 231–240.     

Opponent colour coding is a universal strategy to evaluate the photoreceptor inputs in Hymenoptera

Summary Behavioural tests were carried out with 9 hymenopteran insect species, which ranked certain sets of coloured stimuli according to their subjective similarity to a previously memorized stimulus. Kendall's coefficient is employed for the analysis of correlation between these similarity rankings and the colour distance rankings predicted by various models of neural colour computation. The models are based on the measured spectral sensitivities of photoreceptor colour types and use a variety of simple colour coding systems to derive hypothetical colour distances. The correlation between the predictions of the models and the behavioural results serves as a measure for the likelihood of existence of a colour coding system. In all species, the similarity rankings can be best explained by assuming that colour is coded on a perceptual level by two colour opponent mechanisms. Brightness differences are ignored, indicating that an intensity-coding sub-system is not used in colour discrimination by the insects investigated. The weighting factors of the colour opponent mechanisms differ between species in detail, but not in the principles involved. It is thus possible to employ a standard measure of perceptual colour distance (colour hexagon distance) to predict the capacities of colour discrimination adequately in all the tested insects.     

High sensitivity to short wavelengths in a lizard and implications for understanding the evolution of visual systems in lizards

Progress in developing animal communication theory is frequently constrained by a poor understanding of sensory systems. For example, while lizards have been the focus of numerous studies in visual signalling, we only have data on the spectral sensitivities of a few species clustered in two major clades (Iguania and Gekkota). Using electroretinography and microspectrophotometry, we studied the visual system of the cordylid lizard Platysaurus broadleyi because it represents an unstudied clade (Scinciformata) with respect to visual systems and because UV signals feature prominently in its social behaviour. The retina possessed four classes of single and one class of double cones. Sensitivity in the ultraviolet region (UV) was approximately three times higher than previously reported for other lizards. We found more colourless oil droplets (associated with UV-sensitive (UVS) and short wavelength-sensitive (SWS) photoreceptors), suggesting that the increased sensitivity was owing to the presence of more UVS photoreceptors. Using the Vorobyev-Osorio colour discrimination model, we demonstrated that an increase in the number of UVS photoreceptors significantly enhances a lizard's ability to discriminate conspecific male throat colours. Visual systems in diurnal lizards appear to be broadly conserved, but data from additional clades are needed to confirm this.     

Polarization-based brightness discrimination in the foraging butterfly, Papilio xuthus

The human eye is insensitive to the angular direction of the light e-vector, but several animal species have the ability to discriminate differently polarized lights. How the polarization is detected is often unclear, however. Egg-laying Papilio butterflies have been shown to see false colours when presented with differently polarized lights. Here we asked whether this also holds in foraging butterflies. After training individuals to feed on nectar in front of an unpolarized spectral light, we carried out three dual-choice tests, where the discrimination of (i) the spectral content, (ii) the light intensity, and (iii) the e-vector orientation were investigated. In the first test, the butterflies selected the trained spectrum irrespective of its intensity, and in the second test they chose the light with the higher intensity. The result of the e-vector discrimination test was very similar to that of the second test, suggesting that foraging butterflies discriminate differently polarized lights as differing in brightness rather than as differing in colour. Papilio butterflies are clearly able to use at least two modes of polarization vision depending on the behavioural context.     

Blue–green opponency and trichromatic vision in the greenhouse whitefly (Trialeurodes vaporariorum) explored using light emitting diodes

Visual orientation in the greenhouse whitefly (Trialeurodes vaporariorum Westwood, Hemiptera: Aleyrodidae) is the result of “wavelength‐specific behaviours.” Green–yellow elicits “settling behaviour” while ultraviolet (UV) radiation initiates “migratory behaviour.” The only available physiological study of the photoreceptors' spectral efficiency showed peaks in the green and the UV range and whitefly vision was said to be dichromatic so far. In order to study the visual behaviour of T. vaporariorum, 19 narrow‐bandwidth light emitting diodes (LEDs) covering the UV‐A and visible range were used in combination with light scattering acrylic glass screens in a small‐scale choice arena under greenhouse conditions. Multiple‐choice and dual‐choice assays were performed, resulting in LED‐based behavioural action spectra of settling (green) and migratory behaviour (UV). A potential inhibitory blue–green chromatic mechanism was studied by combining yellow with different bluish LEDs. Intensity dependencies were illustrated by changing LED intensities. Regarding the “settling response,” highest attraction was achieved by a green LED with a centroid wavelength of 550 nm, while a blue LED with 469 nm proved to be most inhibitory. Besides this inhibitory interaction, an intensity dependence was observed within the action spectrum in the green–yellow range. “Migratory behaviour” was elicited the most by the UV LED with the shortest available wavelength of 373 nm. The results provide compelling behavioural evidence for the presence of a green and a yet undescribed blue sensitive photoreceptor and a blue–green opponent mechanism. Furthermore, empirical colour choice models were built and receptor peaks were estimated around 510–520 nm (green), 480–490 nm (blue) and 340–370 nm (UV). Consequently, a trichromatic receptor setup is suggested for T. vaporariorum.     

