Pollinator Reward Research Articles

Food Reward Chemistry Explains a Novel Pollinator Shift and Vestigialization of Long Floral Spurs in an Orchid

During the evolutionary history of flowering plants, transitions between pollinator groups (pollinator shifts) have been frequent,1 and contributed to the spectacular radiation of angiosperms.2 Although the evolution of floral traits during pollinator shifts has been studied in real time under controlled laboratory conditions,3 it is challenging to study in nature and therefore poorly understood.4-7 Using a comparative, multidisciplinary approach, we dissect the evolution of floral traits during a pollinator shift in the long-spurred African orchid Satyrium longicauda. Phylogenetic analysis and ecological experiments revealed a shift from moth- to oil-collecting bee pollination. Remarkably, flowers of the bee-pollinated form are similar in morphology, color, and overall volatile chemistry to those of moth-pollinated forms, but differ in having spurs that are mostly devoid of nectar, and have an elevated presence of the oil-derived compound diacetin, which oil-collecting bees use as a cue for oil presence.8 Experiments demonstrated that long spurs are critical for pollination of a moth-pollinated form, but are not needed for pollination of the bee-pollinated form. We conclude that the pollinator shift in Satyrium was mediated by a switch in chemistry of the pollinator reward. The ancestral presence of diacetin might have served as a pre-adaptation for bee pollination, whereas the current mismatch between flower morphology and bees is due to the retention of vestigial floral spurs. These results elucidate the sequence of floral evolution in the early stages of pollinator shifts and help to explain the assembly of suites of co-varying traits through pre-adaptation and vestigialization.9-12.

The Floral Microbiome: Plant, Pollinator, and Microbial Perspectives

Flowers at times host abundant and specialized communities of bacteria and fungi that influence floral phenotypes and interactions with pollinators. Ecological processes drive variation in microbial abundance and composition at multiple scales, including among plant species, among flower tissues, and among flowers on the same plant. Variation in microbial effects on floral phenotype suggests that microbial metabolites could cue the presence or quality of rewards for pollinators, but most plants are unlikely to rely on microbes for pollinator attraction or reproduction. From a microbial perspective, flowers offer opportunities to disperse between habitats, but microbial species differ in requirements for and benefits received from such dispersal. The extent to which floral microbes shape the evolution of floral traits, influence fitness of floral visitors, and respond to anthropogenic change is unclear. A deeper understanding of these phenomena could illuminate the ecological and evolutionary importance of floral microbiomes and their role in the conservation of plant–pollinator interactions.

Influence of light, dark, temperature and drought on metabolite and ion composition in nectar and nectaries of an epiphytic bromeliad species (Aechmea fasciata).

Research into the influence of stress factors, such as drought, different temperatures and/or varied light conditions, on plants due to climate changes is becoming increasingly important. Epiphytes, like many species of the Bromeliaceae, are particularly affected by this, but little is known about impacts on nectar composition and nectary metabolism. We investigated the influence of drought, different temperatures and light-dark regimes on nectar and nectaries of the epiphytic bromeliad species, Aechmea fasciata, and also the influence of drought with the terrestrial bromeliad, Billbergia nutans. The content of sugars, amino acids and ions in nectar and nectaries was analysed using HPLC. In addition, the starch content and the activities of different invertases in nectaries were determined. Compositions of nectar and nectaries were hardly influenced, neither by light nor dark, nor by different temperatures. In contrast, drought revealed changes in nectar volumes and nectar sugar compositions in the epiphytic bromeliad as well as in the terrestrial bromeliad. In both species, the sucrose-to-hexose ratio in nectar decreased considerably during the drought period. These changes in nectar sugar composition do not correlate with changes in the nectaries. The total sugar, amino acid and ion concentrations remained constant in nectar as well as in nectaries during the drought period. Changes in nectar composition or in the production of floral pollinator rewards are likely to affect plant-pollinator interactions. It remains questionable how far the adaptations of the bromeliads to drought and diverse light or temperature conditions are still sufficient.

