Simple Physical Interactions Yield Social Self-Organization in Honeybees

Social insect colonies show all characteristics of complex adaptive systems (CAS). Their complex behavioral patterns arise from social interactions that are based on the individuals’ reactions to and interactions with environmental stimuli. We study here how social and environmental factors modulate and bias the collective thermotaxis of young honeybees. Therefore, we record their collective decision-making in a series of laboratory experiments and derived a mathematical model of the collective decision-making in young bees from our empirical observations. This model uses only one free parameter that combines the ultimate effects of several aspects of the microscopic individual behavioral mechanisms, such as motion behavior, sensory range, or contact detection, into one single coefficient. We call this coefficient the “social factor.” Our model is capable of capturing the observed aggregation patterns from our empiric experiments with static environments and of predicting the emergent swarm-intelligent behavior of the system in dynamic environments. Besides the fundamental research aspect in studying CAS, our model enables us to predict the effects of a physical stimulus onto the macroscopic collective decision-making that affects several crucial prerequisites for efficient and effective brood production and population growth in honeybee colonies.

Lower pollen nutritional quality delays nest building and egg laying in Bombus terrestris audax micro-colonies leading to reduced biomass gain

The performance of Bombus terrestris micro-colonies fed five diets differing in pollen species composition and level of nine essential amino acids (EAA; leucine, lysine, valine, arginine, isoleucine, phenylalanine, threonine, histidine, methionine) was assessed for 37 days by recording total biomass gain, nest building initiation, brood production (eggs, small and large larvae, pupae, drones), nectar, and pollen collection. Stronger colony performance was linked to higher amino acid levels but no consistent differences in biomass gain were recorded between mono- and poly-species diets. Poorest performance occurred in micro-colonies offered pure oilseed rape (OSR) pollen which contained the lowest EAA levels. Reduced micro-colony development (delayed nest initiation and lower brood production) was related to OSR proportion in the diet and lower EAA levels. Results are discussed in relation to selection of plant species in the design of habitats to promote bee populations.

Lignin concentrations in phloem and outer bark are not associated with resistance to mountain pine beetle among high elevation pines

A key component in understanding plant-insect interactions is the nature of host defenses. Research on defense traits among Pinus species has focused on specialized metabolites and axial resin ducts, but the role of lignin in defense within diverse systems is unclear. We investigated lignin levels in the outer bark and phloem of P. longaeva, P. balfouriana, and P. flexilis; tree species growing at high elevations in the western United States known to differ in susceptibility to mountain pine beetle (Dendroctonus ponderosae; MPB). Pinus longaeva and P. balfouriana are attacked by MPB less frequently than P. flexilis, and MPB brood production in P. longaeva is limited. Because greater lignification of feeding tissues has been shown to provide defense against bark beetles in related genera, such as Picea, we hypothesized that P. longaeva and P. balfouriana would have greater lignin concentrations than P. flexilis. Contrary to expectations, we found that the more MPB-susceptible P. flexilis had greater phloem lignin levels than the less susceptible P. longaeva and P. balfouriana. No differences in outer bark lignin levels among the species were found. We conclude that lignification in Pinus phloem and outer bark is likely not adaptive as a physical defense against MPB.

Experimental increase of worker diversity benefits brood production in ants

Background The reproductive division of labor of eusocial insects, whereby one or several queens monopolize reproduction, evolved in a context of high genetic relatedness. However, many extant eusocial species have developed strategies that decrease genetic relatedness in their colonies, suggesting some benefits of the increased diversity. Multiple studies support this hypothesis by showing positive correlations between genetic diversity and colony fitness, as well as finding effects of experimental manipulations of diversity on colony performance. However, alternative explanations could account for most of these reports, and the benefits of diversity on performance in eusocial insects still await validation. In this study, we experimentally increased worker diversity in small colonies of the ant Lasius niger while controlling for typical confounding factors. Results We found that experimental colonies composed of workers coming from three different source colonies produced more larvae and showed more variation in size compared to groups of workers coming from a single colony. Conclusions We propose that the benefits of increased diversity stemmed from an improved division of labor. Our study confirms that worker diversity enhances colony performance, thus providing a possible explanation for the evolution of multiply mated queens and multiple-queen colonies in many species of eusocial insects.

Traces of a neonicotinoid pesticide stimulates activity in honey bee colonies, but not productivity

How neonicotinoid contamination effects honey bees remains controversial. Studies have yielded contradictory results, and few have examined effects on colony development. Here we report the results of a comprehensive five-year study of the effects of the neonicotinoid imidacloprid on honey bee colonies. Colonies fed 5 ng/g (ppb) imidacloprid showed increased brood production, lower temperature variability, higher CO2 production and had more foragers compared to control colonies. Even so, treatment did not affect adult bee numbers or average hive temperatures, and did not increase food stores, daily food acquisition or colony survivorship. Imidacloprid contamination increased colony metabolism without improving colony productivity, which is consistent with the pharmacological action of imidacloprid as a stimulant. Our findings explain why some studies have reported no, or even positive, effects of neonicotinoids. We provide an explanation for the diverse effects of pesticide contamination on honey bees, and a thorough understanding of how colonies are impacted.

