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Bee learning and communication

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in his lab placed the sugar water on a boat in the middle of a small lake. When scouts returned to the hive to communicate their find, Gould claimed that other bees refused to go with them, even though they frequently flew over the lake to reach pollen sources on the opposite shore. To Gould, these observations suggested that "bees somehow consider information to see if it makes sense before they act on it". No one observed behavior of bees in the hive during this experiment, however, and when the experiment was replicated by Wray et al. in 2008, the researchers found no evidence for Gould's assertion that followers of waggle dances advertising "implausible" sites refused to respond to these dances. The authors' conclusion: "Contrary to prior findings, our results provide no evidence that honeybees assess the plausibility of information contained in waggle dances or reject dances for locations that are unlikely to yield food. Thus, we conclude that the original Lake Experiment should no longer be cited as evidence that honeybees possess cognitive maps, ‘insight’ or ‘imagination’."
180:. He performed an elegant experiment that showed not only that the bees could discriminate colors but that they demonstrated associative learning. He first trained his bees to feed from a small dish filled with a nectar-like sugar water. This dish was placed on a piece of blue colored cardboard so that the color was visible to the bees as they arrived at the dish and fed. Next, von Frisch placed identically sized pieces of cardboard in varying shades of grey, each with a dish, all around the blue piece. Lacking color vision, the bees should visit one or more of the gray pieces as often as the blue piece, but he found the vast majority of the bees flew directly to the blue piece of cardboard on which they had previously obtained their reward. The bees largely ignored the gray pieces which had not been rewarded. Von Frisch repeated the experiment with other colors like violet and yellow and got the same results. Later other researchers used this experimental design to test the color vision of vertebrates. 141:
choice between two patterns (matching and non-matching) placed at different distances (Can bees trained in the task continue to perform correctly when the matching pattern as well as a non-matching pattern are presented in the tunnel?); and a choice between two patterns (Can bees learn which of two sequentially encountered patterns in the tunnel is the pattern to be matched in the decision cylinder?). The researchers found that working memory in the honey bee is both robust and flexible. The experiments demonstrated that bees can choose between alternatives, determine if a stimulus is the same or different than one seen earlier, remember the earlier one for a short period, and generalize this performance to new pairs of stimuli. The bees retained information in working memory for about 5 seconds, and they might have been simultaneously learning a matching and a nonmatching task; further research was needed.
193: 918: 100: 386:(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. 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 551: 2834: 20: 429:. Significant to the argument are the experiments of William F. Towne, of the Kutztown University in Pennsylvania, 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 " 321: 168: 563:
pigments which have maximal absorbance at wavelengths of 350 nm, 440 nm, and 540 nm. Menzel also found that most of the cells he studied had secondary sensitivities that corresponded to wavelength regions at which the other two receptor types were maximally active. He used spectral efficiency experiments to suggest that these secondary sensitivities result from electric coupling between the receptors.
406:. 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. 521:
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. It has also been shown that pheromones are a method of sexual selection between male drones and queens.
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a different colored background. One of the colors was the same as that used during the initial trial, and the others were novel, unrewarded colors. Remarkably, after a single trial and several days without exposure to the rewarded color, bees correctly chose to explore the color used in the first trial more than fifty-percent of the time.
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dish could not be seen but the hive was still visible. When released one by one, the bees appeared disoriented for a few seconds, then flew directly to the dish, 73 of 75 bees reaching it in about 28 seconds. They apparently accomplished this feat by devising a new flight path based on a cognitive map of visible landmarks.
