Conspicuousness, not colour as foraging cue in plant–animal signalling

The global prevalence of red and black fruits has still not been explained. Hypotheses based on innate consumer preferences have been tested and rejected. Though colour itself plays an important role in animal foraging, it is only one component of signals. Another major component are colour contrasts against background achieving the conspicuousness of signals. In order to evaluate which signal component determines consumers behaviour, we measured fruit colour and colour contrasts of 43 species against their natural background under ambient light conditions. Red and black fruits exhibit stronger contrasts and are therefore more conspicuous to consumers than fruits of other colours. Subsequently, trials were carried out to determine whether colour or conspicuousness influences avian food choice. Four bird species strongly preferred contrasting red–green or black–green over uni-coloured red, green, or black fruit displays, while no preference for particular hues was found. We therefore hypothesize that conspicuousness determines avian food selection and define the contrast hypothesis: Diurnal dispersers select fruit colours based on their conspicuousness and not their colour itself.
Because colour vision is an ancient trait, the entire heterogeneous group of frugivorous birds most likely perceives conspicuousness uniformly over evolutionary time spans. Conspicuousness has thus the potential to explain the global prevalence of red and black fruits.     

The colour of flowers in spectrally variable illumination and insect pollinator vision

The spectral reflectance of differently coloured Australian native plant flowers and foliage was measured and plotted in a colour triangle to represent the colour space of the honeybee. Spectral variations in illumination are shown to significantly change plant colours for bee vision without colour constancy. A model of chromatic adaptation based upon the von Kries coefficient law shows a reduction in plant colour shift, with the degree of correction depending upon position in colour space. A set of artificial reflectances is used to map relative colour shift caused by spectrally variable illumination for the entire colour space of the honeybee. The rarity of some flower colours in nature shows a correlation to a larger colour shift for these colours when illuminated by spectrally variable radiation. The model of chromatic adaptation is applied to illuminations used in a behavioural study on honeybee colour constancy by Neumeyer 1981. Surface colours used by Neumeyer are plotted in colour space for the various illuminations. The results show that an illumination-dependent colour shift correlates to a decrease in the frequency of bees correctly choosing a colour to which it was trained. Accepted: 23 February 1998     

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.     

The evolution of the multicoloured face of mandrills: insights from the perceptual space of colour vision

Multicomponent signals consist of several traits that are perceived as a whole. Although many animals rely on multicomponent signals to communicate, the selective pressures shaping these signals are still poorly understood. Previous work has mainly investigated the evolution of multicomponent signals by studying each trait individually, which may not accurately reflect the selective pressures exerted by the holistic perception of signal receivers. Here, we study the design of the multicoloured face of an Old World primate, the mandrill (Mandrillus sphinx), in relation to two aspects of signalling that are expected to be selected by receivers: conspicuousness and information. Using reflectance data on the blue and red colours of the faces of 34 males and a new method of hue vectorisation in a perceptual space of colour vision, we show that the blue hue maximises contrasts to both the red hue and the foliage background colouration, thereby increasing the conspicuousness of the whole display. We further show that although blue saturation, red saturation and the contrast between blue and red colours are all correlated with dominance, dominance is most accurately indicated by the blue-red contrast. Taken together our results suggest that the evolution of blue and red facial colours in male mandrills are not independent and are likely driven by the holistic perception of conspecifics. In this view, we propose that the multicoloured face of mandrills acts as a multicomponent signal. Last, we show that information accuracy increases with the conspicuousness of the whole display, indicating that both aspects of signalling can evolve in concert.     

Structural colour and iridescence in plants: the poorly studied relations of pigment colour

Background

Colour is a consequence of the optical properties of an object and the visual system of the animal perceiving it. Colour is produced through chemical and structural means, but structural colour has been relatively poorly studied in plants.

