Heather M. Whitney

Floral Iridescence, Produced by Diffractive Optics, Acts As a Cue for Animal Pollinators

Iridescence, the change in hue of a surface with varying observation angles, is used by insects, birds, fish, and reptiles for species recognition and mate selection. We identified iridescence in flowers of Hibiscus trionum and Tulipa species and demonstrated that iridescence is generated through diffraction gratings that might be widespread among flowering plants. Although iridescence might be expected to increase attractiveness, it might also compromise target identification because the objects appearance will vary depending on the viewers perspective. We found that bumblebees (Bombus terrestris) learn to disentangle flower iridescence from color and correctly identify iridescent flowers despite their continuously changing appearance. This ability is retained in the absence of cues from polarized light or ultraviolet reflectance associated with diffraction gratings.

Detection and Learning of Floral Electric Fields by Bumblebees

Flowers and Bees Have “Sparks” Plants and their pollinators have very intimate interactions, as emphasized by the many classic cases of coevolution among species of each. Such close relationships require signaling between plant and pollinator. Coordination between plant signals and pollinator perception has been shown to exist in flower color, shape, and odor. Clarke et al. (p. 66, published online 21 February) report the potential for a distinct mode of plant–pollinator communication, electric fields. Natural floral electric fields, which are impacted by visits from naturally charged bees, were easily discriminated by bees, based on their level, pattern, and structure, and improved the rate at which bees remembered the location of a nectar reward. Flower-specific electric fields are used by bumblebees to enhance discrimination and memory of floral rewards. Insects use several senses to forage, detecting floral cues such as color, shape, pattern, and volatiles. We report a formerly unappreciated sensory modality in bumblebees (Bombus terrestris), detection of floral electric fields. These fields act as floral cues, which are affected by the visit of naturally charged bees. Like visual cues, floral electric fields exhibit variations in pattern and structure, which can be discriminated by bumblebees. We also show that such electric field information contributes to the complex array of floral cues that together improve a pollinator’s memory of floral rewards. Because floral electric fields can change within seconds, this sensory modality may facilitate rapid and dynamic communication between flowers and their pollinators.

Conical Epidermal Cells Allow Bees to Grip Flowers and Increase Foraging Efficiency

The plant surface is by default flat, and development away from this default is thought to have some function of evolutionary advantage. Although the functions of many plant epidermal cells have been described, the function of conical epidermal cells, a defining feature of petals in the majority of insect-pollinated flowers, has not. The location and frequency of conical cells have led to speculation that they play a role in attracting animal pollinators. Snapdragon (Antirrhinum) mutants lacking conical cells have been shown to be discriminated against by foraging bumblebees. Here we investigated the extent to which a difference in petal surface structure influences pollinator behavior through touch-based discrimination. To isolate touch-based responses, we used both biomimetic replicas of petal surfaces and isogenic Antirrhinum lines differing only in petal epidermal cell shape. We show that foraging bumblebees are able to discriminate between different surfaces via tactile cues alone. We find that bumblebees use color cues to discriminate against flowers that lack conical cells--but only when flower surfaces are presented at steep angles, making them difficult to manipulate. This facilitation of physical handling is a likely explanation for the prevalence of conical epidermal petal cells in most flowering plants.

Why do so many petals have conical epidermal cells

BACKGROUND The conical epidermal cells found on the petals of most Angiosperm species are so widespread that they have been used as markers of petal identity, but their function has only been analysed in recent years. This review brings together diverse data on the role of these cells in pollination biology. SCOPE The published effects of conical cells on petal colour, petal reflexing, scent production, petal wettability and pollinator grip on the flower surface are considered. Of these factors, pollinator grip has been shown to be of most significance in the well-studied Antirrhinum majus/bumble-bee system. Published data on the relationship between epidermal cell morphology and floral temperature were limited, so an analysis of the effects of cell shape on floral temperature in Antirrhinum is presented here. Statistically significant warming by conical cells was not detected, although insignificant trends towards faster warming at dawn were found, and it was also found that flat-celled flowers could be warmer on warm days. The warming observed is less significant than that achieved by varying pigment content. However, the possibility that the effect of conical cells on temperature might be biologically significant in certain specific instances such as marginal habitats or weather conditions cannot be ruled out. CONCLUSIONS Conical epidermal cells can influence a diverse set of petal properties. The fitness benefits they provide to plants are likely to vary with pollinator and habitat, and models are now required to understand how these different factors interact.

