Broad spectral sensitivities in the honeybee's photoreceptors limit colour constancy

Colour constancy allows for visual systems to be view stimuli independent of changes in spectral illumination. Chromatic adaptation is likely to be an important mechanism in colour constancy and can be explained by use of the von Kries coefficient law. Chromatic adaptation is compared for the honeybee and three hypothetical visual systems. It is shown that the spectral breadth and asymmetry of photoreceptors in the honeybee may limit colour constancy. In particular, it is demonstrated that the absorption of short-wavelength radiation by the cis-band of chromophore is responsible for a poorer correction for bee colours rich in ultraviolet reflectance. The results are discussed in relation to theoretical considerations of von Kries colour constancy and the physiology of eye design in some other species for which colour constancy has been demonstrated. Accepted: 14 August 1999     

The evolutionary adaptation of flower colours and the insect pollinators' colour vision

Summary The evolutionary tuning between floral colouration and the colour vision of flower-visiting Hymenoptera is quantified by evaluating the informational transfer from the signalling flower to the perceiving pollinator. The analysis of 180 spectral reflection spectra of angiosperm blossoms reveals that sharp steps occur precisely at those wavelengths where the pollinators are most sensitive to spectral differences. Straight-forward model calculations determine the optimal set of 3 spectral photoreceptor types for discrimination of floral colour signals on the basis of perceptual difference values. The results show good agreement with the sets of photoreceptors characterized electrophysiologically in 40 species of Hymenoptera.     

Flower colour variation across a hybrid zone in Antirrhinum as perceived by bumblebee pollinators

To assess if pollinators’ behaviour could explain the maintenance of hybrid zones between different flower colour morphs, we analyzed flower colour variation in an Antirrhinum hybrid zone using spectrometry and a model of bee perception. Some colours generated by hybridization were not observed in any Antirrhinum species and even appeared to be rare among angiosperms. Variation in flower colours within the hybrid zone was continuous; the most similar colours were predicted not to be discriminated from one another in natural foraging situations. However, when compared at a scale corresponding to bees’ foraging range, some flower colours could be discriminated from all colours displayed by neighbouring plants. This could affect pollinator behaviour and explain lower visitation rates within the centre of the hybrid zone. Behavioural studies involving bumblebees and plant mixtures of parental and hybrid flower colours carefully characterized with appropriate visual models will be necessary to test this hypothesis. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Lars Chittka     

Biogeography of the yeasts of ephemeral flowers and their insects

We studied specific yeast communities vectored by beetles, drosophilids, and bees that visit ephemeral flowers, mostly in the genus Hibiscus and in the families Convolvulaceae and Cactaceae, in the Neotropical, Nearctic, and Australian biogeographic regions. The communities consist mostly of yeasts in four clades centered around the genera Metschnikowia, Kodamaea, Wickerhamiella, and Starmerella. The largest geographic discontinuity occurs as a function of the nitidulid beetle species that dominate the non-pollinator insect visitors of the flowers. This partitions the New World, where the dominant beetle is in the genus Conotelus, from the Australian biogeographic region, dominated by species of Aethina. Distinct but sympatric insects may also carry radically different yeast communities.     

Bumblebees directly perceive variations in the spectral quality of illumination

Individual bumblebees were tested on a task of walking along a test tunnel to collect sucrose solution in an initial training illumination condition that simulated natural daylight, and in two spectrally different illumination conditions. Compared to the training condition the bees took a significantly longer time to complete the task in conditions that simulated either an ultraviolet negative illumination environment, or an illumination environment that represented blue skylight. In a control condition, bees did not derive this cue from spectral information reflected from the dark background material. This result shows that bees can directly perceive spectral changes in illumination conditions, even in the context of a task that does not require colour processing. This potentially enables the visual system of bees to have prior knowledge about the spectral quality of illumination conditions in which they may forage. The findings are discussed in relation to both theoretical models and empirical evidence of colour constancy, and it is concluded that bees can use multiple mechanisms to solve the dilemma posed by having to find colour targets in the spectrally different illumination conditions that exist for insects visiting flowers.     

