The modelling of flower colour: spectral purity or colour contrast as biologically relevant descriptors of flower colour signals for bees depending upon the perceptual task.

Flower colour is an important mediator of plant-pollinator interactions. While the reflectance of light from the flower surface and background are governed by physical properties, the perceptual interpretation of such information is generated by complex multilayered visual processing. Should quantitative modelling of flower signals strive for repeatable consistency enabled by parameter simplification, or should modelling reflect the dynamic way in which bees are known to process signals? We discuss why colour is an interpretation of spectral information by the brain of an animal. Different species, or individuals within a species, may respond differently to colour signals depending on sensory apparatus and/or individual experience. Humans and bees have different spectral ranges, but colour theory is strongly rooted in human colour perception and many principles of colour vision appear to be common. We discuss bee colour perception based on physiological, neuroanatomical and behavioural evidence to provide a pathway for modelling flower colours. We examine whether flower petals and floral guides as viewed against spectrally different backgrounds should be considered as a simple colour contrast problem or require a more dynamic consideration of how bees make perceptual decisions. We discuss that plants such as deceptive orchids may present signals to exploit bee perception, whilst many plants do provide honest signalling where perceived saturation indicates the probability of collecting nutritional rewards towards the centre of a flower that then facilitates effective pollination.

Flower colour and size signals differ depending on geographical location and altitude region.

Bees are major pollinators of angiosperms and have phylogenetically conserved colour vision but differ in how various key species use achromatic information that is vital for both flower detection and size processing. We modelled green contrast and colour contrast signals from flowers of different countries where there are well established differences in availability of model bee species along altitudinal gradients. We tested for consistency in visual signals as expected from generalization in pollination principles using phylogenetically informed linear models. Patterns of chromatic contrast, achromatic green contrast and flower size differed among the three floras we examined. In Nepal there is a significant positive correlation between flower size and colour contrast in the subalpine region, but a negative correlation at the lower altitudes. At high elevations in Norway, where pollinators other than bees are common, flower size was positively correlated with colour contrast. At low and medium altitudes in Norway and in Australia, we did not observe a significant relationship between size and colour contrast. We thus find that the relationship between size, green and colour contrast cannot be generalized across communities, thus suggesting that flower visual signal adaptations to local pollinators are not limited to chromatic contrast.

Rewardlessness in orchids: how frequent and how rewardless?

About one-third of orchid species are thought to offer no floral reward and therefore attract pollinators through deception. Statements of this idea are common in the botanical literature, but the empirical basis of the estimate is rarely mentioned. We traced citation pathways for the one-third estimate in a sample of the literature and found that the paths lead to empirical foundations that are surprisingly narrow. Moreover, recent measurements have detected minute quantities of sugar available to insect visitors in some orchids thought to be rewardless, raising the possibility of a pollination strategy that is largely deceitful but different to absolute rewardlessness. The orchids are a well-studied group and there is no doubt that rewardlessness is common in the family. However, greater empirical effort is needed to verify rewardlessness in orchids and to explore geographic and environmental variation in the proportion of rewardless species.

Colour evolution within orchids depends on whether the pollinator is a bee or a fly.

Orchids are a classic angiosperm model for understanding biotic pollination. We studied orchid species within two species-rich herbaceous communities that are known to have either hymenopteran or dipteran insects as the dominant pollinators, in order to understand how flower colour relates to pollinator visual systems. We analysed features of the floral reflectance spectra that are significant to pollinator visual systems and used models of dipteran and hymenopteran colour vision to characterise the chromatic signals used by fly-pollinated and bee-pollinated orchid species. In contrast to bee-pollinated flowers, fly-pollinated flowers had distinctive points of rapid reflectance change at long wavelengths and a complete absence of such spectral features at short wavelengths. Fly-pollinated flowers also had significantly more restricted loci than bee-pollinated flowers in colour space models of fly and bee vision alike. Globally, bee-pollinated flowers are known to have distinctive, consistent colour signals. Our findings of different signals for fly pollination is consistent with pollinator-mediated selection on orchid species that results from the distinctive features of fly visual systems.

Floral colours in a world without birds and bees: the plants of Macquarie Island.

We studied biotically pollinated angiosperms on Macquarie Island, a remote site in the Southern Ocean with a predominately or exclusively dipteran pollinator fauna, in an effort to understand how flower colour affects community assembly. We compared a distinctive group of cream-green Macquarie Island flowers to the flora of likely source pools of immigrants and to a continental flora from a high latitude in the northern hemisphere. We used both dipteran and hymenopteran colour models and phylogenetically informed analyses to explore the chromatic component of community assembly. The species with cream-green flowers are very restricted in colour space models of both fly vision and bee vision and represent a distinct group that plays a very minor role in other communities. It is unlikely that such a community could form through random immigration from continental source pools. Our findings suggest that fly pollination has imposed a strong ecological filter on Macquarie Island, favouring floral colours that are rare in continental floras. This is one of the strongest demonstrations that plant-pollinator interactions play an important role in plant community assembly. Future work exploring colour choices by dipteran flower visitors would be valuable.