Reproductive biology and pollinators in two enantiostylous Qualea species (Vochysiaceae) in the Brazilian Cerrado.

Enantiostyly is a floral polymorphism in which two floral forms in the same species differ in deflection of the stigma to right or left position. In monomorphic enantiostylous plants, flowers of the two morphs occur within the same individual, usually in the same proportion. In self-compatible species the function of monomorphic enantiostyly is proposed to increase outcrossing rates and offer a reproductive advantage under pollination limitation. Enantiostylous species are usually self-compatible and show heteranthery, with poricide anthers and pollen as pollinator reward; however, there are families, such as Vochysiaceae, that have different characteristics. We analysed the reproductive system and pollination biology of Qualea parviflora and Q. multiflora, two enantiostylous species from the Brazilian Cerrado that have specific morphological and physiological traits. For this, we characterized flower traits, performed hand pollinations and studied floral visitors. We found no differences between morphs in the proportion of flowers, nectar produced or its concentration, pollen quantity and fruit set. Both species were self-incompatible and quite generalist regarding floral visitors. Enantiostyly in self-incompatible plants seems to confer a reproductive advantage by reducing self-interference resulting from stigma clogging. This novel result helps to expand our knowledge on this complex floral polymorphism and opens new avenues for future research on this topic.

Do food trichomes occur in Pinguicula (Lentibulariaceae) flowers?

Background and AimsFloral food bodies (including edible trichomes) are a form of floral reward for pollinators. This type of nutritive reward has been recorded in several angiosperm families: Annonaceae, Araceae, Calycanthaceae, Eupomatiaceae, Himantandraceae, Nymphaeaceae, Orchidaceae, Pandanaceae and Winteraceae. Although these bodies are very diverse in their structure, their cells contain food material: starch grains, protein bodies or lipid droplets. In Pinguicula flowers, there are numerous multicellular clavate trichomes. Previous authors have proposed that these trichomes in the Pinguicula flower play the role of ‘futterhaare’ (‘feeding hairs’) and are eaten by pollinators. The main aim of this study was to investigate whether the floral non-glandular trichomes of Pinguicula contain food reserves and thus are a reward for pollinators. The trichomes from the Pinguicula groups, which differ in their taxonomy (species from the subgenera: Temnoceras, Pinguicula and Isoloba) as well as the types of their pollinators (butterflies/flies and bees/hummingbirds), were examined. Thus, it was determined whether there are any connections between the occurrence of food trichomes and phylogeny position or pollination biology. Additionally, we determined the phylogenetic history of edible trichomes and pollinator evolution in the Pinguicula species.MethodsThe species that were sampled were: Pinguicula moctezumae, P. esseriana, P. moranensis, P. emarginata, P. rectifolia, P. mesophytica, P. hemiepiphytica, P. agnata, P. albida, P. ibarrae, P. martinezii, P. filifolia, P. gigantea, P. lusitanica, P. alpina and P. vulgaris. Light microscopy, histochemistry, and scanning and transmission electron microscopy were used to address our aims with a phylogenetic perspective based on matK/trnK DNA sequences.Key ResultsNo accumulation of protein bodies or lipid droplets was recorded in the floral non-glandular trichomes of any of the analysed species. Starch grains occurred in the cells of the trichomes of the bee-/fly-pollinated species: P. agnata, P. albida, P. ibarrae, P. martinezii, P. filifolia and P. gigantea, but not in P. alpina or P. vulgaris. Moreover, starch grains were not recorded in the cells of the trichomes of the Pinguicula species that have long spurs, which are pollinated by Lepidoptera (P. moctezumae, P. esseriana, P. moranensis, P. emarginata and P. rectifolia) or birds (P. mesophytica and P. hemiepihytica), or in species with a small and whitish corolla that self-pollinate (P. lusitanica). The results on the occurrence of edible trichomes and pollinator syndromes were mapped onto a phylogenetic reconstruction of the genus.ConclusionFloral non-glandular trichomes play the role of edible trichomes in some Pinguicula species (P. agnata, P. albida, P. ibarrae, P. martinezii, P. filifolia and P. gigantea), which are mainly classified as bee-pollinated species that had originated from Central and South America. It seems that in the Pinguicula that are pollinated by other pollinator groups (Lepidoptera and hummingbirds), the non-glandular trichomes in the flowers play a role other than that of a floral reward for their pollinators. Edible trichomes are symplesiomorphic for the Pinguicula species, and thus do not support a monophyletic group such as a synapomorphy. Nevertheless, edible trichomes are derived and are possibly a specialization for fly and bee pollinators by acting as a food reward for these visitors.

