RESUMO
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.
RESUMO
Floral colours represent a highly diverse communication signal mainly involved in flower visitors' attraction and guidance, but also flower discrimination, filtering non-pollinators and discouraging floral antagonists. The divergent visual systems and colour preferences of flower visitors, as well as the necessity of cues for flower detection and discrimination, foster the diversity of floral colours and colour patterns. Despite the bewildering diversity of floral colour patterns, a recurrent component is a yellow UV-absorbing floral centre, and it is still not clear why this pattern is so frequent in angiosperms. The pollen, anther, stamen, and androecium mimicry (PASAM) hypothesis suggests that the system composed of the flowers possessing such yellow UV-absorbing floral reproductive structures, the flowers displaying central yellow UV-absorbing structures as floral guides, and the pollen-collecting, as well as pollen-eating, flower visitors responding to such signals constitute the world's most speciose mimicry system. In this review, we call the attention of researchers to some hypothetical PASAM systems around the globe, presenting some fascinating examples that illustrate their huge diversity. We will also present new and published data on pollen-eating and pollen-collecting pollinators' responses to PASAM structures supporting the PASAM hypothesis and will discuss how widespread these systems are around the globe. Ultimately, our goal is to promote the idea that PASAM is a plausible first approach to understanding floral colour patterns in angiosperms.
Assuntos
Magnoliopsida , Polinização , Polinização/fisiologia , Reprodução , Flores/fisiologia , Pólen/fisiologia , Magnoliopsida/fisiologiaRESUMO
The flower perianth has various, non-mutually exclusive functions, such as visual signalling to pollinators and protecting the reproductive organs from the elements and from florivores, but how different perianth structures and their different sides play a role in these functions is unclear. Intriguingly, in many species there is a clear colour difference between the different sides of the perianth, with colour patterns or pigmentation present on only one side. Any adaptive benefit from such colour asymmetry is unclear, as is how the asymmetry evolved. In this viewpoint paper, we address the phenomenon of flowers with differently coloured inner and outer perianth sides, focusing on petals of erect flowers. Guided by existing literature and our own observations, we delineate three non-mutually exclusive evolutionary hypotheses that may explain the factors underlying differently coloured perianth sides. The pollen-protection hypothesis predicts that the outer side of petals contributes to protect pollen against UV radiation, especially during the bud stage. The herbivore-avoidance hypothesis predicts that the outer side of petals reduces the flower's visibility to herbivores. The signalling-to-pollinators hypothesis predicts that flower colours evolve to increase conspicuousness to pollinators. The pollen-protection hypothesis, the herbivore-avoidance hypothesis, and the signalling-to-pollinators hypothesis generate largely but not entirely overlapping predictions about the colour of the inner and outer side of the petals. Field and laboratory research is necessary to disentangle the main drivers and adaptive significance of inner-outer petal side colour asymmetry.
Assuntos
Evolução Biológica , Flores , Pigmentação , Polinização , Animais , Cor , Flores/fisiologia , Flores/anatomia & histologia , Herbivoria/fisiologia , Pigmentação/fisiologia , Pólen/fisiologia , Polinização/fisiologiaRESUMO
The tropical Melastomataceae are characterized by poricidal anthers which constitute a floral filter selecting for buzz-pollinating bees. Stamens are often dimorphic, sometimes with discernible feeding and pollinating functions. Rhynchanthera grandiflora produces nectarless flowers with four short stamens and one long stamen; all anthers feature a narrow elongation with an upwards facing pore. We tested pollen transfer by diverse foraging bees and viability of pollen from both stamen types. The impact of anther morphology on pollen release direction and scattering angle was studied to determine the plant's reproductive strategy. Medium-sized to large bees sonicated flowers in a specific position, and the probability of pollen transfer correlated with bee size even among these legitimate visitors. Small bees acted as pollen thieves or robbers. Anther rostrum and pore morphology serve to direct and focus the pollen jet released by floral sonication towards the pollinator's body. Resulting from the ventral and dorsal positioning of the short and long stamens, respectively, the pollinator's body was widely covered with pollen. This improves the plant's chances of outcrossing, irrespective of which bee body part contacts the stigma. Consequently, R. grandiflora is also able to employ bee species of various sizes as pollen vectors. The strategy of spreading pollen all over the pollinator's body is rather cost-intensive but counterbalanced by ensuring that most of the released pollen is in fact transferred to the bee. Thus, flowers of R. grandiflora illustrate how specialized morphology may serve to improve pollination by a functional group of pollinators.
