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.     

Visual modelling suggests a weak relationship between the evolution of ultraviolet vision and plumage coloration in birds

Birds have sophisticated colour vision mediated by four cone types that cover a wide visual spectrum including ultraviolet (UV) wavelengths. Many birds have modest UV sensitivity provided by violet‐sensitive (VS) cones with sensitivity maxima between 400 and 425 nm. However, some birds have evolved higher UV sensitivity and a larger visual spectrum given by UV‐sensitive (UVS) cones maximally sensitive at 360–370 nm. The reasons for VS–UVS transitions and their relationship to visual ecology remain unclear. It has been hypothesized that the evolution of UVS‐cone vision is linked to plumage colours so that visual sensitivity and feather coloration are ‘matched’. This leads to the specific prediction that UVS‐cone vision enhances the discrimination of plumage colours of UVS birds while such an advantage is absent or less pronounced for VS‐bird coloration. We test this hypothesis using knowledge of the complex distribution of UVS cones among birds combined with mathematical modelling of colour discrimination during different viewing conditions. We find no support for the hypothesis, which, combined with previous studies, suggests only a weak relationship between UVS‐cone vision and plumage colour evolution. Instead, we suggest that UVS‐cone vision generally favours colour discrimination, which creates a nonspecific selection pressure for the evolution of UVS cones.     

Contrast versus colour in aposematic signals

Conspicuousness is an important feature of warning coloration. One hypothesis for its function is that it increases signal efficacy by facilitating avoidance learning. An alternative, based on the handicap hypothesis, suggests that the degree of conspicuousness holds information directly about the quality of the prey, and that predators associate and learn about the conspicuousness of the coloration, and not the actual colour pattern. We studied the relative importance of signal contrast and the colours of signals for predator attention during discrimination. We used young chicks, Gallus gallus domesticus, as predators and small blue or red paper cones on either matching or contrasting paper backgrounds as stimuli associated with palatable or unpalatable chick crumbs. In four treatment groups, birds could use either cone and/or background colour, cone colour only, background colour only or cone-to-background contrast as cues for discrimination. Only birds in the contrast treatment failed to learn their discrimination task. Birds that had a choice between cone and background colour as cues used the cone colour and they learned the task faster than did birds that had to use background colour as a cue. The results suggest that birds primarily attend to the colours of signals and disregard contrast in discrimination tasks; they thus fail to support a handicap function of conspicuous aposematic coloration. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

What influences aggression and foraging activity in social birds? Measuring individual,group and environmental characteristics

For specialised feeders, accessing food resources may impact on the performance of appetitive foraging and social behaviours at individual and population levels. Flamingos are excellent examples of social species with complex, species-specific feeding strategies. As attainment of coloured plumage depends upon intake of dietary carotenoids, and as study of free-ranging flamingos shows that foraging is disrupted by aggression from other birds, we investigated the effect of four feeding styles on foraging and aggression in captive lesser flamingos. We evaluated individual and group differences in foraging and aggression when birds consumed bespoke “flamingo pellet” from a bowl, an indoor feeding pool and an outdoor feeding section of their pool. Natural foraging (when birds were feeding irrespective of the presence of pellet) was recorded for comparison with artificial feeding styles. One-minute long video footage of the birds' activities in these different locations, recorded between 2013 and 2016, was used to evaluate behaviour. Total number of seconds engaged in feeding and in aggression was recorded by continuous sampling. The colour of individual birds was scored from 1 (mainly white) to 4 (mainly pink). For natural filter feeding in the outdoor pool, maximum foraging was twice as much as bowl feeding, whilst aggression was less than half as much as other feeding methods. Overall, a more restricted feeding style significantly predicted aggression, along with increasing group size. Plumage colour significantly influenced aggression (brightest flamingos were more aggressive) and showed a non-significant trend with foraging (brighter birds fed less than paler birds). No sex effect on feeding or aggression was found. This study enhances our understanding of husbandry and species' biology impacts on captive behaviour and provides data-based evidence to improve food presentation. For flamingos, implementation of spacious outdoor feeding areas can encourage natural foraging patterns by reducing excess aggression and enhances welfare by improving flock social stability.     

