Editing Bee learning and communication (section)

==Communication==
{{see also|Bumblebee communication}}
Foragers communicate their floral findings in order to recruit other worker bees of the [[beehive|hive]] to forage in the same area. The factors that determine recruiting success are not completely known but probably include evaluations of the quality of [[nectar]] and/or [[pollen]] brought in.

There are two main hypotheses to explain how foragers recruit other workers—the [[waggle dance|"waggle dance" or "dance language"]] theory and the "odor plume" theory. The dance theory is far more widely accepted, and has far more empirical support than the odor theory. Supporters of the dance theory often grant odor a significant role in recruitment, while supporters of the odor theory have claimed that the dance is essentially irrelevant to recruitment. The academic debate between these theories has been polarized and sometimes hostile.<ref>{{Cite journal
| last1 = Munz | first1 = T.
| title = The Bee Battles: Karl von Frisch, Adrian Wenner and the Honey Bee Dance Language Controversy
| doi = 10.1007/s10739-005-0552-1
| journal = Journal of the History of Biology
| volume = 38
| issue = 3
| pages = 535–570
|date=November 2005
| s2cid = 120244044
}} {{closed access}}</ref>

=== Dance communication ===
[[File:Bee waggle dance.png|thumb|Figure-Eight-Shaped waggle dance of the honeybee ([[Apis mellifera]]). A waggle run oriented 45° to the right of ‘up' on the vertical comb indicates a food source 45° to the right of the direction of the sun outside the hive. The abdomen of the dancer appears blurred because of the rapid motion from side to side.]]

It has long been known that successfully foraging [[Western honey bee]]s perform a ''[[waggle dance]]'' upon their return to the [[beehive|hive]]. The laden forager dances on the comb in a circular pattern, occasionally crossing the circle in a zig-zag or waggle pattern. [[Aristotle]] described this behaviour in his ''Historia Animalium''.<ref>{{cite web|url=http://etext.lib.virginia.edu/toc/modeng/public/AriHian.html|title=HISTORIA ANIMALIUM|work=virginia.edu|date=6 September 2008 }}</ref> This waggle pattern of movement was thought to attract the attention of other bees. In 1947,<ref>[http://beesource.com/point-of-view/adrian-wenner/the-honey-bee-dance-language-controversy/ The Honey Bee Dance Language Controversy]</ref> [[Karl von Frisch]] correlated the runs and turns of the dance to the distance and direction of the food source from the hive. He reported that the orientation of the dance is correlated with the relative position of the sun to the food source, and the length of the waggle portion of the run is correlated to the distance of the food from the hive. Von Frisch also reported that the more vigorous the display is, the better the food.  Von Frish published these and many other observations in his 1967 book ''The Dance Language and Orientation of Bees''<ref>von Frisch, K. (1967) ''The Dance Language and Orientation of Bees''. Cambridge, MA: Harvard Univ. Press.</ref> and in 1973 he was awarded the [[Nobel Prize in Physiology or Medicine]] for his discoveries.

Later work has supported Von Frisch's  observations and added many details. It appears that all of the known species and races of [[honey bee]]s exhibit the behavior, but details of its execution vary among the different species. For example, in ''[[Apis florea]]'' and ''[[Apis andreniformis]]'' (the "dwarf honeybees") the dance is performed on the dorsal, horizontal portion of the nest, which is exposed. The runs and dances point directly toward the resource in these species. Each honey bee species has a characteristically different correlation of "waggling" to distance, as well.<ref>{{cite journal | last1 = Dyer | first1 = F.C. | last2 = Seeley | first2 = T.D. | year = 1991 | title = Dance dialects and foraging range in three Asian honey bee species | journal = Behavioral Ecology and Sociobiology | volume = 28 | issue = 4| pages = 227–233 | doi=10.1007/bf00175094| bibcode = 1991BEcoS..28..227D | s2cid = 23263610 }}</ref> Such species-specific behavior suggests that this form of communication does not depend on learning but is rather determined genetically. It also suggests how the dance may have [[Evolution|evolved]].

