Secondary metabolites in floral nectar reduce parasite infections in bumblebees

The synthesis of secondary metabolites is a hallmark of plant defence against herbivores. These compounds may be detrimental to consumers, but can also protect herbivores against parasites. Floral nectar commonly contains secondary metabolites, but little is known about the impacts of nectar chemistry on pollinators, including bees. We hypothesized that nectar secondary metabolites could reduce bee parasite infection. We inoculated individual bumblebees with Crithidia bombi, an intestinal parasite, and tested effects of eight naturally occurring nectar chemicals on parasite population growth. Secondary metabolites strongly reduced parasite load, with significant effects of alkaloids, terpenoids and iridoid glycosides ranging from 61 to 81%. Using microcolonies, we also investigated costs and benefits of consuming anabasine, the compound with the strongest effect on parasites, in infected and uninfected bees. Anabasine increased time to egg laying, and Crithidia reduced bee survival. However, anabasine consumption did not mitigate the negative effects of Crithidia, and Crithidia infection did not alter anabasine consumption. Our novel results highlight that although secondary metabolites may not rescue survival in infected bees, they may play a vital role in mediating Crithidia transmission within and between colonies by reducing Crithidia infection intensities.     

Consumption of a nectar alkaloid reduces pathogen load in bumble bees

Diet has a significant effect on pathogen infections in animals and the consumption of secondary metabolites can either enhance or mitigate infection intensity. Secondary metabolites, which are commonly associated with herbivore defense, are also frequently found in floral nectar. One hypothesized function of this so-called toxic nectar is that it has antimicrobial properties, which may benefit insect pollinators by reducing the intensity of pathogen infections. We tested whether gelsemine, a nectar alkaloid of the bee-pollinated plant Gelsemium sempervirens, could reduce pathogen loads in bumble bees infected with the gut protozoan Crithidia bombi. In our first laboratory experiment, artificially infected bees consumed a daily diet of gelsemine post-infection to simulate continuous ingestion of alkaloid-rich nectar. In the second experiment, bees were inoculated with C. bombi cells that were pre-exposed to gelsemine, simulating the direct effects of nectar alkaloids on pathogen cells that are transmitted at flowers. Gelsemine significantly reduced the fecal intensity of C. bombi 7 days after infection when it was consumed continuously by infected bees, whereas direct exposure of the pathogen to gelsemine showed a non-significant trend toward reduced infection. Lighter pathogen loads may relieve bees from the behavioral impairments associated with the infection, thereby improving their foraging efficiency. If the collection of nectar secondary metabolites by pollinators is done as a means of self-medication, pollinators may selectively maintain secondary metabolites in the nectar of plants in natural populations.     

Possible Synergistic Effects of Thymol and Nicotine against Crithidia bombi Parasitism in Bumble Bees

Floral nectar contains secondary compounds with antimicrobial properties that can affect not only plant-pollinator interactions, but also interactions between pollinators and their parasites. Although recent work has shown that consumption of plant secondary compounds can reduce pollinator parasite loads, little is known about the effects of dosage or compound combinations. We used the generalist pollinator Bombus impatiens and its obligate gut parasite Crithidia bombi to study the effects of nectar chemistry on host-parasite interactions. In two experiments we tested (1) whether the secondary compounds thymol and nicotine act synergistically to reduce parasitism, and (2) whether dietary thymol concentration affects parasite resistance. In both experiments, uninfected Bombus impatiens were inoculated with Crithidia and then fed particular diet treatments for 7 days, after which infection levels were assessed. In the synergism experiment, thymol and nicotine alone and in combination did not significantly affect parasite load or host mortality. However, the thymol-nicotine combination treatment reduced log-transformed parasite counts by 30% relative to the control group (P = 0.08). For the experiment in which we manipulated thymol concentration, we found no significant effect of any thymol concentration on Crithidia load, but moderate (2 ppm) thymol concentrations incurred a near-significant increase in mortality (P = 0.054). Our results tentatively suggest the value of a mixed diet for host immunity, yet contrast with research on the antimicrobial activity of dietary thymol and nicotine in vertebrate and other invertebrate systems. We suggest that future research evaluate genetic variation in Crithidia virulence, multi-strain competition, and Crithidia interactions with the gut microbe community that may mediate antimicrobial activities of secondary compounds.     