Sexual dimorphism in floral scents of the neotropical orchid Catasetum arietinum and its possible ecological and evolutionary significance

Abstract Dioecy in angiosperms is often associated with sexual dimorphism in floral traits other than the sexual organs. Species of the neotropical orchid genus Catasetum produce unisexual flowers characterized by a remarkable morphological sexual dimorphism. Catasetum species emit strong floral perfumes that act as both signal and reward for male euglossine bee pollinators. Although the role of floral perfumes of Catasetum in attracting euglossine pollinators is well investigated, little is known about whether perfumes differ between floral sexes and, if they do, whether this chemical dimorphism influences the pollination ecology of the plants. Taking Catasetum arietinum as a model species, our aim was to observe the behaviour of pollinators on male and female flowers and to compare scent properties (i.e. chemical composition, total amount and temporal fluctuation) of male and female flowers. Floral scent samples were collected by using dynamic headspace methods and were analysed via gas chromatography coupled with mass spectroscopy (GC-MS). Catasetum arietinum is pollinated by males of two Euglossa species (i.e. E. nanomelanotricha and E. securigera). Bees approached male and female inflorescences of C. arietinum in similar proportions but landed significantly more often and spent more time on female flowers, which emitted more scent than male flowers. Furthermore, the amount of scent emitted varied across the different times of sampling, corresponding to the pattern of the diel foraging activity of pollinating bees on male and female flowers. The chemical composition of scents differed significantly between sexes. The two major compounds (Z)-methyl-p-methoxycinnamate and (E)-geranyl geraniol contributed most to this difference. This is the first case of sexual dimorphism reported in orchid floral perfumes. We discuss the influence of sex-specific floral scents on the behaviour of euglossine pollinators and offer new insights into the ecological and evolutionary significance of divergence in floral scents among dioecious plants.

Dimorphic flowers modify the visitation order of pollinators from male to female flowers

Sexual dimorphism is a pervasive form of variation within species. Understanding how and why sexual dimorphism evolves would contribute to elucidating the mechanisms underlying the diversification of traits. In flowering plants, pollinators are considered a driver of sexual dimorphism when they affect female and male plant fitness in distinct ways. Here, we found that flowers appear to manipulate the behavior of pollinators using sexually dimorphic traits in the dioecious tree Eurya japonica. In this plant, female flowers present a higher-quality reward for pollinators, whereas male flowers have a more conspicuous appearance. Plants benefit by inducing pollinators to carry pollen from male to female flowers, and their sexual dimorphism might thus facilitate pollen movement through pollinator behavior. In two-choice experiments, pollinators frequently moved from male to female flowers, whereas computer simulation suggested that sexually dimorphic traits would evolve if pollinators changed behavior depending on the traits of the flowers they had just visited. These results suggest that the floral traits affecting the visiting order of pollinators have evolved in plants. Using E. japonica, we theoretically show that the induction of sequential behavior in pollinators might be crucial to the evolution of sexual dimorphism in flowers, and our experiments support these findings.