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by marking individual cells with a fluorescent dye and recording from these cells as single units. From this analysis he determined that there are three types of receptors in the honey bee eye: 1) UV receptors, 2) blue receptors, and 3) green receptors The three receptors contain three rhodopsin-like
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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
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Menzel then repeated this experiment with another group of bees, keeping all factors the same except that in the second round of testing he gave the bees three initial trials with the rewarded color instead of just one. When, after several days in confinement, the bees were presented with a choice of
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Menzel did several experiments to answer these questions. First, he gave individual bees a single sugar reward on a colored background. He then kept these bees in small cages for several days without any further trials. After a few days, he presented each bee with an array of several dishes, each on
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Interpretations of another test suggested not only the use of a map, but also an ability to remember and combine relevant information. Gould moved a supply of sugar water 25% farther away from a hive each day. The bees communicated the location of the water to each other as usual. Then a researcher
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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. Critics of the odor plume theory counter that most natural nectar sources are relatively large—orchards or entire fields— so, precision may
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Other experiments further document the communicative nature of the waggle dance. For example, dances by robotic dummy bees induced some recruitment. 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
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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
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To test bees ability to distinguish between two different colors, Menzel placed a small dish containing sugar-water in one circle and a second empty dish some distance away on a differently colored circle. A single bee was placed equidistant between the two circles and allowed to choose between the
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The German scientist Randolf Menzel continued the study of color vision in honey bees with more detailed tests. He was curious about whether bees would learn certain colors faster than others. He used lights of various color and intensity to project circles of light on a surface, a set-up like that
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for information they have learned, and utilize it when foraging. In an experimental demonstration, Gould lured some bees to a dish of artificial nectar, then gradually moved it farther from the hive. He marked the trained bees, placed them in a darkened jar, and relocated them to a spot where the
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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
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also showed that they discriminated between rewarding and unrewarding patterns. This knowledge persisted, both after 24 hours of learning and after 7 days. Dr. Muth's studies have also shown that bumblebees do not prefer nor choose against a flower based on its complexity. However, they will learn
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In still other experiments, Menzel explored the timing of bee color learning by testing whether bees register color before, during, or after receiving their sugar-water reward. For this purpose Menzel displayed the color beneath a rewarded dish at different stages of the honey bee feeding process:
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of honey bees, after learning to associate a certain pattern with a reward (delayed matching-to-sample). Bees were shown a pattern at the beginning of a tunnel, and then subjected to a series of variations: in the length of the tunnel (How long can bees retain the pattern in working memory?), in a
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requires such learning. For example, honey bees make few repeat visits to a plant if it provides little in the way of reward. A single forager will visit different flowers in the morning and, if there is sufficient reward in a particular kind of flower, she will make visits to that type of flower
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rely on pheromones for much of their communication with nest mates and rivals. They produce pheromones from their labial glands. 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
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After his work on color preferences, Menzel extended his experiments to study aspects of color learning and memory. He wanted to know how many trials bees need to reliably choose a previously rewarded color when they are presented with several alternatives, and how long they would remember the
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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
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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.
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Menzel's results showed that bees do not learn to discriminate between all color pairs equally well. Bees learned the fastest when violet light was rewarded, and the slowest when the light was green; the other colors fell somewhere in between. This evidence of inherent bias is evolutionarily
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Menzel found that bees register color during both approach and feeding, and that they had to see the color for about a total of about 5 seconds, with best performance usually coming with about three seconds exposure during the approach and two seconds after landing and beginning to feed.
257:. Bumblebees were shown to be able to learn multiple color-food associations and tended to continue to apply what they learned. In another study, Dr. Felicity Muth continued to learn more about these associations. Bumblebees initially preferred yellow 502:
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
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Testing for color vision in honey bees. The majority of bees flew directly to the dish with the blue background as they had been trained to do. Thus, they were able to discriminate between gray and blue backgrounds, showing their capability for color
328:). 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. 225:
This ability to retain information about color-linked rewards for several days after minimal exposure to the rewarded color demonstrates the remarkable facility with which bees learn and retain color information.