The nectar alkaloid, gelsemine, does not affect offspring performance of a native solitary bee, Osmia lignaria (Megachilidae)

Abstract.  1. The ecology and evolution of foliar-feeding insects are thought to be closely tied to plant secondary compounds. Although secondary compounds are also abundant in floral nectar, their role in mediating pollinator preference and performance remains relatively unexplored.
2. This study tested the effects of an alkaloid, gelsemine, found in the nectar of Carolina jessamine ( Gelsemium sempervirens L., Loganiaceae), on the performance of a native solitary bee ( Osmia lignaria lignaria Say, Megachilidae). Nectar gelsemine reduces visits from pollinators, including O. lignaria lignaria , and gelsemine is toxic to vertebrates and possibly non-native honey bees ( Apis mellifera L., Apidae). To test the hypothesis that the deterrent effects of nectar gelsemine reflect negative consequences for pollinator performance, O. lignaria lignaria offspring provisions were supplemented with nectar containing different gelsemine concentrations. Effects on larval development time, prepupa cocoon mass, adult emergence, and adult mass were measured.
3. Nectar gelsemine had no effect on any measure of offspring performance. Thus, although gelsemine deters foraging by adult bees, this behaviour did not optimize offspring performance under the experimental conditions of this study. In contrast, sugar added to nectar treatments increased offspring mass.
4. While adult pollinators may avoid nectar with secondary compounds, this could hinder offspring performance by reducing sugar in provisions if nectar is limiting in the environment. Preference-performance trade-offs, which are studied extensively with foliar herbivores, have seldom been tested for pollinating plant consumers. Future studies of nectar secondary compounds and insect pollinator preference and performance may help to integrate studies of foliage-consuming insect herbivores with nectar-consuming insect pollinators.     

Manduca sexta experience high parasitoid pressures in the field but minor fitness costs of consuming plant secondary compounds

1. Plant‐herbivore coevolutionary interactions have led to a range of plant defenses that minimize insect damage and a suite of counter adaptations that allow herbivores to feed on defended plants. Consuming plant secondary compounds results in herbivore growth and developmental costs but can have beneficial effects such as deterrence or harm of parasitoid enemies. Therefore, the role of secondary compounds on herbivore fitness must be considered in the context of the abundance and level of harm from natural enemies and the costs herbivores incur feeding on plant secondary compounds.
2. In this study, I combined field measurements of Cotesia congregata wasp parasitism pressure with detailed measurements of the costs of plant secondary compounds across developmental stages in the herbivore host, Manduca sexta.
3. I show that C. congregata parasitoids exert large negative selective pressures, killing 31%–57% of M. sexta larvae in the field. Manduca sexta developed fastest during instars most at risk for parasitoid oviposition but growth was slowed by consumption of plant secondary compounds. The negative effects of consuming plant secondary compounds as larvae influenced adult size traits but there were no immune, survival, or fecundity costs.
4. These results suggest that developmental costs experienced by M. sexta herbivores consuming defensive compounds are minor in comparison to the strong negative survival pressures from abundant parasitoid enemies.