Male flowers of Aconitum compensate for toxic pollen with increased floral signals and rewards for pollinators

Many plants require animal pollinators for successful reproduction; these plants provide pollinator resources in pollen and nectar (rewards) and attract pollinators by specific cues (signals). In a seeming contradiction, some plants produce toxins such as alkaloids in their pollen and nectar, protecting their resources from ineffective pollinators. We investigated signals and rewards in the toxic, protandrous bee-pollinated plant Aconitum napellus, hypothesizing that male-phase flower reproductive success is pollinator-limited, which should favour higher levels of signals (odours) and rewards (nectar and pollen) compared with female-phase flowers. Furthermore, we expected insect visitors to forage only for nectar, due to the toxicity of pollen. We demonstrated that male-phase flowers emitted more volatile molecules and produced higher volumes of nectar than female-phase flowers. Alkaloids in pollen functioned as chemical defences, and were more diverse and more concentrated compared to the alkaloids in nectar. Visitors actively collected little pollen for larval food but consumed more of the less-toxic nectar. Toxic pollen remaining on the bee bodies promoted pollen transfer efficiency, facilitating pollination.

Diversity of nectar amino acids in the Fritillaria (Liliaceae) genus: ecological and evolutionary implications

Nectar is considered to be a primary food reward for most pollinators. It mostly contains sugars, but also has amino acids. The significance of the concentration and composition of amino acids in nectar is often less understood than that of its volume, sugar concentration and composition. However, there is a trend towards a broader approach in ecological research, which helps to understand nectar properties in an ecological context. The genus Fritillaria, exhibiting great diversity in flower morphology, nectar composition, and dominant pollinators, allows for the possibility to study some of the above. We studied the concentration and composition of amino acids in the nectar of 38 Fritillaria species attracting different groups of pollen vectors (bees, flies, passerines, and hummingbirds). The flowers of fritillaries produced nectar with a varying composition and concentration of amino acids. These differences were mostly associated with the pollinator type. The nectar of passerine bird-pollinated species was rich in amino acids, whereas humming bird-pollinated produced low amino acid nectar. Contrary to previous reports nectar of the insect-pollinated species did not contain a higher amount of proline. Two non-protein amino acids, sarcosine and norvaline, were detected in the floral nectar for the first time.

Floral micromorphology and nectar composition of the early evolutionary lineage Utricularia (subgenus Polypompholyx, Lentibulariaceae)

Utricularia (Lentibulariaceae) is a genus comprising around 240 species of herbaceous, carnivorous plants. Utricularia is usually viewed as an insect-pollinated genus, with the exception of a few bird-pollinated species. The bladderworts Utricularia multifida and U. tenella are interesting species because they represent an early evolutionary Utricularia branch and have some unusual morphological characters in their traps and calyx. Thus, our aims were to (i) determine whether the nectar sugar concentrations and composition in U. multifida and U. tenella are similar to those of other Utricularia species from the subgenera Polypompholyx and Utricularia, (ii) compare the nectary structure of U. multifida and U. tenella with those of other Utricularia species, and (iii) determine whether U. multifida and U. tenella use some of their floral trichomes as an alternative food reward for pollinators. We used light microscopy, histochemistry, and scanning and transmission electron microscopy to address those aims. The concentration and composition of nectar sugars were analysed using high-performance liquid chromatography. In all of the examined species, the floral nectary consisted of a spur bearing glandular trichomes. The spur produced and stored the nectar. We detected hexose-dominated (fructose + glucose) nectar in U. multifida and U. tenella as well as in U. violacea. In both U. multifida and U. tenella, there were trichomes that blocked the entrance into the throat and spur. Because these trichomes were rich in chromoplasts and contained lipid droplets, they may form an additional visual attractant. Bearing in mind the phylogenetic hypothesis for the genus, we suggest that an early ancestor of Utricularia had a nectariferous spur flower with a lower lip that formed a wide landing platform for bee pollinators.

Subtle Chemical Variations with Strong Ecological Significance: Stereoselective Responses of Male Orchid Bees to Stereoisomers of Carvone Epoxide.