Other experiments further document the communicative nature of the waggle dance. For example, dances by robotic dummy bees induced some recruitment.<ref>{{cite journal | last1 = Michelsen | first1 = A. | last2 = Anderson | first2 = B. B. | last3 = Kirchner | first3 = W. H. | last4 = Lindauer | first4 = M. | year = 1989 | title = Honeybees can be recruited by a mechanical model of a dancing bee | journal = Naturwissenschaften | volume = 76 | issue = 6| pages = 277–280 | doi = 10.1007/BF00368642 | bibcode=1989NW.....76..277M| s2cid = 22396565 }}</ref> Research has also shown that the dance may vary with the environmental context, a finding that may explain why the results of some earlier studies were inconsistent.<ref>Visscher, P.K. and Tanner, D.A. (2004). Sensory aspects of recruitment-dance performance in honey bees ''(Apis mellifera)''. in: Hartfelder, K.H, De Jong, D. et al. eds. (2004) Proceedings of the 8th IBRA International Conference on Tropical Bees and VI Encontro sobre Abelhas. Ribierao Preto: USP/FM</ref><ref>{{cite journal | last1 = Sherman | first1 = G. | last2 = Visscher | first2 = P.K. | year = 2002 | title = Honeybee colonies achieve fitness through dancing | journal = Nature | volume = 419 | issue = 6910| pages = 920–922 | doi=10.1038/nature01127|bibcode = 2002Natur.419..920S | pmid=12410309| s2cid = 4431247 }}</ref>

===Odor plume ===
While many researchers believe that bee dances give enough information to locate resources, proponents of the odor plume theory argue that the dance gives little, or no actual guidance to a [[nectar source]]. They argue that bees instead are primarily recruited by odor. The purpose of the dance is simply to gain attention to the returning worker bee so she can share the odor of the nectar with other workers who will then follow the odor trail to the source. Most scientists agree that odor is used in recruitment to resources, but they differ strongly in opinion as to the information content of the dance.{{citation needed|date=November 2015}}

The primary lines of evidence used by the odor plume advocates are
#experiments with odorless sugar sources which show that worker bees are unable to recruit to those sources<ref>{{cite web|url=http://www.beesource.com/point-of-view/adrian-wenner/experiments-on-directing-bee-flight-by-odors/|title=Experiments on Directing Bee Flight by Odors|work=beesource.com|date=24 February 2016 }}</ref> and
#logical difficulties of a small-scale dance (a few centimeters across) giving directions precise enough to hold the other bees on course during a flight that could be several kilometers long. Misreading by even a few degrees would lead the bee off course by hundreds of meters at the far end. {{Citation needed|date=October 2010}}

Neither of these points invalidate the dance theory, but simply suggest that odor might be involved, which is indeed conceded by all proponents of dance theory.{{citation needed|date=November 2015}} Critics of the odor plume theory counter that most natural nectar sources are relatively large—orchards or entire fields— so, precision may not be necessary or even desirable. They have also challenged the reproducibility of the odorless source experiment.

Odor learning in bees is usually tested by the [[proboscis extension reflex]]. Significant to the argument are the experiments of William F. Towne, of the Kutztown University in Pennsylvania,<ref>{{cite journal |last1=Towne |first1=William F. |title=Honeybees can learn the relationship between the solar ephemeris and a newly-experienced landscape |journal=Journal of Experimental Biology |date=1 December 2008 |volume=211 |issue=23 |pages=3737–3743 |doi=10.1242/jeb.003640 |url=https://faculty.kutztown.edu/towne/Towne_2008_JEB_211_3737-3743.pdf |access-date=3 November 2022 |publisher=Kutztown University of Pennsylvania |pmid=19011214 |s2cid=11997761 |language=en-us}}</ref> in which hives are moved to "mirror image" terrain settings, and the bees are thereby fooled into dancing about the wrong location for a nectar source.  Foragers were successfully recruited to the wrong location, but only when the sun was obscured by clouds, forcing them to rely on terrain-based navigation rather than "[[solar ephemeris]]"-based navigation. As the cloud cover broke up, more and more bees corrected their dances to indicate the actual location of nectar, and forager visits shifted to the correct location.

