Brain biogenic amines and reproductive dominance in bumble bees (Bombus terrestris)

To begin to explore the role of biogenic amines in reproductive division of labor in social insects, brain levels of dopamine, serotonin, and octopamine were measured in bumble bee (Bombus terrestris) workers and queens that differ in behavioral and reproductive state. Levels of all three amines were similar for mated and virgin queens. Young workers that developed with or without a queen had similar amine levels, but in queenright colonies differences in biogenic amine levels were associated with differences in behavior and reproductive physiology. Dominant workers had significantly higher octopamine levels compared with workers of lower dominance status but of similar size, age, and ovary state. High dopamine levels were associated with the last stages of oocyte development irrespective of worker social status and behavior. These results suggest that biogenic amines are involved in behavioral and physiological aspects of regulation of reproduction in bumble bees. Accepted: 10 December 1999     

Reproductive division of labor, dominance, and ecdysteroid levels in hemolymph and ovary of the bumble bee Bombus terrestris

To determine whether ecdysteroids are associated with reproductive division of labor in Bombus terrestris, we measured their levels in hemolymph and ovaries of queens and workers. Queens heading colonies had large active ovaries with high ecdysteroid content, whereas virgin gynes and mated queens before and after diapause had undeveloped ovaries with low ecdysteroid content. The hemolymph ecdysteroid titer was rather variable, but in a pooled analysis of mated queens before and after diapause versus colony-heading queens, ecdysteroid titers were higher in the latter group. In workers, agonistic behavior, ovarian activity, ovarian ecdysteroid content, and hemolymph ecdysteroid titers were positively correlated, and were lowest when a queen was present. In queenless workers, ecdysteroid levels were elevated in dominant workers, and were also influenced by the presence of brood and by group demography; hormone levels were higher in bees kept in larger groups. These findings are consistent with the premise that in B. terrestris the ovary is the primary site of ecdysteroid synthesis, and they show that ecdysteroids levels vary with the social environment.     

Small worker bumble bees (Bombus impatiens) are hardier against starvation than their larger sisters

In bumble bees (Bombus spp.), where workers within the same colony exhibit up to a tenfold difference in mass, labor is divided by body size. Current adaptive explanations for this important life history feature are unsatisfactory. Within the colony, what is the function of the smaller workers? Here, we report on the differential robustness to starvation of small and large worker bumble bees (Bombus impatiens); when nectar is scarce, small workers remain alive significantly longer than larger workers. The presence of small workers, and size variation in general, might act as insurance against times of nectar shortage. These data may provide a novel, adaptive explanation, independent of division of labor, for size polymorphism within the worker caste.     

Detection of Deformed wing virus, a honey bee viral pathogen, in bumble bees (Bombus terrestris and Bombus pascuorum) with wing deformities

Honey bees (Apis mellifera) productively infected with Deformed wing virus (DWV) through Varroa destructor (V. destructor) during pupal stages develop into adults showing wing and other morphological deformities. Here, we report for the first time the occurrence of bumble bees (Bombus terrestris, Bombus pascuorum) exhibiting wing deformities resembling those seen in clinically DWV-infected honey bees. Using specific RT-PCR protocols for the detection of DWV followed by sequencing of the PCR products we could demonstrate that the bumble bees were indeed infected with DWV. Since such deformed bumble bees are not viable DWV infection may pose a serious threat to bumble bee populations.     

A comprehensive phylogeny of the bumble bees (Bombus)

Bumble bees ( Bombus Latreille) occupy a wide diversity of habitats, from alpine meadows to lowland tropical forest, yet they appear to be similar in morphology throughout their range, suggesting that behavioural adaptations play a more important role in colonizing diverse habitats. Notwithstanding their structural homogeneity, bumble bees exhibit striking inter- and intraspecific variation in colour pattern, purportedly the outcome of mimetic evolution. A robust phylogeny of Bombus would provide the framework for elucidating the history of their wide biogeographical distribution and the evolution of behavioural and morphological adaptations, including colour pattern. However, morphological studies of bumble bees have discovered too few phylogenetically informative characters to reconstruct a robust phylogeny. Using DNA sequence data, we report the first nearly complete species phylogeny of bumble bees, including most of the 250 known species from the 38 currently recognized subgenera. Bayesian analysis of nuclear (opsin, EF-1α, arginine kinase, PEPCK) and mitochondrial (16S) sequences results in a highly resolved and strongly supported phylogeny from base to tips, with clear-cut support for monophyly of most of the conventional morphology-based subgenera. Most subgenera fall into two distinct clades ( short-faced and long-faced ) associated broadly with differences in head morphology. Within the short-faced clade is a diverse New World clade, which includes nearly one-quarter of the currently recognized subgenera, many of which are restricted to higher elevations of Central and South America. The comprehensive phylogeny provides a firm foundation for reclassification and for evaluating character evolution in the bumble bees.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 161–188.     

