waggle

　

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1. …‘を'振る,揺り動かす(wag) / 揺れ動く / 揺れ,振ること

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出典(authority):フリー百科事典『ウィキペディア（Wikipedia）』「2012/12/12 10:58:51」(JST)

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This article is about bee behavior. For the beer, see Wells & Young's Brewery. For the football play, see bootleg play.

The waggle dance - the direction the bee moves in relation to the hive indicates direction; if it moves vertically upwards the direction to the source is directly towards the Sun, the duration of the waggle part of the dance signifies the distance.

Video demonstrating the waggle dance of the honeybee

Waggle dance is a term used in beekeeping and ethology for a particular figure-eight dance of the honey bee. By performing this dance, successful foragers can share with other members of the colony, information about the direction and distance to patches of flowers yielding nectar and pollen, to water sources, or to new housing locations. A waggle dance with a very short waggle run used to be characterized as a distinct (round) recruitment dance (see below). Austrian ethologist and Nobel laureate Karl von Frisch was one of the first who translated the meaning of the waggle dance.

Description

Figure-Eight-Shaped Waggle Dance of the Honeybee (Apis mellifera). A waggle run oriented 45° to the right of ‘up’ on the vertical comb (A) indicates a food source 45° to the right of the direction of the sun outside the hive (B). The abdomen of the dancer appears blurred because of the rapid motion from side to side.

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. For cavity-nesting honey bees, 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.

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 Apis mellifera bee 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”. He did, however, admit ignorance as to the purpose of the dancing. 35 years prior to Unhoch's observations, Ernst Spitzner observed bees dancing and interpreted it as transmitting forage resource odors to other nestmates. Even Aristotle, in addition to describing flower constancy behavior, suspected that some form of communication occurred between foragers within a nest:

"On each trip the bee does not fly from a flower of one kind to a flower of another, but flies from one violet, say, to another violet, and never meddles with another flower until it has got back to the hive; on reaching the hive they throw off their load, and each bee on her return is followed by three or four companions. What it is that they gather is hard to see, and how they do it has not been observed".

Jürgen Tautz also writes about it in his book "The Buzz about Bees":

Page 112: 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. Tautz,J.: The Buzz about Bees - Biology of a Superorganism (photos by H. R. Heilmann) Springer Heidelberg & Berlin, 2008

Controversy

Workers of Apis mellifera carnica on honeycomb.

The Dance Language vs. The Waggle Dance

As defined by von Frisch, Tanzsprache (German for "dance language") is the information about direction, distance, and quality of a resource (such as food or nesting sites) contained within the waggle dance. Though von Frisch insisted on the direct connection between the Tanzsprache and the performance of the waggle dance, recent criticism holds that potential foragers need not correctly translate the dance language from the waggle dance to successfully forage. In an experiment on the honeybee Apis mellifera, most individuals who thoroughly followed a waggle dance ignored the resource direction and location information. Instead, 93% of the foragers returned to foraging areas they had previous knowledge of.

Bees that follow a waggle dance can successfully forage without decoding the dance language information in several ways:

• Dance follower may use olfactory information from the dancer and find either the same resource or a different one with a similar scent.
• Following a dance may simply trigger foraging behavior. A forager may then search randomly for resources.
• Following a dance may reactivate private knowledge of a resource. After reactivation, the forager may return to the known resource.
• Using information communicated in the waggle dance is more useful to foragers when private information about resources is lacking.

Is Dance Language a Language?

The use of the word “language” may lead to misrepresentations of the waggle dance. The Swiss linguist Ferdinand de Saussure proposed a system of language a sign is made up two chief components. The signifier is the physical or phonetic representation of a sign. The signified is the conceptual component. If the dance language followed the Saussurian dyadic model of semiotics, the signifier would be the waggle dance and the signified would be the location of the foraging resource. Though the dance language may or may not follow this sort of pattern, it is not considered to be a language with syntactical grammar or a set of symbols.

Efficiency and Adaptation

The waggle dance may be less efficient than once thought. Some bees observe over 50 waggle runs without successfully foraging, while others will forage successfully after observing 5 runs.

The waggle dance may be adaptive in some environments and not in others. In temperate habitats, honey bee colonies routinely perform the waggle dance, but can still successfully forage when the dance is experimentally obscured. In tropical habitats, honey bee foraging is severely impaired if waggle dancing is prevented. This is thought to be due to the patchiness of resources in tropical environment versus the homogeneity of resources in temperate environments. In the tropics, food resources can come in the form of flowering trees which are rich in nectar but are scattered sparsely and bloom only briefly. Thus, in tropical zones information about forage location might be more valuable than in temperate zones.

Evolution

Ancestors to modern honeybees most likely performed excitatory movements to encourage other nestmates to forage. These excitatory movements include shaking, zig-zagging, buzzing and crashing into nestmates. Similar behavior is observed in other Hymenoptera including stingless bees, wasps, bumblebees and ants.