Behavioural evidence of colour vision in free flying stingless bees.

Colour vision was first demonstrated with behavioural experiments in honeybees 100 years ago. Since that time a wealth of quality physiological data has shown a highly conserved set of trichromatic colour receptors in most bee species. Despite the subsequent wealth of behavioural research on honeybees and bumblebees, there currently is a relative dearth of data on stingless bees, which are the largest tribe of the eusocial bees comprising of more than 600 species. In our first experiment we tested Trigona cf. fuscipennis, a stingless bee species from Costa Rica in a field setting using the von Frisch method and show functional colour vision. In a second experiment with these bees, we use a simultaneous colour discrimination test designed for honeybees to enable a comparative analysis of relative colour discrimination. In a third experiment, we test in laboratory conditions Tetragonula carbonaria, an Australian stingless bee species using a similar simultaneous colour discrimination test. Both stingless bee species show relatively poorer colour discrimination compared to honeybees and bumblebees; and we discuss the value of being able to use these behavioural methods to efficiently extend our current knowledge of colour vision and discrimination in different bee species.

A calibrated gray scale for forensic ultraviolet photography.

The human eye is an important tool for observing evidence, and visual evidence in turn may be documented onto a photographic medium. The human eye is generally sensitive only to a narrow band of the electromagnetic spectrum from about 700 nm (red) to about 400 nm (violet/blue). It is possible to extend the range of radiations over which evidence may be documented by using the natural ultraviolet sensitivity of photographic films. However, photographing evidence with ultraviolet radiation ultimately involves presenting the information to subjects who may have no prior experience at viewing these wavelengths. This study shows that it is necessary to use a calibrated ultraviolet reflecting gray scale to allow meaningful interpretation of results.

Biological significance of distinguishing between similar colours in spectrally variable illumination: bumblebees (Bombus terrestris) as a case study.

Individual bumblebees were trained to choose between rewarded target flowers and non-rewarded distractor flowers in a controlled illumination laboratory. Bees learnt to discriminate similar colours, but with smaller colour distances the frequency of errors increased. This indicates that pollen transfer might occur between flowers with similar colours, even if these colours are distinguishable. The effect of similar colours on reducing foraging accuracy of bees is evident for colour distances high above discrimination threshold, which explains previous field observations showing that bees do not exhibit complete flower constancy unless flower colour between species is distinct. Bees tested in spectrally different illumination conditions experienced a significant decrease in their ability to discriminate between similar colours. The extent to which this happens differs in different areas of colour space, which is consistent with a von Kries-type model of colour constancy. We find that it would be beneficial for plant species to have highly distinctive colour signals to overcome limitations on the bees performance in reliably judging differences between similar colours. An exception to this finding was flowers that varied in shape, in which case bees used this cue to compensate for inaccuracies of colour vision.

Ocular filtering of ultraviolet radiation and the spectral spacing of photoreceptors benefit Von Kries colour constancy.

Ocular filters in the eyes of many vertebrates, including humans, absorb wavelengths shorter than approximately 400nm. These filters prevent the beta-band of a visual pigment from being exposed to ultraviolet radiation, essentially narrowing the spectral sensitivity of the different photoreceptor classes. A comparison of different hypothetical visual systems is used to show that von Kries colour constancy is improved by ocular filtration of ultraviolet radiation, whilst there is no reduction in colour discrimination. Furthermore, it is shown that the asymmetric spectral spacing of different photoreceptor classes present in the human visual system may benefit colour constancy. The results are interpreted in relation to predictions of von Kries colour constancy for a standard human observer.

Limits to the salience of ultraviolet: lessons from colour vision in bees and birds.

Ultraviolet is an important component of the photic environment. It is used by a wide variety of animals and plants in mutualistic communication, especially in insect and flower inter-relationships. Ultraviolet reflections and sensitivity are also becoming well considered in the relationships between vertebrates and their environment. The relative importance of ultraviolet vis à vis other primary colours in trichromatic or tetrachromatic colour spaces is discussed, and it is concluded that ultraviolet is, in most cases, no more important that blue, green or red reflections. Some animals may use specific wavebands of light for specific reactions, such as ultraviolet in escape or in the detection of polarised light, and other wavebands in stimulating feeding, oviposition or mating. When colour vision and, thus, the input from more than a single spectral receptor type are concerned, we point out that even basic predictions of signal conspicuousness require knowledge of the neuronal wiring used to evaluate the signals from all receptor types, including the ultraviolet. Evolutionary analyses suggest that, at least in arthropods, ultraviolet sensitivity is phylogenetically ancient and undergoes comparatively little evolutionary fine-tuning. Increasing amounts of ultraviolet in the photic environment, as caused by the decline of ozone in the atmosphere, are not likely to affect colour vision. However, a case for which ultraviolet is possibly unique is in the colour constancy of bees. Theoretical models predict that bees will perform poorly at identifying pure ultraviolet signals under conditions of changing illumination, which may explain the near absence of pure ultraviolet-reflecting flowers in nature.