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Odor is essential and even necessary at various stages of the recruitment process, including once a recruited forager reaches the vicinity of the resource while some scientists think that dancing may be a simple
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and retain this information for several days at least; they communicate the location and nature of sources of food; they adjust their foraging to the times at which food is available; they may even form
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Wray MK, Klein BA, Seeley TD. 2011. Honey bees use the social information in waggle dances more fully when foraging errors are more costly. Behavioral Ecology. 23:125-131. doi:10.1093/beheco/arr165
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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
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Menzel, R. and Backhaus, W. 1989. Color vision in honey bees: Phenomena and physiological mechanisms. In D. Stavenga and R. Hardie (eds): Facets of vision. Berlin-Heidelberg-New York: 281-297
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dishes. The bees quickly learned to choose the color signaling the dish with the reward, and Menzel was able to measure how quickly the bees learned this task with various color differences.
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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
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Muth, Felicity; Keasar, Tamar; Dornhaus, Anna (March 2015). "Trading off short-term costs for long-term gains: how do bumblebees decide to learn morphologically complex flowers?".
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A seminal paper by Menzel (1975) described the morphology and spectral sensitivity of the honey bee eye that underlie their color vision. He examined color-coding in the honey bee
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Menzel, R. and Backhaus, W. 1991. Colour Vision in Insects. In P. Gouras (ed): Vision and Visual Dysfunction. The Perception of Colour. London: MacMillan Press, 262-288.
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movement that conveys no information. Others see the dance as conveying information, but doing it poorly compared to other means and potentially used backup approach.
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cells were found to have long visual nerve fibers that penetrated the lamina with deep tree-like branchings. Blue and green receptor cells had more shallow fibers.
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reasonable, given that bees forage for differently-colored nectar-bearing flowers, many of which are to be found in green foliage which does not signal reward.
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to forage in the same area. The factors that determine recruiting success are not completely known but probably include evaluations of the quality of
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Leoncini I, Le Conte Y, Costagliola G, Plettner E, Toth AL, Wang M, Huang Z, Becard JM, Crauser D, Slessor KN, Robinson GE (December 14, 2004).
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used by von Frisch except that, by using light instead of cardboard, Menzel was able to easily change the intensity and color of the circles.
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The Austrian zoologist Karl von Frisch began the exploration of color vision in honey bees when, in 1919, he asked whether or not bees have
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Michelsen, A.; Anderson, B. B.; Kirchner, W. H.; Lindauer, M. (1989). "Honeybees can be recruited by a mechanical model of a dancing bee".
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Riley, J.R.; Greggers, U.; Smith, A.D.; Reynolds, D.R.; Menzel, R. (2005). "The flight paths of honeybees recruited by the waggle dance".
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quickly mature to become foragers. It appears that this control system is an example of decentralized decision making in the bee colony.
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Later work has supported Von Frisch's observations and added many details. It appears that all of the known species and races of
346:. The laden forager dances on the comb in a circular pattern, occasionally crossing the circle in a zig-zag or waggle pattern. 1567: 1241: 948: 715: 364: 1266: 889:
Munz, T. (November 2005). "The Bee Battles: Karl von Frisch, Adrian Wenner and the Honey Bee Dance Language Controversy".
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An American specialist in bee cognition, Dr. Felicity Muth, has studied the mechanism behind the associative learning in
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not be necessary or even desirable. They have also challenged the reproducibility of the odorless source experiment.
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and later Randolf Menzel began asking questions about color vision and various aspects of color learning in bees.
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Genetic Control of the Honey Bee (apis mellifera) Dance Language: Segregating Dance Forms in a Backcrossed Colony
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Jarau, Stefan; Dambacher, Jochen; Twele, Robert; Aguilar, Ingrid; Francke, Wittko; Ayasse, Manfred (2010-09-01).
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Frisch, K. von. 1956. Bees; their vision, chemical senses, and language. Ithaca, N.Y., Cornell University Press.
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irrelevant to recruitment. The academic debate between these theories has been polarized and sometimes hostile.
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experiments with odorless sugar sources which show that worker bees are unable to recruit to those sources and
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Menzel, R; Blakers, M (1975). "Colour receptors in the bee eye — Morphology and spectral sensitivity".