The power and efficiency of brood incubation in queenless microcolonies of bumble bees (Bombus terrestris L.)

1. Ground‐nesting colonies of bumble bees incubate their brood at > 30 °C if floral forage provides sufficient energy and the thermogenic power of the colony can counteract cool soil conditions. To explore the basis of incubation, the thermogenic power and sugar consumption of orphaned nests of bumble bee workers (microcolonies) were investigated under laboratory conditions. 2. This study tested experimentally the effect of variation in worker number (ranging from four to 12 adults) on a microcolony's capacity to regulate brood temperature and recover from acute cold exposure. Microcolonies were provided with ad libitum sugar syrup and minimal insulation and maintained at an ambient temperature of c. 25 °C. Energy conversion efficiency was estimated by comparing sugar consumption with the power required for artificial incubation. The joint energetics of foraging and incubation were modelled in wild colonies to explore the effect of colony size and landscape quality on thermoregulation. 3. The results showed that all sizes of microcolonies regulated brood temperature at c. 31 °C under laboratory conditions, which required 96 mW of thermogenic power. It was estimated that individual workers of B. terrestris generated an incubatory power of 35 mW. The smallest microcolonies had the highest conversion efficiency (57%), apparently because few workers were required for incubation. 4. Modelling indicated that small microcolonies of three to seven adult workers have the capacity for normal brood incubation in the wild, but that the minimum viable colony size increases as floral forage becomes poorer or more distant. 5. These preliminary findings suggest the feasibility of identifying the minimum conditions (forage quality, soil temperature, and colony size) necessary for brood incubation by queenright colonies in the wild.     

Sunflower pollen reduces a gut pathogen in worker and queen but not male bumble bees

1. Social insect castes and sexes differ in many ways, including morphology, behavior, and sometimes ploidy level. Recent studies have found that consuming sunflower pollen reduces the gut pathogen Crithidia bombi in workers of the common eastern bumble bee (Bombus impatiens). Here, this work is extended to the reproductive individuals that represent colony fitness – males and queens – to assess if the medicinal effects of sunflower pollen vary with bee caste and sex. 2. This study examined the effect of sunflower pollen compared to a diverse wildflower pollen mix on infection in worker, male, and daughter queen commercial B. impatiens. Bees were infected, fed either sunflower pollen or wildflower pollen for 7 days, and then infection levels were assessed. 3. Compared to wildflower pollen, sunflower pollen dramatically reduced Crithidia infection in workers and daughter queens, but not males. Infection levels were very low for both diets in males; this could be due to low pollen consumption or other mechanisms. 4. Reducing Crithidia infection in young queens before they undergo hibernation is important for population dynamics since infected queens are less likely to survive hibernation, and those that do are less likely to successfully establish a nest the following spring. Because sunflowers bloom in late summer when new queens are emerging, sunflowers could provide an important dietary component for queens during this critical life stage. Deepening our understanding of how diet impacts pathogens in reproductive bees, as well as workers, is crucial to maintain healthy pollinator populations.     

Bumble bees (Bombus terrestris) store both food and information in honeypots

Social insect foragers often transmit information about foodsources to nest mates. In bumble bees (Bombus terrestris), forexample, successful foragers use excited motor displays anda pheromone as communication signals. In addition, bees couldmake use of an indirect pathway of information flow, via thehoney stores. We show here that, indeed, bees in the nest continuouslymonitor honeypots and sample their contents, thus obtaininginformation on supply and demand of nectar. When there is aninflux of nectar into the nest, the colony deploys more workersfor foraging. The number of new foragers depends on sugar concentration.Foragers returning with high-quality sugar solution displaymore "excited runs" on the nest structure. The recruits' response,however, does not depend on modulated behavior by foragers:more workers start to forage with high quality of incoming nectar,even when this nectar is brought by a pipette. Moreover, weshow that the readiness of bees to respond to recruitment signalsor incoming nectar also depends on colony demand. When colonynectar stores are full, the response of bees to equal amountsof nectar influx is smaller than when stores are empty. Whencolony nectar stores are depleted, foragers spend more timerunning excitedly and less time probing pots in the nest andrun with higher average speed, possibly to disperse the alertingpheromone more efficiently. However, more bees respond to nectarinflux to empty stores, whether or not this is accompanied byforager signals. Thus, honeypots serve to store informationas well as food.     

