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 their hive mates information about the direction and distance to patches of flowers yielding nectar and pollen, to water sources, or to new housing locations. [1][2] Thus the waggle dance is a mechanism whereby successful foragers can recruit other bees in their colony to good locations for collecting various resources. It was once thought that bees had two distinct recruitment dances — round dances and waggle dances — the former for indicating nearby targets and the latter for indicating distant targets, but it is now known that a round dance is simply a waggle dance with a very short waggle run (see below). Austrian ethologist and Nobel laureate Karl von Frisch was one of the first who translated the meaning of the waggle dance. [3] On the right side is a video explaining the waggle dance in depth, as well as the experimentation that went into discovering the 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. [4]The direction and duration of waggle runs are closely correlated with the direction and distance of the patch of flowers being advertised by the dancing bee. Flowers 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. [1] The farther the target, the longer the waggle phase, with a rate of increase of about 75 milliseconds per 100 meters. Waggle dancing bees that have been in the hive 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. [5]When scientists placed a dead Apis mellifera bee on flowers they discovered that bees performed far fewer waggle dances upon returning to their nest. This is likely to be because they associate the dead bee with the presence of a predator on the flower and so it is better for other bees to not forage there. [6][7]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”. [3] He did, however, admit ignorance as to purpose of the dancing. 35 years before that, Ernst Spitzner observed bees dancing and interpreted it as transmitting forage resource odors to other nestmates. [3] 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".[8] 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
The dance language, as defined by von Frisch, is the information about direction, distance, and quality of a resource (such as food or nesting sites) contained within the waggle dance. Karl von Frisch named the dance language “Tanzsprache” in his native tongue. [9]Though von Frisch insisted on the direct connection between the dance language and the waggle dance, recent criticism holds that potential foragers need not correctly translate the dance language from the waggle dance to successfully forage. [9]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. [9]Bees that follow a waggle dance can successfully forage without decoding the dance language information in several ways [10]: - 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 the known resource.
- Using information communicated in the waggle dance is more useful to foragers when private information about resources is lacking.
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. [11] 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. [10]
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. [10]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 sparsely scattered and bloom for short periods of time. Thus, in tropical zones information about forage location might be more valuable than in temperate zones. [12] EvolutionAncestors 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. [10]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. [10]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. [13][14] 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. [15] Applications to operations researchIn line with recent work in swarm intelligence research involving optimization algorithms inspired by the behavior of social insects and animals such as fish, birds, and ants, recently there has been research on using bee waggle dance behavior for efficient fault-tolerant routing. [16] From the abstract of Wedde, Farooq, and Zhang (2004) [17]: 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.[18][19]
See also References- ^ a b Riley, J.; Greggers, U.; Smith, A.; Reynolds, D.; 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. edit
- ^ Seeley, T.D., P.K. Visscher, and K.M. Passino. (2006) Group decision making in honey bee swarms. American Scientist. 94:220-229.
- ^ a b c Frisch, Karl von. (1967) The Dance Language and Orientation of Bees. Cambridge, Mass.: The Belknap Press of Harvard University Press.
- ^ Thom, C.; Gilley, D.; Hooper, J.; Esch, H. (2007). "The scent of the waggle dance". PLoS biology 5 (9): e228. doi:10.1371/journal.pbio.0050228. PMC 1994260. PMID 17713987. edit
- ^ Bozic, J.; Abramson, C.; Bedencic, M. (2006). "Reduced ability of ethanol drinkers for social communication in honeybees (Apis mellifera carnica Poll.)". Alcohol (Fayetteville, N.Y.) 38 (3): 179–183. doi:10.1016/j.alcohol.2006.01.005. PMID 16905444. edit
- ^ Walker, Matt (31 July 2009). "Honeybees warn of Risky Flowers". BBC Earth News. Retrieved 18 January 2010
- ^ Abbott, K. R.; Dukas, R. (2009). "Honeybees consider flower danger in their waggle dance". Animal Behaviour 78 (3): 633. doi:10.1016/j.anbehav.2009.05.029. edit
- ^ Aristotle, Historia animalium, IX, 40, Becker 624b; modified from the translation by D.W. Thompson in The Works of Aristotle, Clarendon, Oxford, 1910.
- ^ a b c Grüter, C.; Balbuena, M.; Farina, W. (2008). "Informational conflicts created by the waggle dance". Proceedings. Biological sciences / the Royal Society 275 (1640): 1321–1327. doi:10.1098/rspb.2008.0186. PMC 2602683. PMID 18331980. edit
- ^ a b c d e Grüter, C.; Farina, W. (2009). "The honeybee waggle dance: can we follow the steps?". Trends in ecology & evolution (Personal edition) 24 (5): 242–247. doi:10.1016/j.tree.2008.12.007. PMID 19307042. edit
- ^ Saussure, Ferdinand de (1916), "Nature of the Linguistics Sign", in: Charles Bally & Albert Sechehaye (Ed.), Cours de linguistique générale, McGraw Hill Education.
- ^ Dornhaus, A.; Chittka, L. (2004). "Why do honey bees dance?". Behavioral Ecology and Sociobiology 55: 395. doi:10.1007/s00265-003-0726-9. edit
- ^ Gould; Towne (1989). "On the Evolution of the Dance Language: Response to Dyer and Seeley". American Naturalist (The American Society of Naturalists) 134 (1): 156–159. doi:10.1086/284972. edit
- ^ Dyer; Seeley (1991). "Dance Dialects and Foraging Range in Three Asian Honey Bee Species". Behavioral Ecology and Sociobiology (Springer) 28 (4): 227–233. doi:10.2307/4600541. JSTOR 4600541. edit
- ^ Su, S.; Cai, F.; Si, A.; Zhang, S.; Tautz, J.; Chen, S.; Giurfa, M. (2008). Giurfa, Martin. ed. "East learns from West: Asiatic honeybees can understand dance language of European honeybees". PloS one 3 (6): e2365. Bibcode 2008PLoSO...3.2365S. doi:10.1371/journal.pone.0002365. PMC 2391287. PMID 18523550. edit
- ^ Crina, Grosan; Abraham Ajith. (2006) Stigmergic Optimization: Inspiration, Technologies and Perspectives. Studies in Computational Intelligence. Vol. 31. pp. 1-24. Springer Berlin / Heidelberg. ISBN 978-3-540-34689-0
- ^ Wedde, H. F.; Farooq, M.; Pannenbaecker, T.; Vogel, B.; Mueller, C.; Meth, J.; Jeruschkat, R. (2005). BeeAdHoc. pp. 153. doi:10.1145/1068009.1068034. edit
- ^ Nakrani, S.; Tovey, C. (2004). "On Honey Bees and Dynamic Server Allocation in Internet Hosting Centers". Adaptive Behavior 12: 223. doi:10.1177/105971230401200308. edit
- ^ C. Tovey, (2004) "The Honey Bee Algorithm: A Biological Inspired Approach to Internet Server Optimization". Engineering Enterprise. Spring 2004, pp.13-15.
Further reading- Gould JL (1975). "Honey bee recruitment: the dance-language controversy". Science 189: 685−693.
- 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. 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.
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