The scent of the waggle dance

The waggle dance of honey bee (Apis mellifera L.) foragers communicates to nest mates the location of a profitable food source. We used solid-phase microextraction and gas chromatography coupled with mass spectrometry to show that waggle-dancing bees produce and release two alkanes, tricosane and pentacosane, and two alkenes, Z-(9)-tricosene and Z-(9)-pentacosene, onto their abdomens and into the air. Nondancing foragers returning from the same food source produce these substances in only minute quantities. Injection of the scent significantly affects worker behavior by increasing the number of bees that exit the hive. The results of this study suggest that these compounds are semiochemicals involved in worker recruitment. By showing that honey bee waggle dancers produce and release behaviorally active chemicals, this study reveals a new dimension in the organization of honey bee foraging.     

Informational conflicts created by the waggle dance

The honeybee (Apis mellifera) waggle dance is one of the most intriguing animal communication signals. A dancing bee communicates the location of a profitable food source and its odour. Followers may often experience situations in which dancers indicate an unfamiliar location but carry the scent of a flower species the followers experienced previously at different locations. Food scents often reactivate bees to resume food collection at previously visited food patches. This double function of the dance creates a conflict between the social vector information and the private navigational information. We investigated which kind of information followers with field experience use in this situation and found that followers usually ignored the spatial information encoded by the waggle dance even if they followed a dance thoroughly (five waggle runs or more). They relied on private information about food source locations instead (in 93% of all cases). Furthermore, foragers preferred to follow dancers carrying food odours they knew from previous field trips, independently of the spatial information encoded in the dance. Surprisingly, neither odour identity nor the location indicated by the dancer was an important factor for the reactivation success of a dance. Our results contrast with the assumption that (i) followers usually try to decode the vector information and (ii) dances indicating an unfamiliar location are of little interest to experienced foragers.     

Working against gravity: horizontal honeybee waggle runs have greater angular scatter than vertical waggle runs

The presence of noise in a communication system may be adaptive or may reflect unavoidable constraints. One communication system where these alternatives are debated is the honeybee (Apis mellifera) waggle dance. Successful foragers communicate resource locations to nest-mates by a dance comprising repeated units (waggle runs), which repetitively transmit the same distance and direction vector from the nest. Intra-dance waggle run variation occurs and has been hypothesized as a colony-level adaptation to direct recruits over an area rather than a single location. Alternatively, variation may simply be due to constraints on bees' abilities to orient waggle runs. Here, we ask whether the angle at which the bee dances on vertical comb influences waggle run variation. In particular, we determine whether horizontal dances, where gravity is not aligned with the waggle run orientation, are more variable in their directional component. We analysed 198 dances from foragers visiting natural resources and found support for our prediction. More horizontal dances have greater angular variation than dances performed close to vertical. However, there is no effect of waggle run angle on variation in the duration of waggle runs, which communicates distance. Our results weaken the hypothesis that variation is adaptive and provide novel support for the constraint hypothesis.     

Importance of wing movements for information transfer during honey bee waggle dance

There is growing evidence indicating that dancing honeybees can transfer some information about the found food source by means of wing movements. However, the available data are limited and inconclusive in the case of the frequency of wing beats. Therefore, in this study, the hypothesis that the wing beats convey information about the foraging distance was re‐examined. Honeybee dances were recorded using a high‐speed camera, and foraging distances were decoded from the duration of waggle phases. Dancing honeybees moved their wings for almost half (47%) of the duration of waggle phases. The number of wing‐beating pulses and the combined duration of wing beating were strongly positively correlated with the duration of waggle phases (p < .0014), whereas the mean frequency of wing beats, the mean duration of wing‐beating pulses and the mean number of wing beats in one wing‐beating pulse were not correlated with the duration of waggle phases (p > .05). Nevertheless, both the mean frequency of wing beats and the mean number of wing beats in one wing‐beating pulse were positively correlated with the mean frequency of abdomen waggles (p < .0014). They were also positively correlated with the mean frequency of wing‐beating pulses (p < .0014). The correlation matrix revealed that there were two groups of dance components that were positively correlated within groups, but negatively correlated between groups. One of the groups provided information about distance to the food source. We hypothesise that the other group, including the number of wing beats in one pulse, the frequency of wing beats, the frequency of wing‐beating pulses and the frequency of abdomen waggles, may provide information about the motivational state of foragers.     