Factors influencing dominance behaviour in queenless bumblebee workers (Bombus terrestris)

ABSTRACT. The establishment of a dominance order within seventy eight groups of four callow workers was investigated. The dominant worker establishes its position by means of overt aggression. Aggression starts 20—40 h after confinement. Dominance order becomes established within 2 days of removal from the colony. Once established, the dominance order is maintained by dominant-subordinate interactions.
The dominant worker starts ovipositing at an age of about 6 days. Subordinates rarely lay eggs. In 25% of the groups the dominant worker is superseded by another worker before any oviposition occurs. Once the dominance order is established, the dominant worker inhibits endocrine activity in subordinates.
Differences in body size and in endocrine activity are thought to be responsible for the ranking of workers in the dominance hierarchy. Ranking seems to depend more on the activity of the corpora allata (CA) than on the activity of the ovaries. Ovariectomized workers do not differ from control workers in the performance of dominance behaviour. Dominant ovariectomized workers do not construct egg cells, but they do defend these if they are present.     

Bombus folsomi and the origin of Philippine bumble bees (Hymenoptera: Apidae)

Abstract The type female of Bombus folsomi (Frison) is redescribed and synomymized with B.ephippiatus Say. It is concluded that the putative type locality of B.folsomi , Mt Kinabalu in Borneo, is in error and that the genus appears to be absent from Borneo. This lends support to the hypothesis that bumble bees immigrated to the Philippines by an unusual northern route through Taiwan. More rigorous testing of this idea will require further cladistic analysis of the genus Bombus .     

Ontogeny of worker body size distribution in bumble bee (Bombus impatiens) colonies

1. Bumble bees exhibit worker size polymorphisms; highly related workers within a colony may vary up to 10‐fold in body mass. As size variation is an important life history feature in bumble bees, the distribution of body sizes within the colony and how it fluctuates over the colony cycle were analysed. 2. Ten commercially purchased colonies of Bombus impatiens (Cresson) were reared in ad libitum conditions. The size of all workers present and newly emerging workers (callows) was recorded each week. 3. The average size of bumble bee workers did not change with colony age, but variation in body size tended to decrease over time. The average size of callows did not change with population size, but did tend to decrease with colony age. In all measures, there was considerable variation among colonies. 4. Colonies of B. impatiens usually produced workers with normally distributed body sizes throughout the colony life cycle. Unlike most polymorphic ants, there was no increase in worker body size with colony age or colony size. This provides the first, quantitative data on the ontogeny of bumble bee worker size distribution. The potential adaptive significance of this size variation is discussed.     

Foraging errors play a role in resource exploration by bumble bees (Bombus terrrestris)

If the cognitive performance of animals reflects their particular ecological requirements, how can we explain appreciable variation in learning ability amongst closely related individuals (e.g. foraging workers within a bumble bee colony)? One possibility is that apparent ‘errors’ in a learning task actually represent an alternative foraging strategy. In this study we investigate the potential relationship between foraging ‘errors’ and foraging success among bumble bee (Bombus terrestris) workers. Individual foragers were trained to choose yellow, rewarded flowers and ignore blue, unrewarded flowers. We recorded the number of errors (visits to unrewarded flowers) each bee made during training, then tested them to determine how quickly they discovered a more profitable food source (either familiar blue flowers, or novel green flowers). We found that error prone bees discovered the novel food source significantly faster than accurate bees. Furthermore, we demonstrate that the time taken to discover the novel, more profitable, food source is positively correlated with foraging success. These results suggest that foraging errors are part of an ‘exploration’ foraging strategy, which could be advantageous in changeable foraging environments. This could explain the observed variation in learning performance amongst foragers within social insect colonies.     