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Visscher, P.K. and Tanner, D.A. (2004). Sensory aspects of recruitment-dance performance in honey bees
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Dyer, F.C.; Seeley, T.D. (1991). "Dance dialects and foraging range in three Asian honey bee species".
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exhibit the behavior, but details of its execution vary among the different species. For example, in
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A number of experiments have demonstrated color recognition, discrimination and memory in honey bees
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Note: much of the research on the two competing hypotheses of communication has been restricted to
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Currently the group is working on four projects in the area of mid-air collision avoidance...
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for most of the day, unless the plants stop producing nectar or weather conditions change.
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https://www.apiservices.biz/en/articles/sort-by-popularity/562-why-do-honeybees-dance-2000
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Sebeok (1990). Essays in Zoosemiotics. Toronto: Toronto Semiotic Circle. ISSN 0838-5858 .
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Sherman, G.; Visscher, P.K. (2002). "Honeybee colonies achieve fitness through dancing".
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Boogert, Neeltje Janna; Hofstede, Frouke Elisabeth; Monge, Ingrid Aguilar (2006-05-01).
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Foragers communicate their floral findings in order to recruit other worker bees of the
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Anthophila. Some species have been studied more extensively than others, in particular
1816: 1512: 1477: 823:"Bees remember flowers for more than one reason: pollen mediates associative learning" 668: 633: 2837: 2544: 2512: 2405: 2256: 2244: 1969: 1964: 1959: 1949: 1903: 1872: 1754: 1620: 1517: 1388: 1380: 1328: 1215: 1157: 1089: 910: 803: 772:"Colour learning when foraging for nectar and pollen: bees learn two colours at once" 711: 673: 445: 333: 1709: 1169: 1037: 994: 875: 485:
Research that was published in November 2004, by scientists under the leadership of
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colors they almost always chose the color that was used on the first three trials.
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Gould, J L "A honey of a question, are bees intelligent?" Discover, 1986, August
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There are two main hypotheses to explain how foragers recruit other workers—the
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Proceedings of the National Academy of Sciences of the United States of America
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Proceedings of the National Academy of Sciences of the United States of America
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Miller, J A "Biology" Science News; 4/23/1983, Vol. 123 Issue 17, p271, 1/6p
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Swarming bees require good communication to all congregate in the same place
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of their surroundings. They also communicate with each other by means of a "
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Miller, Julie Ann (April 23, 1983). "Do Bees Plan Ahead Intelligently?".
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Behavioral Neurobiology: The Cellular Organization of Natural Behavior
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The primary lines of evidence used by the odor plume advocates are
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Muth, Felicity; Papaj, Daniel; Leonard, Anne (1 September 2015).
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Carew, Thomas J. (2000). "9. Associative Learning in Honeybees".
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those unique traits if the reward, the pollen, is great enough.
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Giurfa, Martin; Zhang, Shaowu; Jenett, Arnim; Menzel, Randolf;
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Muth, Felicity; Papaj, Daniel; Leonard, Anne (January 2016).
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Certain morphologies distinguished the receptor types.