Mutations perturbing petal cell shape and anthocyanin synthesis influence bumblebee perception of Antirrhinum majus flower colour

We wished to understand the effects on pollinator behaviour of single mutations in plant genes controlling flower appearance. To this end, we analysed snapdragon flowers (Antirrhinum majus), including the mixta and nivea mutants, in controlled laboratory conditions using psychophysical tests with bumblebees. The MIXTA locus controls petal epidermal cell shape, and thus the path that incident light takes within the pigment-containing cells. The effect is that mixta mutant flowers are pink in comparison to the wild type purple flowers, and mutants lack the sparkling sheen of wild type flowers that is clearly visible to human observers. Despite their fundamentally different appearance to humans, and even though bees could discriminate the flowers, inexperienced bees exhibited no preference for either type, and the flowers did not differ in their detectability in a Y-maze—either when the flowers appeared in front of a homogeneous or a dappled background. Equally counterintuitive effects were found for the non-pigmented, UV reflecting nivea mutant: even though the overall reflectance intensity and UV signal of nivea flowers is several times that of wild type flowers, their detectability was significantly reduced relative to wild type flowers. In addition, naïve foragers preferred wild type flowers over nivea mutants, even though these generated a stronger signal in all receptor types. Our results show that single mutations affecting flower signal can have profound effects on pollinator behaviour—but not in ways predictable by crude assessments via human perception, nor simple quantification of UV signals. However, current models of bee visual perception predict the observed effects very well.

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.

The interaction of temperature and sucrose concentration on foraging preferences in bumblebees

Several authors have found that flowers that are warmer than their surrounding environment have an advantage in attracting pollinators. Bumblebees will forage preferentially on warmer flowers, even if equal nutritional reward is available in cooler flowers. This raises the question of whether warmth and sucrose concentration are processed independently by bees, or whether sweetness detectors respond to higher sugar concentration as well as higher temperature. We find that bumblebees can use lower temperature as a cue to higher sucrose reward, showing that bees appear to process the two parameters strictly independently. Moreover, we demonstrate that sucrose concentration takes precedence over warmth, so that when there is a difference in sucrose concentration, bees will typically choose the sweeter feeder, even if the less sweet feeder is several degrees warmer.

BIOLOGICAL EFFECTS OF PHYSICALLY CONDITIONED WATER

Abstract Physically conditioned water, made by passage between the poles of a magnet or by injecting a weak electrical signal, is used for the prevention and removal of lime-scale, but also has diverse biological effects. In this investigation, we have shown both stimulations and inhibitions of the multiplication of yeast cells, depending on the degree of conditioning. Weakly conditioned water is stimulatory, but strongly conditioned water is inhibitory. Conditioned water also increases the toxicity of heavy metal ions such as copper and cobalt. We suggest that these effects are due to colloidal impurities, which have been activated by the conditioning process, interacting with structural calcium in the cell membranes to make them more permeable. The stimulations may be due to the inward leakage of small amounts of calcium to stimulate metabolism by acting as a “secondary messenger”. The inhibitions may be due to more severe damage to the membranes allowing the entry of larger and more toxic quantities of calcium and, if present, other noxious materials. This is discussed in relation to the beneficial and adverse effects of conditioned water on the growth and well-being of higher organisms. Possible applications include stimulating the growth of organisms at optimum levels of conditioning, inhibiting unwanted microorganisms at higher levels and increasing the efficacy of biocides. Attention is also drawn to a similarity between the biological effects of conditioned water and those of weak electromagnetic fields. We discuss the possibility that some of the effects of electromagnetic fields on living organisms are due to their interacting with colloidal cell components and membrane surfaces by a mechanism analogous to the conditioning of water. This is discussed in relation to ion cyclotron resonance phenomena and an explanation based on our proposals given for the hitherto unexplained differences in biological responses to the resonant frequencies for calcium and potassium ions.

Floral temperature and optimal foraging: is heat a feasible floral reward for pollinators?

As well as nutritional rewards, some plants also reward ectothermic pollinators with warmth. Bumble bees have some control over their temperature, but have been shown to forage at warmer flowers when given a choice, suggesting that there is some advantage to them of foraging at warm flowers (such as reducing the energy required to raise their body to flight temperature before leaving the flower). We describe a model that considers how a heat reward affects the foraging behaviour in a thermogenic central-place forager (such as a bumble bee). We show that although the pollinator should spend a longer time on individual flowers if they are warm, the increase in total visit time is likely to be small. The pollinators net rate of energy gain will be increased by landing on warmer flowers. Therefore, if a plant provides a heat reward, it could reduce the amount of nectar it produces, whilst still providing its pollinator with the same net rate of gain. We suggest how heat rewards may link with plant life history strategies.

Function of blue iridescence in tropical understorey plants

The blue colouration seen in the leaves of Selaginella willdenowii is shown to be iridescent. Transmission electron microscopy studies confirm the presence of a layered lamellar structure of the upper cuticle of iridescent leaves. Modelling of these multi-layer structures suggests that they are responsible for the blue iridescence, confirming the link between the observed lamellae and the recorded optical properties. Comparison of blue and green leaves from the same plant indicates that the loss of the blue iridescence corresponds to a loss of the multi-layer structure. The results reported here do not support the idea that iridescence in plants acts to enhance light capture of photosynthetically important wavelengths. The reflectance of light in the range 600–700 nm is very similar for both iridescent and non-iridescent leaves. However, owing to the occurrence of blue colouration in a wide variety of shade dwelling plants it is probable that this iridescence has some adaptive benefit. Possible adaptive advantages of the blue iridescence in these plants are discussed.