The safety of thiamethoxam to pollinating bumble bees (Bombus terrestris L.) when applied to tomato plants through drip irrigation

Thiamethoxam, mainly sold under the trademark of Actara, is a neonicotinoid widely used in covered vegetables for the control of aphids and whiteflies. In these crops, and particularly in covered tomatoes, bumble-bees are used for cross-pollination as an alternative to labour intensive manual techniques. In this study, made on tomatoes grown in separated greenhouse plots in Murcia, Southern Spain, thiamethoxam was applied through drip irrigation at a rate of 200 g ai/ha, and as a split application of the same rate, to evaluate the effects on pollinating bumble bees compared to a foliar application of a toxic standard. The results showed that the toxic foliar standard had a clear effect on the pollination of tomato flowers, declining to zero pollination two weeks after application, whereas both the single and split drip irrigation applications of Actara had no effect on pollination when compared to the control plots. The count of dead adults and larvae did not show any differences between the treatments, whereas the measurement of sugar water consumption was shown to correlate well with pollination. The consumption of sugar water declined in the toxic standard plots by 69% with respect to the control, whilst the decline in lower dose drip irrigation application was only 3%. In regard to hive weight, and number of adults and brood after destructive sampling; there were no statistical differences between the treatments but a negative effect of the foliar treatment was observed. Based on these results we can conclude that a split application of Actara applied in drip irrigation to the soil/substrate has no effect on the bumble-bees used in tomatoes for pollination.     

Historical biogeography, divergence times, and diversification patterns of bumble bees (Hymenoptera: Apidae: Bombus)

Bumble bees (Bombus) are a cold-adapted, largely alpine group that can elucidate patterns of Holarctic historical biogeography, particularly in comparison to the alpine plants with which they likely coevolved. A recently published molecular phylogeny of bumble bees provides uniquely comprehensive species sampling for exploring historical patterns of distribution and diversification. Using this phylogeny and detailed data on extant distributions, I reconstruct the historical distribution of bumble bees in a temporal framework, estimating divergence times using fossil data and molecular rates derived from the literature. The nearly comprehensive phylogeny allows assessment of the tempo of diversification within the bumble bees using lineage-through-time plots and diversification statistics, which have been performed with special consideration to confidence intervals. These analyses reveal movements of Bombus concordant with geographic and climatic events of the late Cenozoic. The initial diversification of extant bumble bee lineages was estimated at around 25 to 40 Ma, near the Eocene-Oligocene boundary 34 Ma, a period of dramatic global cooling. Dispersal-vicariance analysis (DIVA) predicted an Old World Bombus ancestor, with early diversification events largely restricted to the eastern Old World. The numerous intercontinental dispersal events occurred mostly in the direction of Old World to New World and North America to South America. Early movements from the Palearctic into the Nearctic most likely took place after 20 Ma and may have coincided with a period of Miocene cooling that gave rise to taiga habitat across Beringia. Subsequent dispersal between these regions is estimated to have occurred among boreal and tundra-adapted species mostly in the last 5 million years. Radiations are estimated in both Nearctic and Neotropical regions at approximately 6 to 8 Ma and after 3.5 Ma, concordant with the opening of land corridors between the continents.     

Primer effect of queen pheromone on juvenile hormone biosynthesis in adult worker honey bees

Juvenile hormone synthesis in adult worker honey bees was measured by an in vitro corpora allata bioassay. Adult queenless workers exhibit higher rates of juvenile hormone biosynthesis than queenright workers. Hormone synthesis is not correlated with the volume of the glands. Extract of queen mandibular glands, applied to a dummy, reduces juvenile hormone biosynthesis in caged queenless workers to the level of queenright workers. The same result was obtained with synthetic (E)-9-oxo-2-decenoic acid, the principal component of the queen mandibular gland secretion. This pheromonal primer effect may function as a key regulating element in maintaining eusocial colony homeostasis. The presence of brood does not affect the hormone production of the corpora allata.Abbreviations BSA bovine serum albumin - CA Corpora allata - JH juvenile hormone - 9-ODA (E)-9-oxo-2-decnoic acid     

Variation in gut microbial communities and its association with pathogen infection in wild bumble bees (Bombus)