     

The ecological significance of toxic nectar

Although plant-herbivore and plant-pollinator interactions have traditionally been studied separately, many traits are simultaneously under selection by both herbivores and pollinators. For example, secondary compounds commonly associated with herbivore defense have been found in the nectar of many plant species, and many plants produce nectar that is toxic or repellent to some floral visitors. Although secondary compounds in nectar and toxic nectar are geographically and phylogenetically widespread, their ecological significance is poorly understood. Several hypotheses have been proposed for the possible functions of toxic nectar, including encouraging specialist pollinators, deterring nectar robbers, preventing microbial degradation of nectar, and altering pollinator behavior. All of these hypotheses rest on the assumption that the benefits of toxic nectar must outweigh possible costs; however, to date no study has demonstrated that toxic nectar provides fitness benefits for any plant. Therefore, in addition to these adaptive hypotheses, we should also consider the hypothesis that toxic nectar provides no benefits or is tolerably detrimental to plants, and occurs due to previous selection pressures or pleiotropic constraints. For example, secondary compounds may be transported into nectar as a consequence of their presence in phloem, rather than due to direct selection for toxic nectar. Experimental approaches are necessary to understand the role of toxic nectar in plant-animal interactions.     

The Only African Wild Tobacco,Nicotiana africana: Alkaloid Content and the Effect of Herbivory

Herbivory in some Nicotiana species is known to induce alkaloid production. This study examined herbivore-induced defenses in the nornicotine-rich African tobacco N. africana, the only Nicotiana species indigenous to Africa. We tested the predictions that: 1) N. africana will have high constitutive levels of leaf, flower and nectar alkaloids; 2) leaf herbivory by the African bollworm Helicoverpa armigera will induce increased alkaloid levels in leaves, flowers and nectar; and 3) increased alkaloid concentrations in herbivore-damaged plants will negatively affect larval growth. We grew N. africana in large pots in a greenhouse and exposed flowering plants to densities of one, three and six fourth-instar larvae of H. armigera, for four days. Leaves, flowers and nectar were analyzed for nicotine, nornicotine and anabasine. The principal leaf alkaloid was nornicotine (mean: 28 µg/g dry mass) followed by anabasine (4.9 µg/g) and nicotine (0.6 µg/g). Nornicotine was found in low quantities in the flowers, but no nicotine or anabasine were recorded. The nectar contained none of the alkaloids measured. Larval growth was reduced when leaves of flowering plants were exposed to six larvae. As predicted by the optimal defense theory, herbivory had a localized effect and caused an increase in nornicotine concentrations in both undamaged top leaves of herbivore damaged plants and herbivore damaged leaves exposed to one and three larvae. The nicotine concentration increased in damaged compared to undamaged middle leaves. The nornicotine concentration was lower in damaged leaves of plants exposed to six compared to three larvae, suggesting that N. africana rather invests in new growth as opposed to protecting older leaves under severe attack. The results indicate that the nornicotine-rich N. africana will be unattractive to herbivores and more so when damaged, but that potential pollinators will be unaffected because the nectar remains alkaloid-free even after herbivory.     

Norditerpene alkaloid concentrations in tissues and floral rewards of larkspurs and impacts on pollinators

Plant secondary compounds mediate interactions with insects and other animals. The norditerpene alkaloids are significant secondary compounds in Delphinium (larkspur) species which are divided into two classes: the 7, 8-methylenedioxylycoctonine (MDL-type) and N-(methylsuccinimido) anthranoyllycoctonine (MSAL-type), and are known to be toxic to herbivorous insects and livestock. Alkaloid concentrations were measured in a whole plant context in vegetative and floral tissues as well as rewards (pollen and nectar) in Delphinium barbeyi and Delphinium nuttallianum. Alkaloid concentrations differed between vegetative tissues, floral tissues and floral rewards. Alkaloid concentrations in floral parts were consistent with optimal defense theory, with tissues more closely tied to plant fitness, such as fruits, being more heavily defended than foliage. However, alkaloid concentrations were significantly lower in nectar compared to other tissues. The norditerpene alkaloids influenced the activity of bumble bees, the dominant pollinator of larkspur, but the effects were concentration dependent. Alkaloids in nectar are found at concentrations that have no effect on bee activity; however, if alkaloid concentrations in nectar were similar to those in foliage bee activity would be reduced significantly. These results suggest that nectar with low alkaloid concentrations may be beneficial to plant fitness by limiting adverse effects on pollinator activity.     