Dynamic range compression in the honey bee auditory system toward waggle dance sounds

Honey bee foragers use a "waggle dance" to inform nestmates about direction and distance to locations of attractive food. The sound and air flows generated by dancer's wing and abdominal vibrations have been implicated as important cues, but the decoding mechanisms for these dance messages are poorly understood. To understand the neural mechanisms of honey bee dance communication, we analyzed the anatomy of antenna and Johnston's organ (JO) in the pedicel of the antenna, as well as the mechanical and neural response characteristics of antenna and JO to acoustic stimuli, respectively. The honey bee JO consists of about 300-320 scolopidia connected with about 48 cuticular "knobs" around the circumference of the pedicel. Each scolopidium contains bipolar sensory neurons with both type I and II cilia. The mechanical sensitivities of the antennal flagellum are specifically high in response to low but not high intensity stimuli of 265-350 Hz frequencies. The structural characteristics of antenna but not JO neurons seem to be responsible for the non-linear responses of the flagellum in contrast to mosquito and fruit fly. The honey bee flagellum is a sensitive movement detector responding to 20 nm tip displacement, which is comparable to female mosquito. Furthermore, the JO neurons have the ability to preserve both frequency and temporal information of acoustic stimuli including the "waggle dance" sound. Intriguingly, the response of JO neurons was found to be age-dependent, demonstrating that the dance communication is only possible between aged foragers. These results suggest that the matured honey bee antennae and JO neurons are best tuned to detect 250-300 Hz sound generated during "waggle dance" from the distance in a dark hive, and that sufficient responses of the JO neurons are obtained by reducing the mechanical sensitivity of the flagellum in a near-field of dancer. This nonlinear effect brings about dynamic range compression in the honey bee auditory system.     

Do honey bees encode distance information into the wing vibrations of the waggle dance?

An optical technique detected the wing vibration frequency of worker honey bees in an observation hive during the straight run of the waggle dance. Wing oscillation frequencies were recorded from dancing bees after they had visited a feeding station located from 50 to 1600 m from the hive. The bees vibrated their wings more rapidly after they visited nearby stations than when they foraged at more distant feeding stations. For example, the mean frequency of 315 Hz at 50 m dropped to only 207 Hz at 1600 m. Wing vibration frequency appears to be another factor to be added to the elements in the dance known to indicate the distance bees must fly to food sources. These known elements include the duration of the straight run and the number of wagtail movements in the run.     

Honey bee dance communication: waggle run direction coded in antennal contacts?

The behaviour of 38 honeybee dance followers and the patterns of antennal contact between followers and dancer were monitored during ten waggle runs for a feeding site 1200 m from the hive. The analysis was restricted to waggle runs with a maximum of 5 followers, allowing the followers to choose between different positions around the dancer. At the beginning of the waggle run, followers are rather evenly spaced around the dancer. During the waggle run, the followers tend to accumulate at the rear end of the dancer. At the end of the waggle run, all followers are found in a ±60° arc behind the dancer. The body orientation angles of the followers depend on their position relative to the dancer. The follower bees have intense antennal contact with the dancer. At least one temporal parameter of the contact pattern may inform the followers about their position relative to the dancer, may guide the dance followers to the rear end of the dancer and may allow them to extract information about the location of the food source advertised by the dance. The role of antennal contact for dance communication appears to have been underestimated in previous studies. Accepted: 20 February 1999     

Using the waggle dance to determine the spatial ecology of honey bees during commercial crop pollination

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Honey bee waggle dance communication increases diversity of pollen diets in intensively managed agricultural landscapes