Noradrenaline and selective attention

Rats depleted of forebrain noradrenaline by intracerebral injection of four micrograms of 6-hydroxydopamine into the dorsal noradrenergic bundle were examined on their ability to ignore irrelevant stimuli. In the latent inhibition paradigm normal rats were pre-exposed to visual and auditory stimuli in the absence of reward and such pre-exposure was found to slow subsequent learning of a successive discrimination task using these stimuli. Noradrenaline depletion blocked the usual latent inhibition effect, thus suggesting that the lesioned animals were impaired in ignoring irrelevant stimuli. A second paradigm, a nonreversal shift, involved training rats on a two-dimension discrimination task with one dimension relevant and the other irrelevant. Nonreversal shift (in which the initially irrelevant dimension became the sole relevant one) was significantly improved by 6-hydroxydopamine lesion. It is thus concluded that strong evidence has been presented in favour of a role for the dorsal noradrenergic bundle in attentional filtering processes.     

Spatial, colour and contrast response characteristics of mechanisms which mediate discrimination of pattern orientation and magnification

We have measured response times for the detection of a single target presented against a set of reference elements which are characterised by combinations of four different stimulus parameters; colour, contrast polarity, magnification and orientation. The aim of the experiments was to determine the response characteristics of visual mechanisms which mediate target detection through the discrimination of orientation and magnification. In the first experiments, we determined sensitivity to differences in colour and contrast polarity, and show that the mechanisms responsible for the discrimination of orientation and of magnification are both selective in their responses to colour and to contrast polarity. There are, nonetheless, residual interactions between patterns of different contrast polarities and between those of different colour, and in the latter case, weak interactions persist under equiluminance conditions. In a second set of experiments, we examined the interactions between orientation and magnification. We conclude that the responses of visual mechanisms which mediate target detection through discrimination of orientation are markedly dependent on stimulus magnification whereas those which mediate detection through discrimination of magnification are, in contrast, relatively insensitive to stimulus orientation.     

Spatial orientation in Japanese quails (Coturnix coturnix japonica)

Finding a given location can be based on a variety of strategies, for example on the estimation of spatial relations between landmarks, called spatial orientation. In galliform birds, spatial orientation has been demonstrated convincingly in very young domestic chicks. We wanted to know whether adult Japanese quails (Coturnix coturnix japonica) without food deprivation are also able to use spatial orientation. The quails had to learn the relation of a food location with four conspicuous landmarks which were placed in the corners of a square shaped arena. They were trained to find mealworms in three adjacent food cups in a circle of 20 such cups. The rewarded feeders were located during training between the same two landmarks each of which showed a distinct pattern. When the birds had learned the task, all landmarks were displaced clockwise by 90 degrees. When tested in the new situation, all birds redirected their choices with respect to the landmark shift. In subsequent tests, however, the previously correct position was also chosen. According to our results, quails are using conspicuous landmarks as a first choice for orientation. The orientation towards the previously rewarded location, however, indicates that the neuronal representation of space which is used by the birds also includes more fine grain, less conspicuous cues, which are probably also taken into account in uncertain situations. We also presume that the rare orientation towards never rewarded feeders may be due to a foraging strategy instead of being mistakes.     