Raising the Sugar Content – Orchid Bees Overcome the Constraints of Suction Feeding through Manipulation of Nectar and Pollen Provisions

Unlike most other bees, the long-tongued orchid bees ingest nectar using suction feeding. Although long tongues allow exploitation of flowers with deep spurs, the energy intake rate is optimal at 10–20% lower nectar sugar concentrations compared to that of lapping bees. This constraint might be compensated by a higher digestive throughput. Additionally, orchid bees might evaporate water from regurgitated droplets of crop contents. We found male Euglossa championi (n = 10) and Euglossa dodsoni (n = 12) to regularly regurgitate droplets of crop content to the base of their proboscis, generating a fluid film between the proximal parts of the galeae, glossa and labial palps. Rhythmic movements of the proboscis may help to increase convection. There was a significant change in sugar concentration between the initially imbibed solution and the resulting crop content (P<0.05) and the time individual bees had engaged in this liquid exposure behavior was positively correlated with the resulting crop sugar concentration. Female Euglossa townsendi and Euglossa viridissima showed the same behavior. Additionally, they manipulated their nectar-enriched pollen provisions for extensive periods of time before deposition in brood cells. The deposited pollen loads (n = 14) showed a significantly higher sugar concentration than the sugar-water available to the bees (P<0.001). Thus, both male and female euglossines show behaviors that promote evaporative water loss from nectar. We suggest that the behaviors have evolved in concert with suction feeding on dilute nectar from deep floral tubes.     

Honeybees prefer warmer nectar and less viscous nectar,regardless of sugar concentration

The internal temperature of flowers may be higher than air temperature, and warmer nectar could offer energetic advantages for honeybee thermoregulation, as well as being easier to drink owing to its lower viscosity. We investigated the responses of Apis mellifera scutellata (10 colonies) to warmed 10% w/w sucrose solutions, maintained at 20–35°C, independent of low air temperatures, and to 20% w/w sucrose solutions with the viscosity increased by the addition of the inert polysaccharide Tylose (up to the equivalent of 34.5% sucrose). Honeybee crop loads increased with nectar temperature, as did the total consumption of sucrose solutions over 2 h by all bees visiting the feeders. In addition, the preference of marked honeybees shifted towards higher nectar temperatures with successive feeder visits. Crop loads were inversely proportional to the viscosity of the artificial nectar, as was the total consumption of sucrose solutions over 2 h. Marked honeybees avoided higher nectar viscosities with successive feeder visits. Bees thus showed strong preferences for both warmer and less viscous nectar, independent of changes in its sugar concentration. Bees may benefit from foraging on nectars that are warmer than air temperature for two reasons that are not mutually exclusive: reduced thermoregulatory costs and faster ingestion times due to the lower viscosity.     

Punch in the gut: parasite tolerance of phytochemicals reflects host diet

Gut parasites of plant-eating insects are exposed to antimicrobial phytochemicals that can reduce infection. Trypanosomatid gut parasites infect insects of diverse nutritional ecologies as well as mammals and plants, raising the question of how host diet-associated phytochemicals shape parasite evolution and host specificity. To test the hypothesis that phytochemical tolerance of trypanosomatids reflects the chemical ecology of their hosts, we compared related parasites from honey bees and mosquitoes – hosts that differ in phytochemical consumption – and contrasted our results with previous studies on phylogenetically related, human-parasitic Leishmania. We identified one bacterial and 10 plant-derived substances with known antileishmanial activity that also inhibited honey bee parasites associated with colony collapse. Bee parasites exhibited greater tolerance of chrysin – a flavonoid found in nectar, pollen and plant resin-derived propolis. In contrast, mosquito parasites were more tolerant of cinnamic acid – a product of lignin decomposition present in woody debris-rich larval habitats. Parasites from both hosts tolerated many compounds that inhibit Leishmania, hinting at possible trade-offs between phytochemical tolerance and mammalian infection. Our results implicate the phytochemistry of host diets as a potential driver of insect–trypanosomatid associations and identify compounds that could be incorporated into colony diets or floral landscapes to ameliorate infection in bees.     