Odor is essential and even necessary at various stages of the recruitment process, including once a recruited forager reaches the vicinity of the resource<ref>{{cite journal | last1 = Riley | first1 = J.R. | last2 = Greggers | first2 = U. | last3 = Smith | first3 = A.D. | last4 = Reynolds | first4 = D.R. | last5 = Menzel | first5 = R. | year = 2005 | title = The flight paths of honeybees recruited by the waggle dance | journal = Nature | volume = 435 | issue = 7039| pages = 205–207 | doi=10.1038/nature03526|bibcode = 2005Natur.435..205R | pmid=15889092| s2cid = 4413962 }}</ref> while some scientists think that dancing may be a simple [[idiothetic]] movement that conveys no information.<ref>{{cite web|url=https://archive.today/20131014004716/http://naturalscience.com/ns/articles/01-13/ns_jdo.html|title=Why do honeybees dance?}}</ref> Others see the dance as conveying information, but doing it poorly compared to other means and potentially used backup approach.<ref>{{Cite journal |url=https://www.newscientist.com/article/mg20327262.400-rethinking-the-bees-waggle-dance.html |archive-url=https://web.archive.org/web/20090923193107/http://www.newscientist.com/article/mg20327262.400-rethinking-the-bees-waggle-dance.html?full=true |archive-date=2009-09-23 |title=Rethinking the bee's waggle dance |date=18 September 2009 |first=Caroline |last=Williams |issue=2726 |journal=[[New Scientist]] |url-status=dead }} {{subscription required}}</ref>

Note: much of the research on the two competing hypotheses of communication has been restricted to [[Western honey bee]]s (see the work of F.C. Dyer<ref>[http://www.msu.edu/~fcdyer/pubs.html Publications of Fred C. Dyer]. {{webarchive |url=https://web.archive.org/web/20060914171408/http://www.msu.edu/~fcdyer/pubs.html |date=September 14, 2006 }}</ref> though). Other species of ''[[Apis (genus)|Apis]]'' use variants on the same theme, and other types of [[bee]]s use other methods altogether.

===Trophallaxis===
The exchange of food, [[trophallaxis]], can be used to communicate the quality of a food source, temperature, a need for water, and the condition of the queen (Sebeok, 1990).

====Primer pheromones====
{{for|more background on this topic|List of honey bee pheromones}}

Research that was published in November 2004, by scientists under the leadership of [[Zachary Huang]], [[Michigan State University]] indicates that so called primer [[pheromone]]s play an important part in how a honey bee [[colony (biology)|colony]] adjusts its distribution of labor most beneficially.  In order to survive as a bee colony of sometimes 50,000–100,000 individual bees, the communal structure has to be adaptable to seasonal changes and the availability of food.  The division of labor has to adjust itself to the resources available from foraging.  While the division of labor in a bee colony is quite complex, the work can be roughly seen as work inside the hive and outside the hive.  Younger bees play a role inside the hive while older bees play a role outside the hive mostly as [[foraging|foragers]].  Huang's team found that forager bees gather and carry a chemical called [[ethyl oleate]] in the stomach.  The forager bees feed this primer pheromone to the worker bees, and the chemical keeps them in a nurse bee state.  The pheromone prevents the nurse bees from maturing too early to become forager bees.  As forager bees die off, less of the ethyl oleate is available and nurse bees more quickly mature to become foragers.  It appears that this control system is an example of decentralized decision making in the bee colony.

Other bees like ''[[Trigona corvina]]'' rely on pheromones for much of their communication with nest mates and rivals.<ref name=":0">{{Cite journal |last1=Boogert |first1=Neeltje Janna |last2=Hofstede |first2=Frouke Elisabeth |last3=Monge |first3=Ingrid Aguilar |date=2006-05-01 |title=The use of food source scent marks by the stingless bee Trigona corvina (Hymenoptera: Apidae): the importance of the depositor's identity |journal=Apidologie |language=en |volume=37 |issue=3 |pages=366–375 |doi=10.1051/apido:2006001 |s2cid=4873234 |issn=0044-8435|doi-access=free }}</ref> They produce pheromones from their labial glands.<ref name=":1">{{Cite journal|title = The Trail Pheromone of a Stingless Bee, Trigona corvina (Hymenoptera, Apidae, Meliponini), Varies between Populations|journal = Chemical Senses|date = 2010-09-01|issn = 0379-864X|pmid = 20534775|pages = 593–601|volume = 35|issue = 7|doi = 10.1093/chemse/bjq057|first1 = Stefan|last1 = Jarau|first2 = Jochen|last2 = Dambacher|first3 = Robert|last3 = Twele|first4 = Ingrid|last4 = Aguilar|first5 = Wittko|last5 = Francke|first6 = Manfred|last6 = Ayasse|doi-access = free}}</ref> The function of signaling depends on the profitability, but they commonly will scent mark a food source either for self-orientation, to deter rivals or to direct a nest mate to the resource. Once an individual finds a good food source, they will return to the same source for many days. If an individual detects the scent of a rival bee, they will avoid the plant in order to avoid conflict and to save time.<ref name=":0" /> It has also been shown that pheromones are a method of sexual selection between male drones and queens.<ref name=":1" />