Lignin accumulation in phloem and outer bark is not associated with resistance to mountain pine beetle in high elevation pines

A key component in understanding plant-insect interactions is the nature of host defenses. Research on defense traits among Pinus species has focused on specialized metabolites and axial resin ducts, but the role of lignin in defense within diverse systems is unclear. We investigated lignin levels in the outer bark and phloem of P. longaeva , P. balfouriana , and P. flexilis ; high elevation species in the western United States known to differ in susceptibility to mountain pine beetle ( Dendroctonus ponderosae ; MPB). Relative to P. flexilis , P. longaeva and P. balfouriana are attacked by MPB less frequently, and MPB brood production in P. longaeva is limited. Because greater lignification of feeding tissues has been shown to provide defense against bark beetles in related genera, such as Picea , we hypothesized that P. longaeva and P. balfouriana would have greater lignin concentrations than P. flexilis . Contrary to expectations, we found that the more MPB-susceptible P. flexilis had greater phloem lignin levels than the less susceptible P. longaeva and P. balfouriana . No differences in outer bark lignin levels among the species were found. We conclude that lignification in Pinus phloem and outer bark is likely not adaptive as a physical defense against MPB.

Sublethal concentrations of clothianidin affect honey bee colony growth and hive CO2 concentration

The effects of agricultural pesticide exposure upon honey bee colonies is of increasing interest to beekeepers and researchers, and the impact of neonicotinoid pesticides in particular has come under intense scrutiny. To explore potential colony-level effects of a neonicotinoid pesticide at field-relevant concentrations, honey bee colonies were fed 5- and 20-ppb concentrations of clothianidin in sugar syrup while control colonies were fed unadulterated syrup. Two experiments were conducted in successive years at the same site in southern Arizona, and one in the high rainfall environment of Mississippi. Across all three experiments, adult bee masses were about 21% lower among colonies fed 20-ppb clothianidin than the untreated control group, but no effects of treatment on brood production were observed. Average daily hive weight losses per day in the 5-ppb clothianidin colonies were about 39% lower post-treatment than in the 20-ppb clothianidin colonies, indicating lower consumption and/or better foraging, but the dry weights of newly-emerged adult bees were on average 6–7% lower in the 5-ppb group compared to the other groups, suggesting a nutritional problem in the 5-ppb group. Internal hive CO2 concentration was higher on average in colonies fed 20-ppb clothianidin, which could have resulted from greater CO2 production and/or reduced ventilating activity. Hive temperature average and daily variability were not affected by clothianidin exposure but did differ significantly among trials. Clothianidin was found to be, like imidacloprid, highly stable in honey in the hive environment over several months.

Honeybee colonies compensate for pesticide-induced effects on royal jelly composition and brood survival with increased brood production

Sublethal doses of pesticides affect individual honeybees, but colony-level effects are less well understood and it is unclear how the two levels integrate. We studied the effect of the neonicotinoid pesticide clothianidin at field realistic concentrations on small colonies. We found that exposure to clothianidin affected worker jelly production of individual workers and created a strong dose-dependent increase in mortality of individual larvae, but strikingly the population size of capped brood remained stable. Thus, hives exhibited short-term resilience. Using a demographic matrix model, we found that the basis of resilience in dosed colonies was a substantive increase in brood initiation rate to compensate for increased brood mortality. However, computer simulation of full size colonies revealed that the increase in brood initiation led to severe reductions in colony reproduction (swarming) and long-term survival. This experiment reveals social regulatory mechanisms on colony-level that enable honeybees to partly compensate for effects on individual level.

Experimental increase of worker diversity benefits brood production in ants

The reproductive division of labor of eusocial insects, whereby one or several queens monopolize reproduction, evolved in a context of high genetic relatedness. However, many extant eusocial species have developed strategies that decrease genetic relatedness in their colonies, suggesting some benefits of the increased diversity. Multiple studies support this hypothesis by showing positive correlations between genetic diversity and colony fitness, as well as finding effects of experimental manipulations of diversity on colony performance. However, alternative explanations could account for most of these reports, and the benefits of diversity on fitness in eusocial insects still await validation. In this study, we experimentally increased worker diversity in the ant Lasius niger while controlling for typical confounding factors. We found that experimental colonies composed of workers coming from three different source colonies produced more larvae and showed more variation in size compared to groups of workers coming from a single colony. We propose that the benefits of increased diversity stemmed from an improved division of labor. Our study confirms that worker diversity enhances colony performance, thus providing a possible explanation for the evolution of multiply mated queens and multiple-queen colonies in many species of eusocial insects.