1846: 1720: 567: 183: 1148:(23). Kutztown University of Pennsylvania: 3737–3743. 627: 448:(see the work of F.C. Dyer though). Other species of 1781:"The Sensory Basis of the Honeybee's Dance Language" 1575:
Ibbotson, M.R.; Hung, Y.; Meffin, H.; N. Boeddeker;
1304: 1686:"The Elusive Honey Bee Dance "Language" Hypothesis" 454:use variants on the same theme, and other types of 119:take the same form in honey bees as they do in the 853: 332:It has long been known that successfully foraging 324:Figure-Eight-Shaped waggle dance of the honeybee ( 1779:Kirchner, Wolfgang H.; Towne, William F. (2000). 1566:(On the idiothetic hypothesis) O'Dea, JD (2000) " 1528: 2850: 820: 769: 23:Bees learn and communicate in a variety of ways. 1063: 545: 425:Odor learning in bees is usually tested by the 242: 1116:"Experiments on Directing Bee Flight by Odors" 144: 2769:Association for the Study of Animal Behaviour 2413: 1832: 1437: 1433: 1431: 1429: 1633:"Neuroscience of vision and aerial robotics" 731: 729: 727: 701: 699: 697: 695: 693: 691: 689: 687: 229: 155:. Beginning in the early 1900s, scientists 2774:International Society for Applied Ethology 2420: 2406: 1839: 1825: 1426: 972: 961:The Dance Language and Orientation of Bees 497:play an important part in how a honey bee 361:The Dance Language and Orientation of Bees 1614: 1537:(17). Society for Science &#38: 271. 1511: 1501: 1374: 1322: 838: 797: 787: 754: 667: 657: 1255:Williams, Caroline (18 September 2009). 1254: 949:The Honey Bee Dance Language Controversy 747: 745: 743: 741: 724: 684: 549: 319: 235:during approach, feeding and departure. 191: 166: 98: 18: 478:For more background on this topic, see 162: 16:Cognitive and sensory processes in bees 2851: 1802:"Pleasure chemical controls bee dance" 1683: 315: 2401: 1820: 1799: 1352: 1350: 1300: 1298: 1133:Towne, William F. (1 December 2008). 1132: 963:. Cambridge, MA: Harvard Univ. Press. 738: 705: 365:Nobel Prize in Physiology or Medicine 888: 473: 184:Color learning rates and preferences 1684:Wenner, Adrian M. (November 2002). 1440:Journal of Comparative Physiology A 1257:"Rethinking the bee's waggle dance" 975:Behavioral Ecology and Sociobiology 533:suggest that honey bees may have a 136:A 2005 three-part study tested the 42:, that is the insects in the seven 13: 1561: 1347: 1295: 309:"waggle dance" or "dance language" 66:are sensitive to odors (including 14: 2880: 1923:Subspecies, Breeds and Phenotypes 1800:Hayes, Jacqui (16 January 2007). 891:Journal of the History of Biology 628:Zhang, S; Bock F; Si A; Tautz J; 70:), tastes, and colors, including 2833: 2832: 916: 493:indicates that so called primer 350:described this behaviour in his 281: 1469: 1417: 1408: 1399: 1276: 1248: 1234: 1183: 1142:Journal of Experimental Biology 1126: 1108: 1057: 1044: 1001: 966: 953: 942: 924: 461: 363:and in 1973 he was awarded the 208: 111:, and many of the phenomena of 2478:Bee learning and communication 1909:Bee learning and communication 1677:Johnson, R. N. et al. (2002) 882: 847: 814: 763: 621: 458:use other methods altogether. 