Behavioural evidence for colour vision in stomatopod crustaceans

If an organism can be taught to respond in a particular way to a wavelength of light, irrespective of that light's intensity, then it must be able to perceive the colour of the stimulus. No marine invertebrate has yet been shown to have colour vision. Stomatopod crustaceans (mantis shrimps) are colourful animals and their eyes have many adaptations which indicate that they are capable of such spectral analysis. We adopted an associative learning paradigm to attempt to demonstrate colour vision. Stomatopods readily learnt to choose some colours from arrays of greys, even when the correct choice colours were darker than the ones they had been trained to. Possible mechanisms underlying colour vision in these animals, and their ecological significance are discussed. A simple model is presented which may help interpret the complex-stomatopod colour vision system and explain some of the learning anomalies.Abbreviations ND neutral density - OD optical density - R8 Retinular cell 8 - R1–7 Retinular cells 1–7 - R1D Distally placed R1–7 retinular cells in mid-band row 1 - e.g. R1P Proximally placed R1–7 retinular cells in mid-band row 1 - D/P Estimate of chromatic signal ratio     

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.     

Rhabdom evolution in butterflies: insights from the uniquely tiered and heterogeneous ommatidia of the Glacial Apollo butterfly, Parnassius glacialis

The eye of the Glacial Apollo butterfly, Parnassius glacialis, a 'living fossil' species of the family Papilionidae, contains three types of spectrally heterogeneous ommatidia. Electron microscopy reveals that the Apollo rhabdom is tiered. The distal tier is composed exclusively of photoreceptors expressing opsins of ultraviolet or blue-absorbing visual pigments, and the proximal tier consists of photoreceptors expressing opsins of green or red-absorbing visual pigments. This organization is unique because the distal tier of other known butterflies contains two green-sensitive photoreceptors, which probably function in improving spatial and/or motion vision. Interspecific comparison suggests that the Apollo rhabdom retains an ancestral tiered pattern with some modification to enhance its colour vision towards the long-wavelength region of the spectrum.     

Frequency-dependent physiological trade-offs between competing colour morphs

Evolutionary theory suggests that alternative colour morphs (i.e. genetically controlled phenotypes) may derive similar fitness under frequency-dependent selection. Here we experimentally demonstrate opposing effects of frequency-dependent social environments on plasma hormone levels (testosterone and corticosterone) and immune function between red- and black-headed male morphs of the Gouldian finch (Erythrura gouldiae). Red-headed males are highly sensitive to changes in the social environment, especially towards the relative density of their own aggressive morph, exhibiting high stress responses and immunosuppression in socially competitive environments. In contrast, the non-aggressive black-headed males follow a more passive strategy that appears to buffer them against social stresses. The differential effect of hormones on aggressive behaviour and immune performance reinforces the contrasting behavioural strategies employed by these colour morphs, and highlights the importance of the social environment in determining the individual basis of behavioural and physiological responses.     

Evolution of colour vision in mammals

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.     

Background colouration influences body colour segregation in mollies

When given a choice of swimming against a black background near an empty black end compartment or swimming against a white background near an empty white compartment, black morph mollies Poecilia latipinna spent significantly more time against the black background. The presence of a shoal (either white or black mollies) in the black compartment increased the preference of the test fish for the black background. However, black molly test fish spent significantly less time against a black background if a black shoal was present in the white compartment than in assays in which a white shoal was in the white compartment. In assays where test fish were given the choice between two black shoals, test fish spent significantly more time near the black shoal on the black background. Similarly, when given the choice between two white shoals, test fish spent significantly more time against the black background. The results suggest that the presence of cryptic backgrounds alter shoalmate choices in fish that assort according to body colouration.     

