Decentralized control of drone comb construction in honey bee colonies

Honey bee colonies furnish their nests with two types of comb distinguished by cell size: large cells for rearing males (drone comb) and small cells for rearing workers (worker comb). The bees actively regulate the relative quantity of each type, a behavior likely to be important in setting a colony's sex ratio. Experimental analysis of the information pathways and control mechanisms responsible for this regulation found the following results. The amount of drone comb in a nest is governed by negative feedback from drone comb already constructed. This feedback depends on the workers having direct contact with the drone comb in their nest, but does not depend on the queen's contact with the comb. The comb itself, rather than the brood within it, is sufficient to provide the negative feedback, although the brood may also contribute to the effect. These findings show that drone comb regulation does not depend on the queen acting as a centralized information gatherer and behavioral controller. Instead, the evidence points to a decision-making process distributed across the population of worker bees, a control architecture typical of colony organization in honey bees and other large-colony insect societies. Received: 24 May 1997 / Accepted after revision: 30 August 1997     

The honey bee shaking signal: function and design of a modulatory communication signal

This study explores the meaning and functional design of a modulatory communication signal, the honey bee shaking signal, by addressing five questions: (I) who shakes, (II) when do they shake, (III) where do they shake, (IV) how do receivers respond to shaking, and (V) what conditions trigger shaking. Several results confirm the work of Schneider (1987) and Schneider et al. (1986a): (I) most shakers were foragers (at least 83%); (II) shaking exhibited a consistent temporal pattern with bees producing the most signals in the morning (0810–1150 hours) just prior to a peak in waggle dancing activity; and (IV) bees moved faster (by 75%) after receiving a shaking signal. However, this study differs from previous work by providing a long-term, temporal, spatial, and vector analysis of individual shaker behavior. (III) Bees producing shaking signals walked and delivered signals in all areas of the hive, but produced the most shaking signals directly above the waggle dance floor. (IV) Bees responded to the signal by changing their direction of movement. Prior to receiving a signal, bees selected from the waggle dance floor moved, on average, towards the hive exit. After receiving a signal, some bees continued moving towards the exit but others moved directly away from the exit. During equivalent observation periods, non-shaken bees exhibited a strong tendency to move towards the hive exit. (V) Renewed foraging activity after food dearth triggered shaking signals, and, the level of shaking is positively correlated with the duration of food dearth. However, shaking signal levels also increased in the morning before foraging had begun and in the late afternoon after foraging had ceased. This spontaneous afternoon peak has not previously been reported. The shaking signal consequently appears to convey the general message “reallocate labor to different activities” with receiver context specifying a more precise meaning. In the context of foraging, the shaking signal appears to activate (and perhaps deactivate) colony foraging preparations. The generally weak response elicited by modulatory signals such as the shaking signal may result from a high receiver response threshold which allows the receiver to integrate multiple sources of information and which thereby increases the probability that receiver actions will be appropriate to colony needs. Received: 21 March 1997 / Accepted after revision: 30 August 1997     

Worker allocation in insect societies: coordination of nectar foragers and nectar receivers in honey bee (Apis mellifera) colonies

Nectar collection in the honey-bee is partitioned. Foragers collect nectar and take it to the nest, where they transfer it to receiver bees who then store it in cells. Because nectar is a fluctuating and unpredictable resource, changes in worker allocation are required to balance the work capacities of foragers and receivers so that the resource is exploited efficiently. Honey bee colonies use a complex system of signals and other feedback mechanisms to coordinate the relative and total work capacities of the two groups of workers involved. We present a functional evaluation of each of the component mechanisms used by honey bees – waggle dance, tremble dance, stop signal, shaking signal and abandonment – and analyse how their interplay leads to group-level regulation. We contrast the actual regulatory system of the honey bee with theory. The tremble dance conforms to predicted best use of information, where the group in excess applies negative feedback to itself and positive feedback to the group in shortage, but this is not true of the waggle dance. Reasons for this and other discrepancies are discussed. We also suggest reasons why honey bees use a combination of recruitment plus abandonment and not switching between subtasks, which is another mechanism for balancing the work capacities of foragers and receivers. We propose that the waggle and tremble dances are the primary regulation mechanisms, and that the stop and shaking signals are secondary mechanisms, which fine-tune the system. Fine-tuning is needed because of the inherent unreliability of the cues, queueing delays, which foragers use to make recruitment decisions. Received: 15 December 1998 / Received in revised form: 6 March 1999 / Accepted: 12 March 1999     