Analysis from avian visual perspective reveals plumage colour differences among females of capuchino seedeaters (Sporophila)

Females of the closely related capuchino seedeaters are difficult to distinguish from one another based on human visual perception of colouration and morphology. We examined plumage colour differences among females of four species, the tawny‐bellied seedeater Sporophila hypoxantha, the dark‐throated seedeater S. ruficollis, the rufous‐rumped seedeater S. hypochroma, and the chesnut‐bellied seedeater S. cinnamomea. Reflectance values were measured on museum skins, and interspecific differences were analyzed using the Vorobyev‐Osorio avian colour discrimination model. Interspecific distances in the colour space calculated by the model were considerably higher than the threshold for colour discrimination, indicating the presence of colour differences among species that should be detectable by birds. Colour differences between S. hypoxantha and the other three species were the largest. A Discriminant Function Analysis showed that UV‐wavelength was particularly important in species separation. Our results indicate that female plumage of these four species is considerably divergent in colour; these differences being imperceptible to the human eye, thus representing previously unknown morphological evolution in these species.     

Selective stimulus control in discrimination of lateral mirror images by pigeons

Four monocularly and two binocularly viewing pigeons were trained to peck a key when it displayed one stimulus (S+) but not to peck when it displayed another stimulus (S−). S+ and S− were a lateral mirror-image pair of two-coloured stimuli. When tested for transfer with the untrained eye open, two of the monocular birds pecked more during S− than S+, the other two continuing to favour S+. During generalization tests on the wavelength dimension all monocular birds pecked much more often during one S+ colour than during the other. The colour controlling pecking was that displayed on the side of the key facing the open eye during S+ presentations. Both binocular birds developed asymmetrical responses to the key, one favouring the left, the other the right side of the key. Generalization tests on the wavelength dimension showed selective control by the colour displayed on the favoured side of the key during S+ presentations. The results are interpreted as supporting the view that pigeons learn to discriminate lateral mirror images by developing asymmetrical observing responses that convert the left-right difference between the mirror images into a difference more easily discriminable.     

Iris colour of American kestrels varies with age,sex, and exposure to PCBs

For most species of birds, little is known about the pattern and significance of intraspecific variation in iris colour. In early winter, captive American kestrels could be subjectively placed into at least two age categories: 0.5-yr-old birds had all-brown irides, whereas those of older birds were red-brown. As part of a toxicological study on kestrels, we quantified iris colour objectively using a digital camera to examine potential variation due to age, sex, and exposure to polychlorinated biphenyls (PCBs). Red, green, and blue values, plus an overall measure of colour using principal components analysis (PC1), were derived for breeding and nonbreeding kestrels 1.5-6.5 yr old (fed PCBs), plus offspring 0.5 yr old (exposed to PCBs only in ovo). Age category (0.5, 1.5, and 2.5+ yr) and PCB exposure consistently had an effect on colour, while sex was significant only for red and almost so for PC1. ANOVA with age as a covariate revealed that the amount of red continued to increase throughout life, but PCBs suppressed the development of that colour.     

Hoverflies are imperfect mimics of wasp colouration

Many Batesian mimics are considered to be inaccurate copies of their models, including a number of hoverfly species which appear to be poor mimics of bees and wasps. This inaccuracy is surprising since more similar mimics are expected to deceive predators more frequently and therefore have greater survival. One suggested explanation is that mimics which appear inaccurate to human eyes may be perceived differently by birds, the probable agents of selection. For example, if patterns contain an ultra-violet (UV) component, this would be visible to birds but overlooked by humans. So far, indirect comparisons have been made using human and bird responses to mimetic stimuli, but direct colour measurements of mimetic hoverflies are lacking. We took spectral readings from a wide range of hoverfly and wasp patterns. They show very low reflectance in the UV range, and do not display any human-invisible colour boundaries. We modelled how the recorded spectra would be perceived by both birds and humans. While colour differences between wasps and hoverflies are slightly more distinct according to human visual abilities, bird vision is capable of discriminating the two taxa in almost all cases. We discuss a number of factors that might make the discrimination task more challenging for a predator in the field, which could explain the apparent lack of selection for accurate colour mimicry.     