Different enantiomers of chiral compounds within floral perfumes usually trigger distinct responses in insects; however, this has frequently been neglected in studies investigating semiochemicals in plant-pollinator interactions. Approximately 1000 neotropical plants produce floral perfumes as the only reward for pollinators, i.e. male euglossine bees. The chiral compound carvone epoxide is a key component of the scent bouquet of many perfume-rewarding plants that are pollinated by males of Eulaema. Here, we tested the biological activity of the four carvone epoxide stereoisomers to four Eulaema species occurring in the Atlantic Rainforest of NE-Brazil. We determined the stereochemistry of carvone epoxide in the floral scent of several Catasetum species, tested whether the antennae of bees respond differentially to these stereoisomers and investigated if there is a behavioural preference for any of the stereoisomers. We found that 1) Catasetum species emit only the (-)-trans-stereoisomer of carvone epoxide, 2) for E. atleticana and E. niveofasciata antennal responses to the (-)-trans-carvone epoxide were significantly stronger than those to (-)-cis-carvone epoxide, 3) the strength and pattern of antennal responses to all 4 stereoisomers (separately tested) did not differ among Eulaema species, and 4) there were significant differences in attractiveness of the four stereoisomers to the bees species with the (-)-trans-stereoisomer being particularly attractive. We assume (-)-trans-carvone epoxide to be the dominant isomer in perfume-rewarding plants pollinated by Eulaema. The universal occurrence of carvone epoxide in Catasetum species pollinated by Eulaema, suggests that this compound has evolved in perfume-rewarding as a specific attractant for Eulaema bees as pollinators.

Pollen and vegetative secondary chemistry of three pollen-rewarding lupines.

Optimal defense theory predicts that selection should drive plants to disproportionally allocate resources for herbivore defense to tissues with high fitness values. Because pollen's primary role is the transport of gametes, plants may be expected to defend it from herbivory. However, for many animal-pollinated plants, pollen serves a secondary role as a pollinator reward. These dual roles may present a conflict between selection to defend pollen from herbivores and selection to reward pollinators. Here, we investigate whether pollen secondary chemistry in three pollen-rewarding Lupinus species better reflects the need to defend pollen or reward pollinators. Lupinus (Fabaceae) species are nectarless, pollen-rewarding, and produce defensive quinolizidine and/or piperidine alkaloids throughout their tissues. We used gas chromatography to identify and quantitate the alkaloids in four aboveground tissues (pollen, flower, leaf, stem) of three western North American lupines, L. argenteus, L. bakeri, and L. sulphureus, and compared alkaloid concentrations and composition among tissues within individuals. In L. argenteus and L. sulphureus, pollen alkaloid concentrations were 11-35% of those found in other tissues. We detected no alkaloids in L. bakeri pollen, though they were present in other tissues. Alkaloid concentrations were not strongly correlated among tissues within individuals. We detected fewer alkaloids in pollen compared to other tissues, and pollen contained no unique alkaloids. Our results are consistent with the hypothesis that, in these pollen-rewarding species, pollen secondary chemistry may reflect the need to attract and reward pollinators more than the need to defend pollen from herbivory.

Secondary metabolites from nectar and pollen: a resource for ecological and evolutionary studies.