Effect of timing and exposure of sunflower pollen on a common gut pathogen of bumble bees

1. Several bee species are declining due to multiple factors, including pathogens. Ingestion of sunflower (Helianthus annuus) pollen can dramatically reduce the bumble bee gut pathogen Crithidia bombi, but little is known about how timing and exposure to sunflower pollen consumption affects pathogen load. 2. Two experiments were carried out to investigate how exposure to sunflower pollen relative to pathogen exposure affects Crithidia bombi in Bombus impatiens. Foraging trials with pollen‐producing and male‐sterile (pollen absent) sunflower lines were performed to investigate whether sunflower pollen affected pathogen transmission in a single foraging bout, and 7‐day laboratory trials were done to investigate whether timing and duration of exposure to sunflower pollen after infection affected C. bombi. 3. In foraging trials, pollen presence on inflorescences inoculated with C. bombi did not affect transmission (pathogen cell counts of foraging workers) 1 week later, suggesting that a brief experience with sunflower pollen concurrent with pathogen exposure is insufficient to reduce infection. In laboratory trials, consuming sunflower pollen for the first 3.5 days or all 7 days after infection reduced cell counts compared with a negative control pollen, but consuming sunflower pollen starting 3.5 days after infection did not. Consuming sunflower pollen for 7 days was significantly and substantially more effective than any other treatment. Thus, both duration and timing of exposure to sunflower pollen may affect pathogen load. 4. These results are important for understanding ecological disease dynamics in natural settings with free‐flying bumble bees, and may inform decisions about using medicinal diets to manage bumble bee health commercially.     

Crop Pollination Exposes Honey Bees to Pesticides Which Alters Their Susceptibility to the Gut Pathogen Nosema ceranae

Recent declines in honey bee populations and increasing demand for insect-pollinated crops raise concerns about pollinator shortages. Pesticide exposure and pathogens may interact to have strong negative effects on managed honey bee colonies. Such findings are of great concern given the large numbers and high levels of pesticides found in honey bee colonies. Thus it is crucial to determine how field-relevant combinations and loads of pesticides affect bee health. We collected pollen from bee hives in seven major crops to determine 1) what types of pesticides bees are exposed to when rented for pollination of various crops and 2) how field-relevant pesticide blends affect bees’ susceptibility to the gut parasite Nosema ceranae. Our samples represent pollen collected by foragers for use by the colony, and do not necessarily indicate foragers’ roles as pollinators. In blueberry, cranberry, cucumber, pumpkin and watermelon bees collected pollen almost exclusively from weeds and wildflowers during our sampling. Thus more attention must be paid to how honey bees are exposed to pesticides outside of the field in which they are placed. We detected 35 different pesticides in the sampled pollen, and found high fungicide loads. The insecticides esfenvalerate and phosmet were at a concentration higher than their median lethal dose in at least one pollen sample. While fungicides are typically seen as fairly safe for honey bees, we found an increased probability of Nosema infection in bees that consumed pollen with a higher fungicide load. Our results highlight a need for research on sub-lethal effects of fungicides and other chemicals that bees placed in an agricultural setting are exposed to.     

Trichomes as dangerous lollipops: Do lizards also use caterpillar body and frass odor to optimize their foraging?

When attacked by herbivores, plants produce toxic secondary metabolites that function as direct defenses, as well as indirect defenses that attract and reward predators of the offending herbivores. These indirect defenses include both nutritive rewards such as extra floral nectar, as well as informational rewards, such as the production and release of volatile compounds that betray the location of feeding herbivores to predators. Herbivory of Nicotiana attenuata by the tobacco hornworm (Manduca larvae) alters the volatile profiles of both the plant and larval headspace. Herbivory-elicited specific changes in the volatile profiles are detected by arthropod predators of Manduca larvae. The known predators that perceive volatile cues induced by Manduca herbivory of N. attenuata are insects that target Manduca at early developmental stages, when the larvae are still small; large, late-instar larvae may have outgrown these predation risks. However, here we offer evidence that branched chain aliphatic acids derived from the digestion of plant O-acyl sugars from trichomes may betray Manduca larvae to lizard predators during late developmental stages as well.     