28:Bee learning and communication 1: 868:10.1016/j.anbehav.2014.12.024 840:10.1016/j.anbehav.2015.09.029 615: 397: 2427: 1283:Publications of Fred C. Dyer 546:Neurobiology of color vision 524: 480:List of honey bee pheromones 243:Color learning in bumblebees 7: 2053:Apis mellifera sinisxinyuan 1289:September 14, 2006, at the 573: 196:Honey bee collecting pollen 145:Color learning in honeybees 94: 10: 2887: 2864:Western honey bee behavior 1690:Journal of Insect Behavior 1657:Queensland Brain Institute 1637:Queensland Brain Institute 1607:10.1038/s41598-017-14954-0 477: 427:proboscis extension reflex 285: 38:processes in all kinds of 2828: 2782: 2761: 2640: 2535:Evolutionary neuroscience 2435: 2387:Honeybee Discovery Center 2364: 2306: 2298:Diseases of the honey bee 2285: 2237: 2146: 2095: 2032:Apis mellifera scutellata 1922: 1886: 1855: 1850:types and characteristics 1242:"Why do honeybees dance?" 903:10.1007/s10739-005-0552-1 491:Michigan State University 342:upon their return to the 131: 2488:Behavioral endocrinology 2365:Museums and insectariums 2277:Colony collapse disorder 2252:Varroa sensitive hygiene 2039:Apis mellifera siciliana 2018:Apis mellifera monticola 2004:Apis mellifera pomonella 1990:Apis mellifera artemisia 1983:Apis mellifera adansonii 1667:"Honeybee Communication" 1661:University of Queensland 230:Timing in color learning 107:Honey bees are adept at 2683:Irenäus Eibl-Eibesfeldt 2463:Animal sexual behaviour 2171:Horizontal top-bar hive 2081:Apis mellifera unicolor 2060:Apis mellifera sossimai 2046:Apis mellifera simensis 1702:10.1023/A:1021131725124 1503:10.1073/pnas.0407652101 1272:(subscription required) 659:10.1073/pnas.0501440102 580:Bumblebee communication 288:Bumblebee communication 2622:Tool use by non-humans 2575:Philosophical ethology 2520:Comparative psychology 2468:Animal welfare science 2238:Parasites and diseases 2074:Apis mellifera taurica 2067:Apis mellifera syriaca 2025:Apis mellifera remipes 1997:Apis mellifera litorea 1723:Srinivasan, Mandyam V. 1265:(2726). Archived from 959:von Frisch, K. (1967) 789:10.1098/rsbl.2015.0628 710:. Sinauer Associates. 555: 466:The exchange of food, 329: 197: 173: 117:classical conditioning 104: 24: 1376:10.1093/chemse/bjq057 1324:10.1051/apido:2006001 553: 367:for his discoveries. 323: 195: 170: 102: 91:" and in other ways. 22: 2859:Animal communication 2728:William Homan Thorpe 2493:Behavioural genetics 2453:Animal consciousness 2448:Animal communication 2377:Bee Museum of Rhodes 2293:Topics in beekeeping 1899:Honey bee life cycle 932:"HISTORIA ANIMALIUM" 163:Color discrimination 109:associative learning 80:operant conditioning 2483:Behavioural ecology 2372:Malacca Bee Gallery 2272:Deformed wing virus 2163:BS National Beehive 2011:Apis mellifera meda 1945:Carniolan honey bee 1812:on 19 January 2007. 1792:Scientific American 1743:2001Natur.410..930G 1716:on 3 December 2003. 1673:on 22 January 2016. 1627:– via Nature. 1599:2017NatSR...714591I 1579:(6 November 2017). 1494:2004PNAS..10117559L 1212:10.1038/nature03526 1204:2005Natur.435..205R 1122:. 24 February 2016. 