Floral chemistry mediates plant interactions with herbivores, pathogens, and pollinators. The chemistry of floral nectar and pollen, the primary food rewards for pollinators, can affect both plant reproduction and pollinator health. Although the existence and functional significance of nectar and pollen secondary metabolites has long been known, comprehensive quantitative characterizations of secondary chemistry exist for only a few species. Moreover, little is known about intraspecific variation in nectar and pollen chemical profiles. Because the ecological effects of secondary chemicals are dose-dependent, heterogeneity across genotypes and populations could influence floral trait evolution and pollinator foraging ecology. To better understand within- and across-species heterogeneity in nectar and pollen secondary chemistry, we undertook exhaustive LC-MS and LC-UV-based chemical characterizations of nectar and pollen methanol extracts from 31 cultivated and wild plant species. Nectar and pollen were collected from farms and natural areas in Massachusetts, Vermont, and California, USA, in 2013 and 2014. For wild species, we aimed to collect 10 samples from each of three sites. For agricultural and horticultural species, we aimed for 10 samples from each of three cultivars. Our data set (1,535 samples, 102 identified compounds) identifies and quantifies each compound recorded in methanolic extracts, and includes chemical metadata that describe the molecular mass, retention time, and chemical classification of each compound. A reference phylogeny is included for comparative analyses. We found that each species possessed a distinct chemical profile; moreover, within species, few compounds were found in both nectar and pollen. The most common secondary chemical classes were flavonoids, terpenoids, alkaloids and amines, and chlorogenic acids. The most common compounds were quercetin and kaempferol glycosides. Pollens contained high concentrations of hydroxycinnamoyl-spermidine conjugates, mainly triscoumaroyl and trisferuloyl spermidine, found in 71% of species. When present, pollen alkaloids and spermidines had median nonzero concentrations of 23,000μmol/L (median 52% of recorded micromolar composition). Although secondary chemistry was qualitatively consistent within each species and sample type, we found significant quantitative heterogeneity across cultivars and sites. These data provide a standard reference for future ecological and evolutionary research on nectar and pollen secondary chemistry, including its role in pollinator health and plant reproduction. Data are published under a Creative Commons Attribution License (CC BY 3.0 US) and may be freely used if properly cited.

What Do Nectarivorous Bats Like? Nectar Composition in Bromeliaceae With Special Emphasis on Bat-Pollinated Species.

Floral nectar is the most important reward for pollinators and an integral component of the pollination syndrome. Nectar research has mainly focused on sugars or amino acids, whereas more comprehensive studies on the nectar composition of closely related plant species with different pollination types are rather limited. Nectar composition as well as concentrations of sugars, amino acids, inorganic ions, and organic acids were analyzed for 147 species of Bromeliaceae. This plant family shows a high diversity in terms of floral morphology, flowering time, and predominant pollination types (trochilophilous, trochilophilous/entomophilous, psychophilous, sphingophilous, chiropterophilous). Based on the analyses, we examined the relationship between nectar traits and pollination type in this family. Nectar of all analyzed species contained high amounts of sugars with different proportions of glucose, fructose, and sucrose. The total concentrations of amino acids, inorganic cations, and anions, or organic acids were much lower. The analyses revealed that the sugar composition, the concentrations of inorganic cations and anions as well as the concentration of malate in nectar of bat-pollinated species differed significantly from nectar of species with other pollination types. Flowers of bat-pollinated species contained a higher volume of nectar, which results in a total of about 25-fold higher amounts of sugar in bat-pollinated species than in insect-pollinated species. This difference was even higher for amino acids, inorganic anions and cations, and organic acids (between 50 and 100-fold). In general, bat-pollinated plant species invest large amounts of organic and inorganic compounds for their pollinators. Furthermore, statistical analyses reveal that the characteristics of nectar in Bromeliaceae are more strongly determined by the pollinator type rather than by taxonomic groups or phylogenetic relations. However, a considerable part of the variance cannot be explained by either of the variables, which means that additional factors must be responsible for the differences in the nectar composition.

Yeast-Bacterium Interactions: The Next Frontier in Nectar Research.

Beyond its role as a reward for pollinators, floral nectar also provides a habitat for specialized and opportunistic yeasts and bacteria. These microbes modify nectar chemistry, often altering mutualistic relationships between plants and pollinators in ways that we are only beginning to understand. Many studies on this multi-partite system have focused on either yeasts or bacteria without consideration of yeast-bacterium interactions, but recent evidence suggests that such interactions drive the assembly of nectar microbial communities and its consequences for pollination. Unexplored potential mechanisms of yeast-bacterium interactions include the formation of physical complexes, nutritional interactions, antibiosis, signaling-based interactions, and horizontal gene transfer. We argue that studying these mechanisms can elucidate how nectar microbial communities are established and affect plant fitness via pollinators.

