Consistent personality differences in house-hunting behavior but not decision speed in swarms of honey bees (<Emphasis Type="BoldItalic">Apis mellifera</Emphasis>)

Speed-accuracy tradeoffs are a common feature of decision-making processes, both in individual animals and in groups of animals working together to reach a single collective decision. Individual organisms display consistent differences in their “impulsivity,” and vary in their tendency to make rapid, impulsive choices as opposed to slower, more accurate decisions. However, we do not yet know whether groups of animals consistently differ in their tendency to prioritize decision speed over accuracy. We challenged 17 swarms of honey bees (Apis mellifera) to simultaneously choose a new nest site in each of three locations, and measured their decision speeds in each trial. We found that swarms displayed consistent personality differences in the number of waggle dances and shaking signals they performed and in how actively they scouted for new nest sites. However, swarms did not consistently differ in how long they took to choose a nest site. We suggest that house-hunting A. mellifera swarms may place an especially high emphasis on decision accuracy when choosing a nest site, and that chance events—such as the time when each swarm discovers a sufficiently high-quality nest site—may consequently play a greater role in determining a swarm’s decision speed than intrinsic characteristics such as a swarm’s “impulsivity.”     

The control of water collection in honey bee colonies

A honey bee (Apis mellifera) colony adaptively controls the collection of water by its foragers, increasing it when high temperatures necesssitate evaporative cooling inside the hive and decreasing it when the danger of overheating passes. This study analyzes the mechanisms controlling water collection once it has begun, that is, how a colony's water collectors know whether to continue or stop their activity. M. Lindauer suggested that water collectors acquire information about their colony's need for more water by noting how easily they can unload their water to bees inside the hive. In support of this hypothesis, we found that a water collector's ease of unloading does indeed change when her colony's need for water changes. How does a water collector sense the ease of unloading? Multiple variables of the unloading experience change in relation to a colony's water need. Three time-based variables – initial search time, total search time, and delivery time – all change quite strongly. But what changes most strongly is the number of unloading rejections (refusals by receiver bees to take the water), suggesting that this is the primary index of ease of unloading. Why does a water collector's ease of unloading change when her colony's need for water changes? Evidently, what links these two variables is change in the number of water receivers. These are middle-aged bees that receive water just inside the hive entrance, then transport it deeper inside the hive, and finally smear it on the walls of cells or give it to other bees, or both. A colony increases the number of water receivers when its water need increases by having bees engaged in nectar reception and other tasks (and possibly also bees that are not working) switch to the task of water reception. Evidently the activation of additional water receivers does not strongly reduce the number of nectar receivers in a colony, since a colony can increase greatly its water collection without simultaneously decreasing its collection of rich nectar. This study provides a clear example of the way that the members of a social insect colony can use indirect indicators of their colony's labor needs to adaptively control the work that they perform.     

Coordinating a group departure: who produces the piping signals on honeybee swarms?

A swarm of honeybees provides a striking example of an animal group performing a synchronized departure for a new location; in this case, thousands of bees taking off at once to fly to a new home. However, the means by which this is achieved remain unclear. Shortly before takeoff, one hears a crescendo of a high-pitched mechanical signal—worker piping—so we explored the role of this signal in coordinating a swarm’s mass takeoff. Specifically, we examined whether exclusively nest site scouts produce the worker piping signal or whether it is produced in a relay or chain reaction fashion. We found no evidence that bees other than the scouts that have visited the swarm’s chosen nest site produce piping signals. This absence of relay communication in piping suggests that it is a signal that only primes swarms for takeoff and that the release of takeoff is triggered by some other signal or cue; perhaps the takeoff of bees on the swarm periphery as they reach flight temperature in response to piping.     

An equilibrium theory of queen production in honeybee colonies preparing to swarm

Summary An equilibrium model is developed which seeks to explain the regulation of queen rearing in honeybee colonies preparing to swarm. The model postulates that there is a balance between nurse bees becoming inhibited from queen rearing and nurses losing their inhibition, and that whether a colony does or does not rear queens reflects the equilibrium percentage of inhibited nurses. This model leads to a quantitative prediction about the size of a conoly's nurse population at which queen rearing should start. Comparing the model's predictions with empirical observations pinpoints data needed for a more complete explanation of control of queen rearing. In particular, the model suggests a central regulatory role for density-dependent changes in the behaviors involved in queen substance dispersal.     