Survival and health improvement of Nosema infected Apis florea (Hymenoptera: Apidae) bees after treatment with propolis extract

Nosema ceranae is now considered to be an emerging infectious disease of the European honey bee Apis mellifera. Only one antibiotic, Fumagillin, is commercially available to combat Nosema infections. This antibiotic treatment is banned from use in Europe and elsewhere there is a high probability for antibiotic resistance to develop. We are therefore interested in investigating the effects of a natural propolis extract on its ability to reduce N. ceranae infection loads in the dwarf honey bee, Apis florea, a native honey bee with a range that overlaps with Apis cerana and Apis mellifera that is at risk of infection. Experimentally infected caged bees were fed a treatment consisting of 0%, 50%, or 70% propolis extract. All 50% and 70% propolis treated bees had significantly lower infection loads, and the 50% treated bees had higher survival in comparison to untreated bees. In addition, propolis treated bees had significantly higher haemolymph trehalose levels and hypopharyngeal gland protein content similar to levels of uninfected bees. Propolis ethanolic extract treatment could therefore be considered as a possible viable alternative to Fumagillin to improve bee health. This natural treatment deserves further exploration to develop it as a possible alternative to combat N. ceranae infections distributed around the world.     

Can alloethism in workers of the bumblebee, Bombus terrestris, be explained in terms of foraging efficiency?

Bumblebee workers vary greatly in size, unlike workers of most other social bees. This variability has not been adequately explained. In many social insects, size variation is adaptive, with different-sized workers performing different tasks (alloethism). Here we established whether workers of the bumblebee, Bombus terrestris (L.) (Hymenoptera; Apidae), exhibit alloethism. We quantified the size of workers engaging in foraging compared to those that remain in the nest, and confirmed that it is the larger bees that tend to forage (X±SE thorax widths 4.34±0.01 mm for nest bees and 4.93±0.02 mm for foragers). We then investigated whether large bees are better suited to foraging because they are able to transport heavier loads of food back to the nest. Both pollen and nectar loads of returning foragers were measured, demonstrating that larger bees do return with a heavier mass of forage. Foraging trip times were inversely related to bee size when collecting nectar, but were unrelated to bee size for bees collecting pollen. Overall, large bees brought back more nectar per unit time than small bees, but the rate of pollen collection appeared to be unrelated to size. The smallest foragers had a nectar foraging rate close to zero, presumably explaining why foragers tend to be large. Why might larger bees be better at foraging? Various explanations are considered: larger bees are able to forage in cooler conditions, may be able to forage over larger distances, and are perhaps also less vulnerable to predation. Conversely, small workers are presumably cheaper to produce and may be more nimble at within-nest tasks. Further research is needed to assess these possibilities. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Post‐ingestive effects of nectar alkaloids depend on dominance status of bumblebees

Abstract 1. Secondary metabolites have acute or chronic post‐ingestive effects on animals, ranging from death to growth inhibition to reduced nutrient assimilation. 2. Although characterised as toxic, the nectar of Gelsemium sempervirens is not lethal to pollinators, even when the concentration of the nectar alkaloid gelsemine is very high. However, little is known about the sublethal costs of nectar alkaloids. 3. Using a microcolony assay and paired worker bumblebees, the present study measured the effects of artificial nectar containing gelsemine on oocyte development. Oocytes are a sensitive indicator of protein utilisation and general metabolic processes. We also calculated carbohydrate concentrations in the haemolymph to examine energetic costs of gelsemine consumption. 4. High concentrations of gelsemine significantly reduced mean oocyte width in subordinate bees, while dominant bees showed only a trend towards oocyte inhibition. Gelsemine consumption did not reduce carbohydrate concentrations in haemolymph. 5. The cost of ingesting gelsemine may be due to direct toxicity of alkaloids or may be an expense associated with detoxifying gelsemine. Detoxification of alkaloids can require reallocation of resources away from essential metabolic functions like reproduction. The risks associated with nectar alkaloid consumption are tied to both the social and nutritional status of the bee.     