Avian colour vision and avian video playback experiments

Video playback potentially allows the presentation, manipulation, and replication of realistic moving visual stimuli, in a way that is impossible with real animals or static dummies, and difficult even with mechanical models. However, there are special problems attached to the use of this technology; this article concentrates on the problem of accurate colour rendition. Video and television simulate the colour of objects rather than reproduce the spectrum of light that they naturally emit, transmit, or reflect. This simulation is achieved by using relatively narrow waveband light to stimulate the cone cells in the retina in a similar pattern to that produced by the natural object. However, species differ in the spectral tuning of their photoreceptors, so a faithful colour rendition for a human is unlikely to be achieved for another species. This problem is discussed with special reference to birds, a taxon renown for its colourfulness and frequent use in behavioural experiments but which has a very different colour vision from that of humans. We stress that the major pitfalls that can arise when using video playback with avian subjects can also occur in ’normal’ behavioural experiments. However, the problems of faithful colour rendition are particularly severe with video, and the major benefits that the technology brings will only be realised under a limited range of circumstances, with careful validation experiments. Received: 24 January 2000 / Received in revised form: 28 March 2000 / Accepted: 1 April 2000     

The fiddler crab Uca mjoebergi uses colour vision in mate choice

Although the role of colour in mate choice is well known, few tests of colour vision have been based on mating behaviour. Females of the fiddler crab Uca mjoebergi have recently been shown to use claw coloration to recognize conspecific males. In this study I demonstrate that the females use colour vision for this task; preferentially approaching yellow claws over grey claws regardless of their intensity while failing to discriminate between yellow claws differing in intensity. This is one of only a handful of studies confirming the involvement of colour vision in mate choice and the first conclusive evidence in fiddler crabs.     

The effect of environmental colour on the growth,metabolism, physiology and skin pigmentation of the carnivorous freshwater catfish Lophiosilurus alexandri

The growth, physiology and skin pigmentation of pacamã Lophiosilurus alexandri juveniles were evaluated in an experiment using different tank colours (white, yellow, green, blue, brown and black) over an 80 day period. The tank colours did not cause significant differences to final body mass, total length, survival rate, carcass composition (moisture, crude protein, ash, ether extract, calcium, phosphorus, energy), or to plasma protein, triglyceride and cholesterol values. Haematocrit values, however, were highest for fish kept in white tanks (ANOVA P < 0·05), while the greatest haemoglobin levels were recorded for fish kept in blue and brown tanks (P < 0·01). The concentrations of cortisol (P < 0·001) and glucose (P < 0·01) were the most in fish in the black tanks. Tank colour affected skin pigmentation significantly, with fish in white tanks having the highest values of L* (brightness) and the lowest values in blue and black tanks. L*, however, decreased in all treatments throughout the experiment. C*_ab increased significantly over the course of the experiment in fish kept in white tanks. Similar increases of C*_ab were recorded in the other treatments but to a lesser extent. The use of black tanks during the cultivation of L. alexandri caused stress and should be avoided. Cultivation in white and yellow tanks produced individuals with a pale skin colour, while cultivation in blue and black tanks resulted in juveniles with a darker and more pigmented skin.     

Defective colour vision and taste sensitivity to PTC in the Maharashtrian population

Three caste groups of the Maharashtrian population, namely Brahmin (N=58), Maratha (N=989) and Scheduled caste (N=1073), were studied for defective colour vision and for the ability to taste phenylthiocarbamide (PTC). Comparisons of defective colour vision and PTC taste sensitivity were made with other Maharashtrian populations studied previously by various authors. The incidence of defective colour vision is lowest among the Brahmins (3.44%) and highest in the Scheduled caste people (4.28%). Taste sensitivity to PTC is highest among the Brahmins (72.5%) with a value which is close to the Vednagar Brahmins (73.3%) who show the highest frequency of the ‘T’ gene recorded so far in Maharashtra.     