1086:10.1038/nature01127 1078:2002Natur.419..920S 1022:1989NW.....76..277M 1010:Naturwissenschaften 938:. 6 September 2008. 650:2005PNAS..102.5250Z 595:Trained hymenoptera 316:Dance communication 2812:Behavioral Ecology 2733:Nikolaas Tinbergen 2525:Emotion in animals 2503:Cognitive ethology 2267:American foulbrood 1586:Scientific Reports 1452:10.1007/bf00625437 1154:10.1242/jeb.003640 1030:10.1007/BF00368642 987:10.1007/bf00175094 556: 446:Western honey bees 383:Apis andreniformis 352:Historia Animalium 334:Western honey bees 330: 265:when foraging for 198: 174: 105: 25: 2846: 2845: 2738:Jakob von Uexküll 2508:Comfort behaviour 2395: 2394: 2257:Small hive beetle 2246:Varroa destructor 1970:Western honey bee 1965:Russian honey bee 1960:Maltese honey bee 1950:European dark bee 1873:Laying worker bee 1737:(6831): 930–933. 1725:(19 April 2001). 1198:(7039): 205–207. 1072:(6910): 920–922. 717:978-0-87893-084-5 632:(April 5, 2005). 554:Western honey bee 474:Primer pheromones 2876: 2836: 2835: 2798:Animal Cognition 2791:Animal Behaviour 2743:Wolfgang Wickler 2443:Animal cognition 2422: 2415: 2408: 2399: 2398: 2123:Honey extraction 1841: 1834: 1827: 1818: 1817: 1813: 1808:. Archived from 1788: 1770: 1751:10.1038/35073582 1717: 1712:. Archived from 1674: 1669:. Archived from 1655:'s group at the 1650: 1645: 1643: 1628: 1618: 1593:(14591): 14591. 1577:M. V. Srinivasan 1554: 1525: 1515: 1505: 1488:(50): 17559–64. 1464: 1463: 1435: 1424: 1421: 1415: 1412: 1406: 1403: 1397: 1396: 1378: 1354: 1345: 1344: 1326: 1302: 1293: 1280: 1274: 1273: 1270: 1252: 1246: 1245: 1238: 1232: 1231: 1187: 1181: 1180: 1178: 1176: 1139: 1130: 1124: 1123: 1112: 1106: 1105: 1061: 1055: 1052:(Apis mellifera) 1048: 1042: 1041: 1005: 999: 998: 970: 964: 957: 951: 946: 940: 939: 928: 922: 921: 920: 914: 886: 880: 879: 856:Animal Behaviour 851: 845: 844: 842: 827:Animal Behaviour 818: 812: 811: 801: 791: 767: 761: 758: 752: 749: 736: 733: 722: 721: 703: 682: 681: 671: 661: 625: 514:Other bees like 254:Bombus impatiens 214:rewarded color. 2886: 2885: 2879: 2878: 2877: 2875: 2874: 2873: 2849: 2848: 2847: 2842: 2824: 2778: 2757: 2753:Solly Zuckerman 2693:Karl von Frisch 2678:Richard Dawkins 2663:John B. Calhoun 2648:Patrick Bateson 2636: 2570:Pain in animals 2431: 2426: 2396: 2391: 2360: 2308: 2302: 2281: 2233: 2219:Honey extractor 2175:Langstroth hive 2142: 2091: 1930:Africanized bee 1918: 1882: 1851: 1845: 1806:COSMOS magazine 1665: 1653:M.V. Srinivasan 1641: 1639: 1631: 1564: 1562:Further reading 1543:10.2307/3967590 1472: 1467: 1436: 1427: 1422: 1418: 1413: 1409: 1404: 1400: 1363:Chemical Senses 1355: 1348: 1303: 1296: 1291:Wayback Machine 1281: 1277: 1271: 1253: 1249: 1240: 1239: 1235: 1188: 1184: 1174: 1172: 1137: 1131: 1127: 1114: 1113: 1109: 1062: 1058: 1049: 1045: 1006: 1002: 971: 967: 958: 954: 947: 943: 930: 929: 925: 915: 887: 883: 852: 848: 819: 815: 776:Biology Letters 768: 764: 759: 755: 750: 739: 734: 725: 718: 704: 685: 626: 622: 618: 576: 548: 529:Experiments by 527: 517:Trigona corvina 483: 476: 464: 431:solar ephemeris 400: 356:Karl von Frisch 