Assuntos
Abelhas , Melastomataceae , Polinização , Animais , Abelhas/anatomia & histologia , Abelhas/classificação , Biodiversidade , Flores/anatomia & histologia , Melastomataceae/anatomia & histologia , Pólen , ReproduçãoRESUMO
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.
Assuntos
Dípteros/fisiologia , Flores/anatomia & histologia , Magnoliopsida/anatomia & histologia , Animais , Abelhas/fisiologia , Aves/fisiologia , Cor , Flores/genética , Flores/fisiologia , Geografia , Ilhas , Magnoliopsida/genética , Magnoliopsida/fisiologia , Modelos Biológicos , Filogenia , Polinização , TasmâniaRESUMO
Colour is one of the most obvious advertisements of flowers, and occurs in a huge diversity among the angiosperms. Flower colour is responsible for attraction from a distance, whereas contrasting colour patterns within flowers aid orientation of flower visitors after approaching the flowers. Due to the striking differences in colour vision systems and neural processing across animal taxa, flower colours evoke specific behavioural responses by different flower visitors. We tested whether and how yellow flowers differ in their spectral reflectance depending on the main pollinator. We focused on bees and birds and examined whether the presence or absence of the widespread UV reflectance pattern of yellow flowers predicts the main pollinator. Most bee-pollinated flowers displayed a pattern with UV-absorbing centres and UV-reflecting peripheries, whereas the majority of bird-pollinated flowers are entirely UV- absorbing. In choice experiments we found that bees did not show consistent preferences for any colour or pattern types. However, all tested bee species made their first antennal contact preferably at the UV-absorbing area of the artificial flower, irrespective of its spatial position within the flower. The appearance of UV patterns within flowers is the main difference in spectral reflectance between yellow bee- and bird-pollinated flowers, and affects the foraging behaviour of flower visitors. The results support the hypothesis that flower colours and the visual capabilities of their efficient pollinators are adapted to each other.
Assuntos
Abelhas/fisiologia , Flores/química , Flores/fisiologia , Polinização/fisiologia , Animais , Evolução Biológica , Aves , Cor , Pigmentação , Néctar de Plantas , Espectrofotometria Ultravioleta , Raios UltravioletaRESUMO
Ageing brings a number of normal physical changes and sometimes is accompanied by diseases that affect physical functions and abilities. Therefore, the elderly require special consideration in the design of chairs. Requirements for three types of chairs for the elderly are presented. A description of a typical user of each chair provides a reference for the level of ability that must be accommodated. Features of the chairs that promote comfort, safety, ease of ingress and egress and ease of propulsion (where applicable) are discussed in relation to some of the special problems experienced by some of the older members of the population.
RESUMO
Shape sensing is useful in the computer aided prosthetic fitting process for two purposes. 1. To input characteristic prosthetic shapes that have been developed over the years through the experience of prosthetists. 2. To provide an accurate and rapid measurement of the anatomical shape of the stump. This paper describes two instruments which have been built to meet these objectives.