Phenolic profile of pollen collected from different ‘Oblačinska’ sour cherry (Prunus cerasus L.) clones

The presence of pollinators in orchards is crucial to obtain high fruit set and yields of fruits. Despite the fact that sour cherry cultivars are mainly autogamous, insect visits are still of great importance for their propagation. In order to attract and reward pollinators, flowers have to provide adequate nourishment to them. Besides nectar, bees gather pollen, which are a prerequisite for normal colony growth and development of their broods. ‘Oblacinska’ sour cherry (Prunus cerasus L.), an autochthonous cultivar, is the most highly planted cultivar in Serbian commercial orchards. Since the cultivar is actually a mixture of different clones, variability in numerous traits and, particularly, its yields has been reported. Since phenolic compounds are considered to be fundamental pollen chemicals, the aim of this study was to determine the phenolic compounds profile in pollen collected from 15 ‘Oblacinska’ sour cherry clones with varying productivity levels. Solid phase extraction (SPE), combined with ultra-high-performance liquid chromatography coupled with a diode array detector and a triple quadruple mass spectrometer (UHPLC DAD-MS/MS), was used to analyse the polyphenolic profile of pollen. Among 23 components quantified, rutin was the most abundant phenolic compound. It ranged from 98.49 (clone V/P) to 358.83 mg kg(-1) (clone III/9) and was observed to contribute, on average, 56% of the total phenolic compounds in pollen as quantified in different ‘Oblacinska’ sour cherry clones. In addition to this compound, clones contained significant amounts of chlorogenic acid (12.92%), astragalin (8.19%), and hyperoside (5.59%) as well. Cluster analysis grouped pollen clones in four different clusters, which showed that clones III/9, IV/8, and V/P had unique phenolic profiles. Despite the significant differences among the studied clones, the contents of chlorogenic acid, rutin, naringin, hyperoside, astralgin, and phlorizin were distinguishable between the clusters.

Osmophores and floral fragrance in Anacardium humile and Mangifera indica (Anacardiaceae): an overlooked secretory structure in Sapindales.

Flowers of Anacardiaceae and other Sapindales typically produce nectar, but scent, often associated with a reward for pollinators, has surprisingly been mentioned only rarely for members of the family and order. However, flowers of Anacardium humile and Mangifera indica produce a strong sweet scent. The origin and composition of these floral scents is the subject of this study. Screening of potential osmophores on the petals and investigations of their anatomy were carried out by light, scanning and transmission electron microscopy. The composition of the floral fragrance was characterized by gas chromatography–mass spectrometry. In both species, the base of the adaxial side of each petal revealed specialized secretory epidermal cells which are essentially similar in structure and distinct from all other neighbouring cells. These cells also showed evidence of granulocrine secretory mechanisms and slight specific variations in their subcellular apparatus coinciding with the respective composition of the floral fragrance, predominantly composed of sesquiterpenes in A. humile and monoterpenes in M. indica. This study reports the presence of osmophores for the first time in flowers of Anacardiaceae and confirms the link between the ultrastructural features of their secretory cells and the volatiles produced by the flowers. The flowers of most Sapindales, including Anacardiaceae, are nectariferous. However, the presence of osmophores has only been described for very few genera of Rutaceae and Sapindaceae. Both the occurrence of osmophores and fragrance may have largely been overlooked in Anacardiaceae and Sapindales until now. Further studies are needed to better understand the nature and diversity of the interactions of their nectariferous flowers with their pollinators.

Ant-Pollinator Conflict Results in Pollinator Deterrence but no Nectar Trade-Offs.

