A waggle dance consists of one to 100 or more circuits, each of which consists of two phases: the waggle phase and the return phase. A worker bee's waggle dance involves running through a small figure-eight pattern: a waggle run (aka waggle phase) followed by a turn to the right to circle back to the starting point (aka return phase), another waggle run, followed by a turn and circle to the left, and so on in a regular alternation between right and left turns after waggle runs. 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. The direction and duration of waggle runs are closely correlated with the direction and distance of the resource being advertised by the dancing bee. In an experiment with capture and relocation of bees exposed to a waggle dance the bees followed the path that would have taken them to an experimental feeder had they not been displaced. For cavity-nesting honey bees, like the
western honey bee (
Apis mellifera) or
Apis nigrocincta, flowers that are located directly in line with the sun are represented by waggle runs in an upward direction on the vertical combs, and any angle to the right or left of the sun is coded by a corresponding angle to the right or left of the upward direction. The distance between hive and recruitment target is encoded in the duration of the waggle runs. The farther the target, the longer the waggle phase. The more excited the bee is about the location, the more rapidly it will waggle, so it will grab the attention of the observing bees, and try to convince them. If multiple bees are doing the waggle dance, it's a competition to convince the observing bees to follow their lead, and competing bees may even disrupt other bees' dances or fight each other off. In addition, some open-air nesting honeybees such as the
black dwarf honeybee (
Apis andreniformis), whose nests hang from twigs or branches, will perform a horizontal dance on a stage above their nest in order to signal to resources. Waggle dancing bees that have been in the nest for an extended time adjust the angles of their dances to accommodate the changing direction of the sun. Therefore, bees that follow the waggle run of the dance are still correctly led to the food source even though its angle relative to the sun has changed. The consumption of
ethanol by foraging bees has been shown to reduce waggle dance activity and increase occurrence of the
tremble dance. Kevin Abbott and Reuven Dukas of McMaster University in Hamilton, Ontario, Canada discovered that if a dead western honeybee is placed on a flower, bees performed far fewer waggle dances upon returning to the hive. The scientists explain that the bees associate the dead bee with the presence of a predator at the food source. The reduction of the dance repetition frequency, therefore, indicates that the dancing bees perform and communicate a form of risk/benefit analysis. Though first decoded by
Karl von Frisch, dancing behavior in bees had been observed and described multiple times prior. Around 100 years before Frisch's discovery, Nicholas Unhoch described dancing behavior of bees as being an indulgence "in certain pleasures and jollity". Jürgen Tautz also writes about it in his book
The Buzz about Bees (2008): Many elements of the communication used to recruit miniswarms to feeding sites are also observed in "true" swarming behavior. Miniswarms of foragers are not placed under the same selection pressure as are true swarms, because the fate of the entire colony is not at stake. A truly swarming colony has to be quickly led to a new home, or it will perish. The behavior used to recruit to food sources possibly developed from the "true" swarming behavior. ==Mechanism==