浅谈色彩在昆虫绘画中的应用

尝试以彩色绘画的方式表现昆虫的体貌特征,应先认识色彩,了解色彩的特性,以色彩变化的基本规律去观察昆虫体色的微妙变化,分析、找出昆虫体壁每一部分的色彩倾向。绘画时,只有掌握并运用色彩的特性及色彩的对比关系,在平面的纸张上才能表现出昆虫的立体效果。     

Where have all the blue flowers gone: pollinator responses and selection on flower colour in New Zealand Wahlenbergia albomarginata

Although pollinators are thought to select on flower colour, few studies have experimentally decoupled effects of colour from correlated traits on pollinator visitation and pollen transfer. We combined selection analysis and phenotypic manipulations to measure the effect of petal colour on visitation and pollen export at two spatial scales in Wahlenbergia albomarginata. This species is representative of many New Zealand alpine herbs that have secondarily evolved white or pale flowers. The major pollinators, solitary bees, exerted phenotypic selection on flower size but not colour, quantified by bee vision. When presented with manipulated flowers, bees visited flowers painted blue to resemble a congener over white flowers in large, but not small, experimental arrays. Pollen export was higher for blue flowers in large arrays. Pollinator preference does not explain the pale colouration of W. albomarginata, as commonly hypothesized. Absence of bright blue could be driven instead by indirect selection of correlated characters.     

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.     

The ultraviolet colour component enhances the attractiveness of red flowers of a bee-pollinated plant

蜂媒花很少呈红色,这可能是由于蜂类的眼睛缺少红光受体,因而较难搜寻红色目标。尽管有实验研究过蜂类对红色的反应,但所用目标多为纯红色。而人类感受到的红色中其实包含细微的色调差别,这些差别还甚少受到关注。我们在本研究中检验了如下假说：红花中的紫外反射组分能够增加其彩调反差,因而增强其对蜂类的吸引力。紫草科的密花滇紫草(Onosma confertum)由熊蜂传粉,其花为红色,但紫外反射明显。本研究利用该植物设计了表型操控实验,在野外检验了紫外反射对传粉者访花行为的影响。我们还在受控环境下检验了无经验熊蜂的色彩偏好。利用两个色彩模型评估熊蜂的色彩感受,用彩调反差和明度反差两个参数进行估计。研究发现,自然状态下及无经验的饲养熊蜂均偏好访问具有紫外反射的红色目标。色彩模型分析表明,具紫外反射的红花对蜂而言色彩更纯,且与叶片背景的彩调反差更大;而它们与无紫外反射的红花在明度反差方面(以绿光受体的反差估计)表现相似。这些结果表明,密花滇紫草红色花朵中的紫外反射组分增加了其彩调反差,增强了其对熊蜂的吸引力。我们推测,由于某些传粉者(例如蜂类)无红光受体,对由它们传粉的红色花而言,光谱中的副反射特征(例如紫外反射)可能是其花色中的必要组分。     

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.     

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.     

Biological significance of distinguishing between similar colours in spectrally variable illumination: bumblebees (<Emphasis Type="Italic">Bombus terrestris</Emphasis>) 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.     

The quality of flower-based ecosystem services in field margins and road verges from human and insect pollinator perspectives

We contrasted traditionally used indicators of service provision quality, such as overall species richness and growth form composition, to three more specific functional properties: functional diversity, functional intensity, and functional stability. We defined flower colour as a functional trait perceived differently by humans and insect pollinators, and used user specific colour richness, flower size, and species richness within colour group as indicators of these three properties. We asked (1) do field margins and road verges provide flower-based ecosystem services with the quality of permanent grasslands; and (2) do traditional and detailed functional indicators of service provision quality agree on the service quality ranking of habitats?In an agricultural landscape of central and south-eastern Estonia (115 ÿ 95 km area, centroid 26°49⿲43⿳ and 58°54⿲49⿳) we sampled 87 field margins and 111 road verges as linear grassland-substitution habitats, and 84 permanent grasslands to scale their service quality.Linear habitats generally provided service of lower quality than permanent grasslands, but detailed indicators showed less evident contrast among habitat types than the overall species richness and stronger contrast than the proportion of forbs. Detailed indices, however, had strong seasonal dynamics to take into account. Vegetation in the first year field margins had greater colour richness (functional diversity) and species richness within colour groups (functional stability), but the smallest flower size (functional intensity), in contrast to road verges. By the third year of succession, field margins had become more similar to road verges. Indication of service provision quality differed between humans and pollinators, but their estimates were correlated across habitats.We showed that (1) combinations of specific service quality indicators provide more adequate information than generalized richness or growth form system, and (2) single grassland surrogate habitat type is an insufficient service providing substitute for permanent grasslands, although a mosaic of these habitats might be more efficient. Therefore, remnant fragments of semi-natural grasslands should receive top priority attention for conservation and restoration, particularly in agriculture dominated landscapes.     

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.     