Avian colour perception predicts behavioural responses to experimental brood parasitism in chaffinches

Hosts of cuckoos have evolved defences allowing them to discriminate and reject parasite eggs. Mechanisms of discrimination are mostly visually mediated, and have been studied using approaches that do not account for what the receiver (i.e. host) actually can discriminate. Here, for the first time we apply a perceptual model of colour discrimination to study behavioural responses to natural variation in parasite egg appearance in chaffinches Fringilla coelebs. Discrimination of parasite eggs gradually increased with increasing differences in chromatic contrasts as perceived by birds between parasite and host eggs. These results confirm that colour differences of the eggs as perceived by birds are important integral parts of a matching signal used by chaffinch hosts.     

Intradimensional and extradimensional shifts in conditional discrimination

Two groups of pigeons learned a two key conditional discrimination. Color was the conditional stimulus and form the choice stimulus for subjects in one group. Form was the conditional stimulus and color the choice stimulus in the other group. Half the subjects in each group then underwent an intradimensional shift: The conditional stimulus dimension and choice stimulus dimension were unchanged but the correct and incorrect stimulus compounds were reversed. The remaining subjects underwent an extra-dimensional shift in which the conditional stimulus dimension and choice stimulus dimension were reversed. Subjects which experienced an intradimensional shift learned the new conditional discrimination more quickly. It was concluded that subjects followed rules to solve conditional discriminations but also learned the functions of each stimulus dimension.     

Sunset and the orientation of European robins (Erithacus rubecula)

The migratory orientation of European robins (Erithacus rubecula) in autumn was tested immediately after sunset and also after the beginning of astronomical darkness. In twilight tests under clear skies, the birds selected an appropriate migratory direction. During the course of autumn, along with the shift of sunset azimuth, the orientation of birds also shifted, always in a counter-clockwise direction. Although this shift of orientation was not statistically significant, the difference between the mean direction and the sunset was the same for each autumn period. This suggests that the migratory direction was selected on the basis of menotactic orientation re the setting sun. Random directions were observed under solid overcast skies as well as during tests under starry skies, begun after all trace of the sunset position had disappeared.     

Bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>) sacrifice foraging speed to solve difficult colour discrimination tasks

The performance of individual bumblebees at colour discrimination tasks was tested in a controlled laboratory environment. Bees were trained to discriminate between rewarded target colours and differently coloured distractors, and then tested in non-rewarded foraging bouts. For the discrimination of large colour distances bees made relatively fast decisions and selected target colours with a high degree of accuracy, but for the discrimination of smaller colour distances the accuracy decreased and the bees response times to find correct flowers significantly increased. For small colour distances there was also significant linear correlations between accuracy and response time for the individual bees. The results show both between task and within task speed-accuracy tradeoffs in bees, which suggests the possibility of a sophisticated and dynamic decision-making process.     

Mechanisms of dusk orientation in white-throated sparrows (Zonotrichia albicollis): Clock-shift experiments

Summary Several species of night migrating birds, especially North American emberizines, exhibit markedly different orientation behaviour when tested in circular cages under clear skies at dusk as compared with tests performed after complete darkness. During the period between sunset and the first appearance of stars, birds tend to show high levels of well-oriented hopping; birds deprived of exposure to clear skies at dusk hop less and their activity is usually not oriented. There is evidence that visual cues available during the dusk period, but not later, are responsible for this difference, but details of the orientation mechanisms involved are unclear. We performed 3-h fast and slow clock shifts on white-throated sparrows (Zonotrichia albicollis) to address two questions concerning migratory orientation at dusk: (1) Is the better orientation of sparrows tested at dusk a function of the visual cues available at that time, or does it result from circadian changes in motivation?; and (2) Is the dusk orientation based on a time-compensated sun compass?Sparrows subjected to a 3-h slow clock shift were tested with controls on clear, moonless nights beginning immediately after lights-off in the clock shift room and thus about 3.5 h after local sunset. Individuals of both groups performed poorly oriented hopping typical of tests performed after complete darkness. The pooled data from each group were not significantly oriented. These results show that the visual cues available shortly after sunset, not temporal changes in the motivation of the birds, are responsible for the qualitative differences in orientation.Birds exposed to a 3-h fast clock shift were tested with controls on clear evenings between sunset and the first appearance of stars. Both groups showed well-oriented hopping. The mean direction of the pooled tests of controls was 325°, a typical spring orientation direction for this species. The mean direction of the pooled tests of the clock shifted birds (274°) was significantly different from that of controls and the 51° counterclockwise shift is consistent with that predicted by a time-compensated sun compass model.     