Ecologically appropriate xenobiotics induce cytochrome P450s in Apis mellifera

Background

Honey bees are exposed to phytochemicals through the nectar, pollen and propolis consumed to sustain the colony. They may also encounter mycotoxins produced by Aspergillus fungi infesting pollen in beebread. Moreover, bees are exposed to agricultural pesticides, particularly in-hive acaricides used against the parasite Varroa destructor. They cope with these and other xenobiotics primarily through enzymatic detoxificative processes, but the regulation of detoxificative enzymes in honey bees remains largely unexplored.

Methodology/Principal Findings

We used several approaches to ascertain effects of dietary toxins on bee susceptibility to synthetic and natural xenobiotics, including the acaricide tau-fluvalinate, the agricultural pesticide imidacloprid, and the naturally occurring mycotoxin aflatoxin. We administered potential inducers of cytochrome P450 enzymes, the principal biochemical system for Phase 1 detoxification in insects, to investigate how detoxification is regulated. The drug phenobarbital induces P450s in many insects, yet feeding bees with phenobarbital had no effect on the toxicity of tau-fluvalinate, a pesticide known to be detoxified by bee P450s. Similarly, no P450 induction, as measured by tau-fluvalinate tolerance, occurred in bees fed xanthotoxin, salicylic acid, or indole-3-carbinol, all of which induce P450s in other insects. Only quercetin, a common pollen and honey constituent, reduced tau-fluvalinate toxicity. In microarray comparisons no change in detoxificative gene expression was detected in phenobarbital-treated bees. However, northern blot analyses of guts of bees fed extracts of honey, pollen and propolis showed elevated expression of three CYP6AS P450 genes. Diet did not influence tau-fluvalinate or imidacloprid toxicity in bioassays; however, aflatoxin toxicity was higher in bees consuming sucrose or high-fructose corn syrup than in bees consuming honey.

Conclusions/Significance

These results suggest that regulation of honey bee P450s is tuned to chemicals occurring naturally in the hive environment and that, in terms of toxicological capacity, a diet of sugar is not equivalent to a diet of honey.     

Honeydew feeding in the solitary bee Osmia bicornis as affected by aphid species and nectar availability

Like honey bees (Apis mellifera), non-Apis bees could exploit honeydew as a carbohydrate source. In addition to providing carbohydrates, this may expose them to potentially harmful plant products secreted in honeydew. However, knowledge on honeydew feeding by solitary bees is very scarce. Here we determine whether the polylectic solitary bee Osmia bicornis (=O. rufa) collects honeydew under semi-field conditions, and whether this is affected by aphid species and presence of floral nectar. Bees were provided with oilseed rape plants containing flowers and/or colonies of either Myzus persicae or Brevicoryne brassicae. We used the total sugar level of the bee crop as a measure of the individual's nutritional state and the oligosaccharide erlose as indicator for honeydew consumption. Erlose was present in honeydews from both aphid species, while absent in oilseed rape nectar, nor being synthesized by O. bicornis. When bees were confined to a single honeydew type as the only carbohydrate source, consumption of M. persicae honeydew was confirmed for 47% of the bees and consumption of B. brassicae honeydew for only 3%. Increased mortality in the latter treatment provided further evidence that B. brassicae honeydew is an unsuitable food source for O. bicornis. All bees that were given the choice between honeydew and floral nectar showed significantly increased total sugar levels. However, the fact that no erlose was detected in these bees indicates that honeydew was not consumed when suitable floral nectar was available. This study demonstrates that honeydew exploitation by O. bicornis is dependent on honeydew type and the presence of floral nectar.