Swarm cognition in honey bees

We synthesize findings from neuroscience, psychology, and behavioral biology to show that some key features of cognition in the neuron-based brains of vertebrates are also present in the insect-based swarm of honey bees. We present our ideas in the context of the cognitive task of nest-site selection by honey bee swarms. After reviewing the mechanisms of distributed evidence gathering and processing that are the basis of decision making in bee swarms, we point out numerous similarities in the functional organization of vertebrate brains and honey bee swarms. These include the existence of interconnected subunits, parallel processing of information, a spatially distributed memory, layered processing of information, lateral inhibition, and mechanisms of focusing attention on critical stimuli. We also review the performance of simulated swarms in standard psychological tests of decision making: tests of discrimination ability and assessments of distractor effects.     

Brood pheromone stimulates pollen foraging in honey bees (Apis mellifera)

Foraging and the mechanisms that regulate the quantity of food collected are important evolutionary and ecological attributes for all organisms. The decision to collect pollen by honey bee foragers depends on the number of larvae (brood), amount of stored pollen in the colony, as well as forager genotype and available resources in the environment. Here we describe how brood pheromone (whole hexane extracts of larvae) influenced honey bee pollen foraging and test the predictions of two foraging-regulation hypotheses: the indirect or brood-food mechanism and the direct mechanism of pollen-foraging regulation. Hexane extracts of larvae containing brood pheromone stimulated pollen foraging. Colonies were provided with extracts of 1000 larvae (brood pheromone), 1000 larvae (brood), or no brood or pheromone. Colonies with brood pheromone and brood had similar numbers of pollen foragers, while those colonies without brood or pheromone had significantly fewer pollen foragers. The number of pollen foragers increased more than 2.5-fold when colonies were provided with extracts of 2000 larvae as a supplement to the 1000 larvae they already had. Within 1 h of presenting colonies with brood pheromone, pollen foragers responded to the stimulus. The results from this study demonstrate some important aspects of pollen foraging in honey bee colonies: (1) pollen foragers appear to be directly affected by brood pheromone, (2) pollen foraging can be stimulated with brood pheromone in colonies provided with pollen but no larvae, and (3) pollen forager numbers increase with brood pheromone as a supplement to brood without increasing the number of larvae in the colony. These results support the direct-stimulus hypothesis for pollen foraging and do not support the indirect-inhibitor, brood-food hypothesis for pollen-foraging regulation. Received: 5 March 1998 / Accepted after revision: 29 August 1998     

Verification of radionuclide transfer factors to domestic-animal food products, using indigenous elements and with emphasis on iodine

Recent reviews have established benchmark values for transfer factors that describe radionuclide transfer from plants to animal food product such as milk, eggs and meat. They also illustrate the paucity of data for some elements and some food products. The present study quantified transfer data using indigenous elements measured in dairy, poultry and other livestock farms in Canada. Up to 62 elements are reported, with particular emphasis on iodine (I) because of the need to accurately assess the behaviour of ¹²⁹I from disposal of nuclear fuel waste. There was remarkable agreement with the literature values, and for many elements the present study involved many more observations than were previously available. Perhaps the most important observation was that product/substrate concentration ratios (CR) were quite consistent across species, whereas the traditional fractional transfer factors (TF, units of d kg⁻¹ or d L⁻¹) necessarily vary with body mass (feed intake). This suggests that for long-term assessments, it may be advisable to change the models to use CR rather than TF.     

The honey bee’s tremble dance stimulates additional bees to function as nectar receivers

If a forager bee returns to her hive laden with high-quality nectar but then experiences difficulty finding a receiver bee to unload her, she will begin to produce a conspicuous communication signal called the tremble dance. The context in which this signal is produced suggests that it serves to stimulate more bees to function as nectar receivers, but so far there is no direct evidence of this effect. We now report an experiment which shows that more bees do begin to function as nectar receivers when foragers produce tremble dances. When we stimulated the production of tremble dances in a colony and counted the number of bees engaged in nectar reception before and after the period of intense tremble dancing, we found a dramatic increase. In two trials, the number of nectar receivers rose from 17% of the colony’s population before tremble dancing to 30–50% of the population after the dancing. We also investigated which bees become the additional nectar receivers, by looking at the age composition of the receiver bees before and after the period of intense tremble dancing. We found that none of the bees recruited to the task of nectar reception were old bees, most were middle-aged bees, and some were even young bees. It remains unclear whether these auxiliary nectar receivers were previously inactive (as a reserve supply of labor) or were previously active on other tasks. Overall, this study demonstrates that a honey bee colony is able to rapidly and strongly alter its allocation of labor to adapt to environmental changes, and it further documents one of the communication mechanisms underlying this ability. Received: 31 May 1996/Accepted after revision: 9 August 1996     

