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Bee learning and communication
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===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.
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