Scope

This Botanical Briefing describes the mechanisms by which structures can produce colour. In plants, as in animals, the most common mechanisms are multilayers and diffraction gratings. The functions of structural colour are then discussed. In animals, these colours act primarily as signals between members of the same species, although they can also play roles in camouflaging animals from their predators. In plants, multilayers are found predominantly in shade-plant leaves, suggesting a role either in photoprotection or in optimizing capture of photosynthetically active light. Diffraction gratings may be a surprisingly common feature of petals, and recent work has shown that they can be used by bees as cues to identify rewarding flowers.

Conclusions

Structural colour may be surprisingly frequent in the plant kingdom, playing important roles alongside pigment colour. Much remains to be discovered about its distribution, development and function.     

Digital imaging for colour measurement in ecological research

Traditional methods for colour quantification are complicated by the fact that colours change depending on illumination, and that different observers often perceive colours differently. Here we describe a new affordable method, which improves methods relying on human observers, to quantify patterns and colour variations. The procedure combines customized software with the use of digital cameras and commercial photofinishing software. The computer routines correct unavoidable illumination changes during image capturing, making all images comparable. Colours are quantified in a continuous scale of the conventional colour models developed for the human vision system, such as HSB, RGB, CMYK, or Lab, amenable for statistical analyses. We illustrate the use of this technique showing a previously unknown sexual dimorphism in the red-legged partridge, Alectoris rufa , undetectable with the unaided human eye. We also demonstrate that the digital system provides a finer discrimination than human observers for scoring the plumage of partridges belonging to two different subspecies. This method has potential applications in behavioural ecology, physiology, genetics, evolutionary biology, and taxonomy.     

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.     

Animal colour vision--behavioural tests and physiological concepts

Over a century ago workers such as J. Lubbock and K. von Frisch developed behavioural criteria for establishing that non‐human animals see colour. Many animals in most phyla have since then been shown to have colour vision. Colour is used for specific behaviours, such as phototaxis and object recognition, while other behaviours such as motion detection are colour blind. Having established the existence of colour vision, research focussed on the question of how many spectral types of photoreceptors are involved. Recently, data on photoreceptor spectral sensitivities have been combined with behavioural experiments and physiological models to study systematically the next logical question: ‘what neural interactions underlie colour vision ?‘This review gives an overview of the methods used to study animal colour vision, and discusses how quantitative modelling can suggest how photoreceptor signals are combined and compared to allow for the discrimination of biologically relevant stimuli.     

Local adaptation and divergence in colour signal conspicuousness between monomorphic and polymorphic lineages in a lizard

Population differences in visual environment can lead to divergence in multiple components of animal coloration including signalling traits and colour patterns important for camouflage. Divergence may reflect selection imposed by different receivers (conspecifics, predators), which depends in turn on the location of the colour patch. We tested for local adaptation of two genetically and phenotypically divergent lineages of a rock‐inhabiting lizard, Ctenophorus decresii, by comparing the visual contrast of colour patches to different receivers in native and non‐native environments. The lineages differ most notably in male throat coloration, which is polymorphic in the northern lineage and monomorphic in the southern lineage, but also differ in dorsal and lateral coloration, which is visible to both conspecifics and potential predators. Using models of animal colour vision, we assessed whether lineage‐specific throat, dorsal and lateral coloration enhanced conspicuousness to conspecifics, increased crypsis to birds or both, respectively, when viewed against the predominant backgrounds from each lineage. Throat colours were no more conspicuous against native than non‐native rock but contrasted more strongly with native lichen, which occurs patchily on rocks inhabited by C. decresii. Conversely, neck coloration (lateral) more closely matched native lichen. Furthermore, although dorsal coloration of southern males was consistently more conspicuous to birds than that of northern males, both lineages had similar absolute conspicuousness against their native backgrounds. Combined, our results are consistent with local adaptation of multiple colour traits in relation to multiple receivers, suggesting that geographic variation in background colour has influenced the evolution of lineage‐specific coloration in C. decresii.