The waggle dance is thought to have evolved to aid in communicating information about a new nest site, rather than spatial information about foraging sites.

Observations have suggested that different species of honeybees have different "dialects" of the waggle dance, each species or subspecies dance varying by curve or duration. A recent study demonstrated that a mixed colony of Asiatic honeybees (Apis cerana cerana) and European honeybees (Apis mellifera ligustica) were gradually able to understand one another's 'dialects' of waggle dance.

Applications to operations research

In line with recent work in swarm intelligence research involving optimization algorithms inspired by the behavior of social insects (including bees, ants and termites), and vertebrates such as fish and birds, there has recently been research on using bee waggle dance behavior for efficient fault-tolerant routing. From the abstract of Wedde, Farooq, and Zhang (2004):

In this paper we present a novel routing algorithm, BeeHive, which has been inspired by the communicative and evaluative methods and procedures of honey bees. In this algorithm, bee agents travel through network regions called foraging zones. On their way their information on the network state is delivered for updating the local routing tables. BeeHive is fault tolerant, scalable, and relies completely on local, or regional, information, respectively. We demonstrate through extensive simulations that BeeHive achieves a similar or better performance compared to state-of-the-art algorithms.

Another bee-inspired stigmergic computational technique called bee colony optimization is employed in Internet Server Optimization.

The Zigbee RF protocol is named after the waggle dance.

See also

• Animal communication
• Bee learning and communication
• Tremble dance
• Grooming dance

References

Further reading

• Gould JL (1975). "Honey bee recruitment: the dance-language controversy". Science 189 (4204): 685−693. Bibcode 1975Sci...189..685G. doi:10.1126/science.1154023.
• Riley JR, Greggers U, Smith AD, Reynolds DR, Menzel R (2005). "The flight paths of honeybees recruited by the waggle dance". Nature 435 (7039): 205–207. Bibcode 2005Natur.435..205R. doi:10.1038/nature03526. PMID 15889092.
• Seeley TD (1995). The Wisdom of the Hive. Cambridge, MA: Harvard University Press.
• von Frisch K (1967). The Dance Language and Orientation of Bees. Cambridge, MA: Harvard University Press.

External links

• Waggle Dance Infographic - VetSci
• Communication and Recruitment to Food Sources by Apis mellifera — USDA-ARS (accessed 2005-03)
• Honeybee Communication — Kimball's Biology Pages (accessed 2005-09)

English Journal

• Exposed science: Genes, the environment, and the politics of population health.

• Waggle J.
• Global public health.Glob Public Health.2014 Oct;9(9):1119-21. doi: 10.1080/17441692.2014.955043. Epub 2014 Sep 10.
• PMID 25204994

• Decision-making in honeybee swarms based on quality and distance information of candidate nest sites.

• Laomettachit T1, Termsaithong T2, Sae-Tang A3, Duangphakdee O4.
• Journal of theoretical biology.J Theor Biol.2014 Sep 10;364C:21-30. doi: 10.1016/j.jtbi.2014.09.005. [Epub ahead of print]
• In the nest-site selection process of honeybee swarms, an individual bee performs a waggle dance to communicate information about direction, quality, and distance of a discovered site to other bees at the swarm. Initially, different groups of bees dance to represent different potential sites, but ev
• PMID 25218431

• Honey loading for pollen collection: regulation of crop content in honeybee pollen foragers on leaving hive.

• Harano K1, Mitsuhata-Asai A, Sasaki M.
• Die Naturwissenschaften.Naturwissenschaften.2014 Jul;101(7):595-8. doi: 10.1007/s00114-014-1185-z. Epub 2014 Jun 13.
• Before foraging honeybees leave the hive, each bee loads its crop with some amount of honey "fuel" depending on the distance to the food source and foraging experience. For pollen collection, there is evidence that foragers carry additional honey as "glue" to build pollen loads. This study examines
• PMID 24925356

Japanese Journal

• 時空間断面画像を用いたミツバチダンス行動個体の自動検出 (MEとバイオサイバネティックス)

• 藤井 昌樹,礒川 悌次郎,池野 英利 [他]
• 電子情報通信学会技術研究報告 : 信学技報 111(482), 207-212, 2012-03-14
• NAID 40019226111

• 蜜蜂の採餌行動を模した解の選別基準を自律的に調整するBCO

• 古川 まき,鈴木 泰博
• 情報処理学会研究報告. MPS, 数理モデル化と問題解決研究報告 2012-MPS-87(2), 1-6, 2012-02-23
• Bee Colony Optimization(BCO) は蜜蜂の採餌行動を模した最適化手法であり,巡回セールスマン問題 (Travelling Salesman Problem TSP) に適用されている.蜜蜂を模したエージェントが各 1 個の解を持ち,巣内での 8 の字ダンスによる仲間への解情報の伝搬と,巣外の採餌による解の局所的最適化によって最適解を探索する.この BCO に対し,本研究では …
• NAID 110008791132

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