The colour line and the colour scale in the twentieth century

Some more recent evidence supports Du Bois’ prediction that the twentieth century would prove the century of the colour line. It indicates that men have always and everywhere shown a preference for fair complexioned women as sexual partners, whereas males seeking a mate are rarely disadvantaged by a dark complexion. In the employment market in the USA, a dark complexion is a significant disadvantage for both males and females. Though there is no properly comparable evidence from other countries, there appears to be a widespread tendency for any negative valuation of darker skin colour to be incorporated into a scale of socio-economic status. In some situations a colour scale is replacing the colour line.

Du Bois’ reference to differences of colour has been largely superseded in English-speaking countries by references to differences of race. From a policy standpoint, the switch from colour to race has had both positive and negative consequences. From a sociological standpoint, it has made it more difficult to disaggregate the dimensions of social difference and to dispel the confusions engendered by ideas of racial difference.     

Evolution of the visual sensory system in cichlid fishes from crater lake Barombi Mbo in Cameroon

In deep‐water animals, the visual sensory system is often challenged by the dim‐light environment. Here, we focus on the molecular mechanisms involved in rapid deep‐water adaptations. We examined visual system evolution in a small‐scale yet phenotypically and ecologically diverse adaptive radiation, the species flock of cichlid fishes in deep crater lake Barombi Mbo in Cameroon, West Africa. We show that rapid adaptations of the visual system to the novel deep‐water habitat primarily occurred at the level of gene expression changes rather than through nucleotide mutations, which is compatible with the young age of the radiation. Based on retinal bulk RNA sequencing of all eleven species, we found that the opsin gene expression pattern was substantially different for the deep‐water species. The nine shallow‐water species feature an opsin palette dominated by the red‐sensitive (LWS) opsin, whereas the two unrelated deep‐water species lack expression of LWS and the violet‐sensitive (SWS2B) opsin, thereby shifting the cone sensitivity to the centre of the light spectrum. Deep‐water species further predominantly express the green‐sensitive RH2Aα over RH2Aβ. We identified one amino acid substitution in the RH2Aα opsin specific to the deep‐water species. We finally performed a comparative gene expression analysis in retinal tissue of deep‐ vs. shallow‐water species. We thus identified 46 differentially expressed genes, many of which are associated with functions in vision, hypoxia management or circadian clock regulation, with some of them being associated with human eye diseases.     

Diurnal lighting patterns and habitat alter opsin expression and colour preferences in a killifish

Spatial variation in lighting environments frequently leads to population variation in colour patterns, colour preferences and visual systems. Yet lighting conditions also vary diurnally, and many aspects of visual systems and behaviour vary over this time scale. Here, we use the bluefin killifish (Lucania goodei) to compare how diurnal variation and habitat variation (clear versus tannin-stained water) affect opsin expression and the preference to peck at different-coloured objects. Opsin expression was generally lowest at midnight and dawn, and highest at midday and dusk, and this diurnal variation was many times greater than variation between habitats. Pecking preference was affected by both diurnal and habitat variation but did not correlate with opsin expression. Rather, pecking preference matched lighting conditions, with higher preferences for blue at noon and for red at dawn/dusk, when these wavelengths are comparatively scarce. Similarly, blue pecking preference was higher in tannin-stained water where blue wavelengths are reduced. In conclusion, L. goodei exhibits strong diurnal cycles of opsin expression, but these are not tightly correlated with light intensity or colour. Temporally variable pecking preferences probably result from lighting environment rather than from opsin production. These results may have implications for the colour pattern diversity observed in these fish.     

Plant scents modify innate colour preference in foraging swallowtail butterflies

Flower-visiting insects exhibit innate preferences for particular colours. A previous study demonstrated that naive Papilio xuthus females prefer yellow and red, whereas males are more attracted to blue. Here, we demonstrate that the innate colour preference can be modified by olfactory stimuli in a sexually dimorphic manner. Naive P. xuthus were presented with four coloured discs: blue, green, yellow and red. The innate colour preference (i.e. the colour first landed on) of the majority of individuals was blue. When scent from essential oils of either orange flower or lily was introduced to the room, females’ tendency to select the red disc increased. Scents of lavender and flowering potted Hibiscus rosa-sinensis, however, were less effective. Interestingly, the odour of the non-flowering larval host plant, Citrus unshiu, shifted the preference to green in females. In males, however, all plant scents were less effective than in females, such that blue was always the most favoured colour. These observations indicate that interactions between visual and olfactory cues play a more prominent role in females.