Direct and indirect negative interactions between ant guards and pollinators on ant-plants are expected for two reasons. First, aggressive ants may deter pollinators directly. Second, pollinators benefit from plant investment in reproduction whilst ants benefit from plant investment in indirect defense, and resource allocation trade-offs between these functions could lead to indirect conflict. We explored the potential for ant-pollinator conflict in a Mexican myrmecophile, Turnera velutina, which rewards ants with extrafloral nectar and pollinators with floral nectar. We characterized the daily timing of ant and pollinator activity on the plant and used experiments to test for direct and indirect conflict between these two groups of mutualists. We tested for direct conflict by quantifying pollinator responses to flowers containing dead specimens of aggressive ant species, relative to unoccupied control flowers. We assessed indirect conflict by testing for the existence of a trade-off in sugar allocation between ant and pollinator rewards, evidenced by an increase in floral nectar secretion when extrafloral nectar secretion was prevented. Secretion of floral and extrafloral nectar, activity of ants and pollinators, and pollen deposition all overlapped in daily time and peaked within the first 2 h after flowers opened. We found evidence of direct conflict, in that presence of ants inside the flowers altered pollinator behavior and reduced visit duration, although visit frequency was unchanged. We found no evidence for indirect conflict, with no significant difference in the volume or sugar content of floral nectar between control plants and those in which extrafloral nectar secretion was prevented. The presence of ants in flowers alters pollinator behavior in ways that are likely to affect pollination dynamics, though there is no apparent trade-off between plant investment in nectar rewards for pollinators and ant guards. Further studies are required to quantify the effect of the natural abundance of ants in flowers on pollinator behavior, and any associated impacts on plant reproductive success.

Impact of flower rewards on phytophagous insects: importance of pollen and nectar for the development of the pollen beetle (Brassicogethes aeneus)

Entomophilous plants reward pollinators with provision of nutrient-rich foods such as pollen and nectar. These rewards contain compounds that are essential to insect development and can be used by pollinators as well as herbivorous insects. The pollen beetle (Brassicogethes aeneus, syn. Meligethes aeneus) whose larvae develop in oilseed rape flowers (Brassica napus) is known to feed on pollen. Previous studies already showed the importance of pollen on the development of this insect but it seems that other resource, such as nectar, could also be used. The purpose of this study was to assess the respective roles of pollen and nectar on pollen beetle development. We tested their role with behavioural and developmental experiments using flowers where the presence and absence of nectar and pollen varied. Larvae, irrespective of their instar, fed both on anthers and nectar. Nectar did not influence larval development or adult survival while pollen influenced development by increasing both larval and adult weight. However, pollen did not affect larval or adult survival nor development time. These results indicate that pollen beetle larvae are adapted to deal with various diets and can complete their development without pollen or nectar.

Using Nectar-Related Traits to Enhance Crop-Pollinator Interactions

Floral nectar and other reward facilitate crop pollination, and in so doing, increase the amount and breadth of food available for humans. Though abundance and diversity of pollinators (particularly bees) have declined over the past several decades, a concomitant increase in reliance on pollinators presents a challenge to food production. Development of crop varieties with specific nectar or nectar-related traits to attract and retain pollinating insects is an appealing strategy to help address needs of agriculture and pollinators for several reasons. First, many crops have specific traits which have been identified to enhance crop–pollinator interactions. Also, an improved understanding of mechanisms that govern nectar-related traits suggest simplified phenotyping and breeding are possible. Finally, the use of nectar-related traits to enhance crop pollination should complement other measures promoting pollinators and will not limit options for crop production or require any changes by growers (other than planting varieties that are more attractive or rewarding to pollinators). In this article, we review the rationale for improving crop-pollinator interactions, the effects of specific plant traits on pollinator species, and use cultivated sunflowers as a case study. Recent research in sunflower has (i) associated variation in bee visitation with specific floral traits, (ii) quantified benefits of pollinators to hybrid yields, and (iii) used genetic resources in sunflower and other plants to find markers associated with key floral traits. Forthcoming work to increase pollinator rewards should enable sunflower to act as a model for using nectar-related traits to enhance crop–pollinator interactions.