Effects of floral diversity and pollinator behaviour on the persistence of hybrid zones between plants sharing pollinators

Background: Plant hybrid zones often display a large diversity in floral traits due to segregation and recombination occurring in the second and later hybrid generations. Such diversity can have important effects on pollinator behaviour.

Aims: We hypothesised that the number of floral types and their relative densities may influence pollinator preferences and level of flower constancy.

Methods: To test the influence of floral diversity on the number of visits to inflorescences of the most common type and on the level of flower constancy, we monitored bumblebees’ behaviour in experimental arrays of artificial inflorescences with a common (‘parental’) floral type mixed with either one or three uncommon (‘hybrid’) floral types.

Results: We found that pollinators preferentially visited the most common flower colour and showed slightly greater constancy when the number of uncommon floral types was increased.

Conclusions: We conclude that diversity in flower colour could promote moderate pollinator constancy in some types of hybrid zones. Segregation of floral phenotypes due to hybridisation may result in small levels of pollinator-mediated assortative mating and contribute, to a small extent, and most probably in combination with other factors, to the persistence of some hybrid zones.     

Light absorption patterns of intact Rosa flowers in relation to the flower colour

Absorption curves of fresh, intact petals from 18 rose cultivars and 2 species were measured and compared with visual evaluations of their colours and there was a reasonable correlation. The in vivo maxima of anthocyanin absorption were in the range of 520–560 nm. Five patterns of absorption spectrum in the visible region were recognized: (a) maximum range ca. 520–535 rim (red roses); (b) as (a) but low absorbance (pink roses); (c) absorption pattern varying with age of flowers; (d) absorption at long wavelengths in blue roses due to co-pigmentation of cyanin, flavonols; (e) absorption of carotenoids and anthocyanins together in yellow, orange or orange red flowers.     

Pollination and fruit set in two rewardless slipper orchids and their hybrids (Cypripedium,Orchidaceae): large yellow flowers outperform small white flowers in the northern tall grass prairie

• Species with rewardless flowers often have low fruit to flower ratios, although wide temporal and spatial variation in fruiting success can occur. We compared floral phenotypes, insect visitors and fruiting success in four populations of the small white (Cypripedium candidum) and yellow (C. parviflorum) lady’s slipper orchids and their hybrids near the northern extent of North America’s tall grass prairie.
• Flower and fruit numbers were observed for two seasons on marked individuals (n = 1811). Floral traits were measured on 82–140 individuals per taxon and analysed in relation to fruiting success. All insects found inside flowers were collected, inspected for pollen smears and measured for comparison to floral features.
• Among orchid taxa, C. candidum had the smallest flowers, lowest number and variety of insect visitors, and lowest fruit to flower ratios. These measures were intermediate in hybrids and highest in C. parviflorum, despite low flower numbers in the latter. Within orchid taxa, fruit number was positively related to flower number, but fruit to flower ratios decreased slightly, as would be expected if pollinators left unrewarding patches. Potential pollinators included the dipteran Odontomyia pubescens and hymenopterans Andrena spp., Apis mellifera and Lasioglossum zonulum.
• Cypripedium parviflorum had a reproductive advantage over C. candidum across multiple populations and years. Hybrids showed segregation for floral traits, and hybrid fruiting success increased with a deeper intensity of yellow pigment and larger escape routes for floral visitors. These same attributes likely contributed to the relatively high fruit set in C. parviflorum in the study region.

     

The potential for selection on pollen colour dimorphisms in Nigella degenii: morph-specific differences in pollinator visitation, fertilisation success and siring ability

Two subspecies of Nigella degenii (Ranunculaceae) possess a dimorphism in pollen colour and vary extensively in frequency of the two morphs in natural populations. Here we investigate the role of selection on pollen colour during the pollination phase in the two subspecies and its potential contribution to the maintenance of this colour variation. In a combination of common garden experiments and field observations, we obtained data on pollinator visitation rates and explored the effect of pollen colour on fertilisation success and siring ability under conditions of low vs. high pollen competition. In experimental gardens, naïve pollinators responded differently to plants with different pollen colour, but the favoured morph varied between dates and locations, and colour morphs were not visited in a frequency-dependent manner. Donor plants with dark pollen had a reproductive advantage (higher seed set) in single-donor pollinations, but the realised siring ability (measured by progeny morph ratio) was highly variable between different two-donor crosses with no general bias towards the light or dark morph. Therefore, although the dark pollen type appears to have a general selective advantage in terms of fertilisation success, our data are also consistent with a scenario involving the maintenance of both colour morphs, particularly under conditions of high pollen competition, a variable genetic background and/or spatial or temporal variation in the pollinator fauna.