Measurement of nest cavity volume by the honey bee (Apis mellifera)

Behavioral Ecology and Sociobiology -     

Fever in honeybee colonies

 Honeybees, Apis spp., maintain elevated temperatures inside their nests to accelerate brood development and to facilitate defense against predators. We present an additional defensive function of elevating nest temperature: honeybees generate a brood-comb fever in response to colonial infection by the heat-sensitive pathogen Ascosphaera apis. This response occurs before larvae are killed, suggesting that either honeybee workers detect the infection before symptoms are visible, or that larvae communicate the ingestion of the pathogen. This response is a striking example of convergent evolution between this "superorganism" and other fever-producing animals. Received: 2 September 1999 / Accepted in revised form: 28 February 2000     

Imprecision in waggle dances of the honeybee (Apis mellifera) for nearby food sources: error or adaptation?

A curious feature of the honeybee's waggle dance is the imprecision in the direction indication for nearby food sources. One hypothesis for the function of this imprecision is that it serves to spread recruits over a certain area and thus is an adaptation to the typical spatial configuration of the bees' food sources, i.e., flowers in sizable patches. We report an experiment that tests this tuned-error hypothesis. We measured the precision of direction indication in waggle dances advertising a nest site (typically a tree cavity, hence a target that is almost a point) and compared it with that of dances advertising a food source (typically a flower patch, hence a target that covers an area). The precision of dances for a nearby nest site was significantly higher than that of dances for an equidistant feeder. This was demonstrated four times with four colonies. Our evidence therefore supports the hypothesis that the level of precision in the direction indication for nearby food sources is tuned to its optimum without being at its maximum. Received: 9 December 1998 / Received in revised form: 24 February 1999 / Accepted 12 March 1999     

An oligarchy of nest-site scouts triggers a honeybee swarm’s departure from the hive

Animals that travel in groups must synchronize the timing of their departures to assure cohesion of the group. While most activities in large colonies of social insects have decentralized control, certain activities (e.g., colony migration) can have centralized control, with only a special subset of well-informed individuals making a decision that affects the entire colony. We recently discovered that a small minority of individuals in a honeybee colony—an oligarchy—decides when to trigger the departure of a swarm from its hive. The departure process begins with some bees producing the worker-piping signal (the primer for departure) and is followed by these bees producing the buzz-run signal (the releaser for departure). In this study, we determined the identity of these signalers. We found that a swarm’s nest-site scouts search for potential nest cavities prior to the departure of the swarm from its hive. Furthermore, we found that the predeparture nest-site scouts are the sole producers of the worker-piping signal and that they are the first producers of the buzz-run signal. The control of the departure of a honeybee swarm from its hive shows how a small minority of well-informed individuals in a large social insect colony can make important decisions about when a colony should take action.     

Assessing the benefits of cooperation in honeybee foraging: search costs,forage quality,and competitive ability

Summary The foragers in honeybee colonies cooperate by sharing information about rich sources of food. This study examines three hypotheses about the benefits of this cooperation: (H1) it decreases foragers' costs in finding new food sources, (H2) it increases the quality of the food sources located by foragers, and (H3) it increases the ability of a colony's foragers to compete for high-quality food sources. To test each hypothesis, we identified a critical pattern in the foraging process which, if observed, would cast doubt on that hypothesis, and then gathered data to check for these patterns. Our observations do not support the first hypothesis, but do support the second and third. These results, in addition to helping us understand the functional significance of the honeybee's dance language, provide insights into the colonial organization of foraging by honeybees.     

The functional organization of resin work in honeybee colonies

Resin is an important building material in the nests of honeybees, but little is known about how it is handled within the nest and how its collection is controlled. We studied the functional organization of resin work to better understand how a colony adaptively controls its intake of resin. Two hypotheses have been proposed for how resin collectors stay informed of the need for additional resin: (1) the unloading difficulty hypothesis (resin need is sensed indirectly by the unloading delay) and (2) the caulking activity hypothesis (resin need is sensed directly while engaged in using resin). A falsifiable prediction of the latter hypothesis, but not of the former, is that resin collectors not only gather resin outside the hive but also regularly handle resin inside the hive (taking it from other bees and using it to caulk crevices). Consistent with this prediction are our findings that in the resin sector of a colony’s economy, unlike in the pollen, nectar, and water sectors, there is no strict division of labor between the collectors and the users of a material. Over the course of a day, bees seen collecting resin were also commonly seen using resin. Moreover, we found that the unloading locations of resin collectors are unlike those of water and nectar collectors, being deep inside the hive (at the sites of resin use) rather than at the hive entrance. This arrangement facilitates the engagement in resin use by resin collectors. We conclude that the caulking activity hypothesis is well-supported, but that the unloading difficulty hypothesis also remains viable, for we found that resin collectors experience variable delays in getting rid of their loads, from less than 15 min to more than an hour, consistent with this hypothesis. The stage is now set for experimental tests of these two hypotheses. Both may be correct, which if true will imply that social insect workers, despite their small brains, can acquire and integrate information from multiple sources to improve their knowledge of conditions within the colony.