Bacterial gut symbiont communities are critical for the health of many insect species. However, little is known about how microbial communities vary among host species or how they respond to anthropogenic disturbances. Bacterial communities that differ in richness or composition may vary in their ability to provide nutrients or defenses. We used deep sequencing to investigate gut microbiota of three species in the genus Bombus (bumble bees). Bombus are among the most economically and ecologically important non-managed pollinators. Some species have experienced dramatic declines, probably due to pathogens and land-use change. We examined variation within and across bee species and between semi-natural and conventional agricultural habitats. We categorized as ‘core bacteria'' any operational taxonomic units (OTUs) with closest hits to sequences previously found exclusively or primarily in the guts of honey bees and bumble bees (genera Apis and Bombus). Microbial community composition differed among bee species. Richness, defined as number of bacterial OTUs, was highest for B. bimaculatus and B. impatiens. For B. bimaculatus, this was due to high richness of non-core bacteria. We found little effect of habitat on microbial communities. Richness of non-core bacteria was negatively associated with bacterial abundance in individual bees, possibly due to deeper sampling of non-core bacteria in bees with low populations of core bacteria. Infection by the gut parasite Crithidia was negatively associated with abundance of the core bacterium Gilliamella and positively associated with richness of non-core bacteria. Our results indicate that Bombus species have distinctive gut communities, and community-level variation is associated with pathogen infection.     

Patterns of parasite infection in bumble bees (Bombus spp.) of Northern Virginia

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Caste-specific patterns of flower visitation in bumble bees (Bombus kirbyellus) collecting nectar from Polemonium viscosum

ABSTRACT.

• 1 Foraging routes of worker and queen bumble bees (Bombus kirbyellus Curtis) collecting nectar from flowers of the alpine sky pilot, Polemonium viscosum Nutt., were followed and the corolla tube length, corolla diameter, floral scent, and number of flowers on plants visited or bypassed by bees were monitored. Additionally, the number and proportion of flowers visited per inflorescence and distance flown from each to the next were recorded. Queens and workers differed significantly in choice of flowers. However, intra-inflorescence visitation rates and departure distances were similar between castes. Castes differed in the extent to which visitation reflected patch quality versus individual floral traits.
• 2 Both queens and workers failed to visit skunky-flowered plants more often than they failed to visit sweet-flowered ones, and preferred large over small inflorescences. However, queens visited large-flowered plants more often than small-flowered ones, while workers preferred flowers with shorter corolla tubes, regardless of their diameter. Although a number of studies have documented caste specialization on alternate species of host plants, ours is one of the first to show that morphological preferences promote comparable foraging differences between castes on monospecific plant resources.
• 3 Queens, once on a plant, responded to floral traits by probing more flowers on large inflorescences, as well as on those with broader floral form. Workers did not alter intra-inflorescence visitation rate in response to floral traits.
• 4 For workers, no significant relationship was demonstrated between the likelihood of passing by a plant and the number of flowers probed on the previous inflorescence visited. Thus, workers appeared to accept or reject each plant of P. viscosum independently. However, queens passed by fewer plants when leaving rich inflorescences than poor ones. These results suggest that workers use only individual plant acceptability in choosing which plants to visit, whereas queens base plant choice on patch and individual attributes. Such differences between castes in foraging rules when exploiting the same floral resource have received little attention, and provide insights into the heterogeneity of harvestable reward distributions from the perspective of the forager population.

     

Perception,regulation, and fitness effects of pollen phytosterols in the bumble bee,Bombus terrestris

Premise

Many flowering plants depend on insects for pollination and thus attract pollinators by offering rewards, mostly nectar and pollen. Bee pollinators rely on pollen as their main nutrient source. Pollen provides all essential micro- and macronutrients including substances that cannot be synthesized by bees themselves, such as sterols, which bees need for processes such as hormone production. Variations in sterol concentrations may consequently affect bee health and reproductive fitness. We therefore hypothesized that (1) these variations in pollen sterols affect longevity and reproduction in bumble bees and (2) can thus be perceived via the bees' antennae before consumption.

Methods

We studied the effect of sterols on longevity and reproduction of Bombus terrestris workers in feeding experiments and investigated sterol perception using chemotactile proboscis extension response (PER) conditioning.

Results

Workers could perceive several sterols (cholesterol, cholestenone, desmosterol, stigmasterol, β-sitosterol) via their antennae but not differentiate between them. However, when sterols were presented in pollen, and not as a single compound, the bees were unable to differentiate between pollen differing in sterol content. Additionally, different sterol concentrations in pollen neither affected pollen consumption nor brood development or worker longevity.

Conclusions

Since we used both natural concentrations and concentrations higher than those found in pollen, our results indicate that bumble bees may not need to pay specific attention to pollen sterol content beyond a specific threshold. Naturally encountered concentrations might fully support their sterol requirements and higher concentrations do not seem to have negative effects.