The benefits of honey bee dance communication for colony performance in different resource environments are still not well understood. Here, we test the hypothesis that directional dance communication enables honey bee colonies to maintain a diverse pollen diet, especially in landscapes with low resource diversity. To test this hypothesis, we placed 24 Apis mellifera L. colonies with either intact or experimentally disrupted dance communication in eight agricultural landscapes that differed in the diversity of flowering plants and in the dominance of mass‐flowering crops. Pollen from incoming foragers was collected and identified via DNA metabarcoding. Disrupting dance communication affected the way the diversity of honey bee pollen diets was impacted by the dominance of mass‐flowering crops in available flower resources (p = .04). With increasing dominance of mass‐flowering crops in resource environments, foragers of colonies with intact communication foraged on an increasing proportion of available plant genera (p = .01). This was not the case for colonies with disrupted dance communication (p = .5). We conclude that the honey bee dance communication benefits pollen foraging on diverse plant resources and thereby contributes to high quality nutrition in environments with low‐resource diversity.     

The waggle dance of the honey bee: Which bees following a dancer successfully acquire the information?

During the waggle dance of the honeybee, the dancer is able to tell her nestmates the distance and direction to a rich food source (Frisch, 1967). Little is known about how waggle dance followers are able to read the waggle dance in the darkness of a hive. Initial observations showed that not all of the bees that appear to be dance followers behave the same. Some bees maneuver themselves behind the dancer, while others do not. The paths of a single dancer, trained to an artificial food source, and her followers were traced during the waggle runs. The success of these dance followers was compared to their position relative to the dancer. The results of this study show that during a waggle run a dance follower must position itself within a 30° arc behind the dancer in order to obtain the dance information. The results suggest that bees are using the position of their own bodies to determine direction.     

The horizontal magnetic dance of the honeybee is compatible with a single-domain ferromagnetic magnetoreceptor

Although honeybees are able to sense the geomagnetic field, very little is known about the method in which they are able to detect it. The recent discovery of biochemically precipitated magnetite (Fe₃O₄) in bees, however, suggests the possibility that they might use a simple compass organelle for magnetoreception. If so, their orientation accuracy ought to be related to the accuracy of the compass, e.g. it should be poor in weak background fields and enhanced in strong fields. When dancing to the magnetic directions on a horizontal honeycomb, bees clearly show this type of alignment behavior. A least-squares fit between the expected alignment of a compass and this horizontal dance data is consistent with this hypothesis, and implies that the receptors have magnetic moments of 5 × 10^?13 emu, or magnetite volumes near 10^?15 cm³. Additional considerations suggests that these crystals are slightly sub-spherical and single-domain in size, held symmetrically in their receptors, and have a magnetic orientation energy of approximately to 6 kT in the geomagneticfield. A model of a magnetite-based magnetoreceptor consistent with these constraints is discussed.     

Quantitative multi‐locus metabarcoding and waggle dance interpretation reveal honey bee spring foraging patterns in Midwest agroecosystems

We explored the pollen foraging behaviour of honey bee colonies situated in the corn and soybean dominated agroecosystems of central Ohio over a month‐long period using both pollen metabarcoding and waggle dance inference of spatial foraging patterns. For molecular pollen analysis, we developed simple and cost‐effective laboratory and bioinformatics methods. Targeting four plant barcode loci (ITS2, rbcL, trnL and trnH), we implemented metabarcoding library preparation and dual‐indexing protocols designed to minimize amplification biases and index mistagging events. We constructed comprehensive, curated reference databases for hierarchical taxonomic classification of metabarcoding data and used these databases to train the metaxa 2 DNA sequence classifier. Comparisons between morphological and molecular palynology provide strong support for the quantitative potential of multi‐locus metabarcoding. Results revealed consistent foraging habits between locations and show clear trends in the phenological progression of honey bee spring foraging in these agricultural areas. Our data suggest that three key taxa, woody Rosaceae such as pome fruits and hawthorns, Salix, and Trifolium provided the majority of pollen nutrition during the study. Spatially, these foraging patterns were associated with a significant preference for forests and tree lines relative to herbaceous land cover and nonflowering crop fields.