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

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

Predator-specific camouflage in chameleons

A crucial problem for most animals is how to deal with multiple types of predator, which differ in their sensory capabilities and methods of prey detection. For animals capable of rapid colour change, one potential strategy is to change their appearance in relation to the threat posed by different predators. Here, we show that the dwarf chameleon, Bradypodion taeniabronchum, exhibits different colour responses to two predators that differ in their visual capabilities. Using a model of animal colour perception to gain a 'predator's eye view', we show that chameleons showed better background colour matching in response to birds than snakes, yet they appear significantly more camouflaged to the snake visual system because snakes have poorer colour discrimination.     

Comparative psychophysics of bumblebee and honeybee colour discrimination and object detection

Bumblebee (Bombus terrestris) discrimination of targets with broadband reflectance spectra was tested using simultaneous viewing conditions, enabling an accurate determination of the perceptual limit of colour discrimination excluding confounds from memory coding (experiment 1). The level of colour discrimination in bumblebees, and honeybees (Apis mellifera) (based upon previous observations), exceeds predictions of models considering receptor noise in the honeybee. Bumblebee and honeybee photoreceptors are similar in spectral shape and spacing, but bumblebees exhibit significantly poorer colour discrimination in behavioural tests, suggesting possible differences in spatial or temporal signal processing. Detection of stimuli in a Y-maze was evaluated for bumblebees (experiment 2) and honeybees (experiment 3). Honeybees detected stimuli containing both green-receptor-contrast and colour contrast at a visual angle of approximately 5 degrees , whilst stimuli that contained only colour contrast were only detected at a visual angle of 15 degrees . Bumblebees were able to detect these stimuli at a visual angle of 2.3 degrees and 2.7 degrees , respectively. A comparison of the experiments suggests a tradeoff between colour discrimination and colour detection in these two species, limited by the need to pool colour signals to overcome receptor noise. We discuss the colour processing differences and possible adaptations to specific ecological habitats.     

Nonelemental visual learning in honeybees

Free-flying honeybees, Apis mellifera, learn visual stimuli in the appetitive context of food search. Visual compound stimuli are relevant in nautre as bees learn flower images that consist of many visual elements. We studied whether elemental associations between each visual element and the reinforcement (elemental approach) are enough to explain the solving of visual discrimination problems that raise ambiguity at the elemental level. We asked whether bees could solve three different visual discriminations: (1) positive patterning (A−, B−, AB+); (2) negative patterning (A+, B+, AB−); and (3) biconditional discrimination (AB+, CD+, AC−, BD−). In experiments 1 and 2 bees had to discriminate a yellow-violet chequerboard from the yellow or the violet squares alone. In experiment 3, four different gratings combining one colour (yellow or violet) with one orientation (vertical or horizontal) had to be discriminated. In all three problems binary compounds were trained in such a way that each element appeared equally often as rewarded and nonrewarded. Bees could solve the three discrimination problems. They always chose the reinforced stimulus despite ambiguity at the level of the elements. For solving positive patterning, elemental processing could be used. For negative patterning and biconditional discrimination, nonelemental processing strategies (unique-cue or configural approach) are necessary to account for these results. Although we cannot decide between a configural and a unique-cue interpretation, we can clearly reject purely elemental processing in these cases. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.