Antibacterial activity of different natural honeys from Transylvania,Romania

Honey is used in food industry and medicine due to its nutritive, therapeutic and dietetic qualities. The microbiological characteristics of 10 unpasteurized honey samples of known origin, collected from Transylvania beekeepers (Romania) were determined. The antibacterial activity of these types of honey against Escherichia coli, Staphylococcus aureus, Staphylococcus epidermidis, Salmonella enteritidis, Salmonella anatum, Salmonella choleraesuis, Bacillus cereus, Bacillus subtilis subsp. spizizenii and Listeria monocytogenes strains was studied. The most sensitive to the antibacterial activity were the two staphylococus strains (the largest diameter of inhibition zone was 18 mm) and B. subtilis strains (13.5 mm). The strains of B. cereus, E. coli, L. monocytogenes and Salmonella spp. were found to present resistance to some of the honey samples. Manna, sunflower and polyfloral honeys presented high antibacterial activity while acacia and linden honeys had a lower activity in terms of the number of sensible strains. Statistical analysis shows that the type of strains and the type of honey have influence on the diameter of inhibition.     

Comparisons of forager distributions from matched honey bee colonies in suburban environments

We conducted experiments designed to examine the distribution of foraging honey bees (Apis mellifera) in suburban environments with rich floras and to compare spatial patterns of foraging sites used by colonies located in the same environment. The patterns we observed in resource visitation suggest a reduced role of the recruitment system as part of the overall colony foraging strategy in habitats with abundant, small patches of flowers. We simultaneously sampled recruitment dances of bees inside observation hives in two colonies over 4 days in Miami, Florida (1989) and from two other colonies over five days in Riverside, California (1991). Information encoded in the dance was used to determine the distance and direction that bees flew from the hive for pollen and nectar and to construct foraging maps for each colony. The foraging maps showed that bees from the two colonies in each location usually foraged at different sites, but occasionally they visited the same patches of flowers. Each colony shifted foraging effort among sites on different days. In both locations, the mean flight distances differed between colonies and among days within colonies. The flight distances observed in our study are generally shorter than those reported in a similar study conducted in a temperate deciduous forest where resources were less dense and floral patches were smaller.     

The communal crop: modulation of sucrose response thresholds of pre-foraging honey bees with incoming nectar quality

We examined whether the quality (concentration) of incoming sucrose solutions returned by foraging honey bees affected the response thresholds of pre-foraging members of the colony. Six pairs of colonies were given ad libitum access to sucrose solution feeders. A colony from each pair was switched from 20–50% sugar concentration feeders while the other continued to have access to 20% sucrose feeders. Proboscis extension response (PER) scores to an increasing series of sucrose concentrations were significantly higher in pre-foragers of colonies foraging on 20% sucrose throughout compared to pre-foragers in colonies where foraging was switched to 50% sucrose. Although all colonies had honey stores, the concentration of sugar solution in non-foraging bees crops were significantly lower in bees from colonies foraging on 20% sucrose compared to those from colonies foraging on 50% sucrose. Because response thresholds to sugar of young bees were modulated by the concentration of sucrose solution returned to colonies, we repeated the 2000 study of Pankiw and Page that potentially confounded baseline response thresholds with modulated scores due to experience in the colony. Here, we examined PER scores to sucrose in bees within 6 h of emergence, prior to feeding experience, and their forage choice 2 to 3 weeks later. Pollen foragers had higher PER scores as newly emerged bees compared to bees that eventually became nectar foragers. These results confirm those of the 2000 study by Pankiw and Page. Combined, these experiments demonstrate that variation in pre-forager sucrose response thresholds are established prior to emerging as adults but may be modulated by incoming resources later on. Whether this modulation has long-term effects on foraging behavior is unknown but modulation has short-term effects and the potential to act as a means of communication among all bees in the colony.Communicated by M. Giurfa