Assuntos
Cotos de Amputação/patologia , Computadores , Desenho de Prótese , Humanos , Perna (Membro)RESUMO
The two widespread tropical Solanum species S. paniculatum and S. stramoniifolium are highly dependent on the visits of large bees that pollinate the flowers while buzzing them. Both Solanum species do not offer nectar reward; the rewarding of bees is thus solely dependent on the availability of pollen. Flower visitors are unable to visually assess the amount of pollen, because the pollen is hidden in poricidal anthers. In this study we ask whether and how the amount of pollen determines the attractiveness of flowers for bees. The number of pollen grains in anthers of S. stramoniifolium was seven times higher than in S. paniculatum. By contrast, the handling time per five flowers for carpenter bees visiting S. paniculatum was 3.5 times shorter than of those visiting S. stramoniifolium. As a result foraging carpenter bees collected a similar number of pollen grains per unit time on flowers of both species. Experimental manipulation of pollen availability by gluing the anther pores showed that the carpenter bees were unable to detect the availability of pollen by means of chemical cues before landing and without buzzing. Our study shows that the efficiency of pollen collecting on S. paniculatum is based on large inflorescences with short between-flower search times and short handling time of individual flowers, whereas that of S. stramoniifolium relies on a large amount of pollen per flower. Interestingly, large carpenter bees are able to adjust their foraging behaviour to drastically different strategies of pollen reward in otherwise very similar plant species.
Assuntos
Abelhas , Comportamento Animal , Flores , Néctar de Plantas , Pólen , Polinização , Solanum , Animais , Inflorescência , RecompensaRESUMO
In this study we show how inexperienced syrphid flies, Eristalis tenax, orient on artificial flowers by means of floral guides. To test the effect of floral guides such as line and ring markings on the probability and speed of the location of a potential food source, we exploited the spontaneous proboscis reaction triggered by yellow colour stimuli. We tested whether and how fast the flies, when placed on the edge of a circular dummy flower, found a small central yellow spot and touched it with the proboscis extended. The flies found the central yellow spot more often and faster if guide lines from the margin to the yellow spot were present. The effect of guide lines was dependent on the colour of the dummy flower, and independent of the colour of the guide lines, except for yellow guide lines releasing the proboscis reaction. The effect of guide lines was stronger if the yellow spot was hidden in a 2 mm deep depression and thus not as easily visible to the flies. Ring guides had a significant effect on performance only when the intensity of the central yellow spot was low.
RESUMO
The labellar gustatory system of the dronefly Eristalis tenax L. (Syrphidae; Diptera) that enables the fly to discriminate between pollen and nectar is investigated, and the triggering of pollen ingestion is examined. In behavioural preference tests, exhaustively extracted pollen of the sunflower Helianthus annuus is consumed in smaller amounts than untreated pollen, indicating that water-soluble substances are important for acceptance. Dry pollen is preferred over moist pollen in which the grains stick together, suggesting that mechanical properties of the pollen also play a role in its sensory assessment. Electrophysiological studies of the labellar taste hairs reveal that aqueous extracts of pollen (2% w/v) stimulate the salt receptor cell, but not the sugar receptor cell. The response of the water receptor cell remains the same as to pure water (or standard electrolyte, 10 mmol.l-1 KCl). Of the 20 amino acids tested, the salt cell is sensitive only to proline in a submillimolar range. Behavioural experiments support the electrophysiological findings. When KCl is applied at concentrations eliciting salt-cell spike frequencies equal to those produced by pollen extract (which is often accepted), the water receptor cell is inhibited and a pronounced rejection behaviour occurs. This rejection of concentrated salt solution in Eristalis is therefore mainly mediated by the inhibition of the water cell.
Assuntos
Células Quimiorreceptoras/fisiologia , Dípteros/fisiologia , Comportamento Alimentar/fisiologia , Pólen , Paladar/fisiologia , Potenciais de Ação/fisiologia , Animais , Eletrofisiologia , Cloreto de Potássio , Prolina , Estimulação Química , Sacarose , ÁguaRESUMO
Avian plumage has long been used to test theories of sexual selection, with humans assessing the colors. However, many birds see in the ultraviolet (<400 nm), to which humans are blind. Consequently, it is important to know whether natural variation in UV reflectance from plumage functions in sexual signaling. We show that female starlings rank males differently when UV wavelengths are present or absent. Principal component analysis of approximately 1300 reflectance spectra (300-700 nm) taken from sexually dimorphic plumage regions of males predicted preference under the UV+ treatment. Under UV- conditions, females ranked males in a different and nonrandom order, but plumage reflectance in the human visible spectrum did not predict choice. Natural variation in UV reflectance is thus important in avian mate assessment, and the prevailing light environment can have profound effects on observed mating preferences.