318: 290: 284: 251:, specifically 245: 232: 211: 186: 165: 157:Karl von Frisch 147: 134: 97: 17: 12: 11: 5: 2884: 2883: 2872: 2871: 2866: 2861: 2844: 2843: 2841: 2840: 2829: 2826: 2825: 2823: 2822: 2815: 2808: 2805:Animal Welfare 2801: 2794: 2786: 2784: 2780: 2779: 2777: 2776: 2771: 2765: 2763: 2759: 2758: 2756: 2755: 2750: 2745: 2740: 2735: 2730: 2725: 2720: 2718:Desmond Morris 2715: 2710: 2705: 2700: 2695: 2690: 2685: 2680: 2675: 2673:Marian Dawkins 2670: 2668:Charles Darwin 2665: 2660: 2655: 2650: 2644: 2642: 2638: 2637: 2635: 2634: 2629: 2624: 2619: 2614: 2613: 2612: 2607: 2602: 2597: 2587: 2582: 2577: 2572: 2567: 2562: 2557: 2552: 2550:Human ethology 2547: 2542: 2537: 2532: 2527: 2522: 2517: 2516: 2515: 2505: 2500: 2495: 2490: 2485: 2480: 2475: 2470: 2465: 2460: 2458:Animal culture 2455: 2450: 2445: 2439: 2437: 2433: 2432: 2425: 2424: 2417: 2410: 2402: 2393: 2392: 2390: 2389: 2384: 2379: 2374: 2368: 2366: 2362: 2361: 2359: 2358: 2353: 2351:United Kingdom 2348: 2343: 2338: 2333: 2328: 2323: 2318: 2312: 2310: 2304: 2303: 2301: 2300: 2295: 2289: 2287: 2283: 2282: 2280: 2279: 2274: 2269: 2264: 2259: 2254: 2249: 2241: 2239: 2235: 2234: 2232: 2231: 2226: 2224:Queen excluder 2221: 2216: 2211: 2206: 2201: 2199:Wax foundation 2196: 2191: 2186: 2183:Stewarton hive 2156: 2150: 2148: 2144: 2143: 2141: 2140: 2135: 2130: 2125: 2120: 2115: 2110: 2105: 2099: 2097: 2093: 2092: 2090: 2089: 2087:Honey bee race 2084: 2077: 2070: 2063: 2056: 2049: 2042: 2035: 2028: 2021: 2014: 2007: 2000: 1993: 1986: 1979: 1976:Apis laboriosa 1972: 1967: 1962: 1957: 1952: 1947: 1942: 1940:Cape honey bee 1937: 1932: 1926: 1924: 1920: 1919: 1917: 1916: 1911: 1906: 1901: 1896: 1890: 1888: 1884: 1883: 1881: 1880: 1875: 1870: 1865: 1859: 1857: 1853: 1852: 1844: 1843: 1836: 1829: 1821: 1815: 1814: 1797: 1776: 1718: 1696:(6): 859–878. 1681: 1675: 1663: 1629: 1563: 1560: 1559: 1558: 1555: 1526: 1471: 1468: 1466: 1465: 1425: 1416: 1407: 1398: 1369:(7): 593–601. 1346: 1317:(3): 366–375. 1294: 1275: 1269:on 2009-09-23. 1247: 1233: 1182: 1125: 1107: 1056: 1043: 1016:(6): 277–280. 1000: 981:(4): 227–233. 965: 952: 941: 923: 897:(3): 535–570. 881: 846: 813: 762: 753: 737: 723: 716: 683: 644:(14): 5250–5. 619: 617: 614: 613: 612: 607: 602: 597: 592: 590:Grooming dance 587: 582: 575: 572: 547: 544: 526: 523: 475: 472: 463: 460: 419: 418: 414: 399: 396: 394:inconsistent. 326:Apis mellifera 317: 314: 283: 280: 244: 241: 231: 228: 210: 207: 185: 182: 164: 161: 152:Apis mellifera 146: 143: 138:working memory 133: 130: 96: 93: 85:cognitive maps 53:Apis mellifera 46:making up the 15: 9: 6: 4: 3: 2: 2882: 2881: 2870: 2867: 2865: 2862: 2860: 2857: 2856: 2854: 2839: 2831: 2830: 2827: 2821: 2820: 2816: 2814: 2813: 2809: 2807: 2806: 2802: 2800: 2799: 2795: 2793: 2792: 2788: 2787: 2785: 2781: 2775: 2772: 2770: 2767: 2766: 2764: 2760: 2754: 2751: 2749: 2746: 2744: 2741: 2739: 2736: 2734: 2731: 2729: 2726: 2724: 2723:Thomas Sebeok 2721: 2719: 2716: 2714: 2713:Konrad Lorenz 2711: 2709: 2708:Julian Huxley 2706: 2704: 2703:Heini Hediger 2701: 2699: 2696: 2694: 2691: 2689: 2686: 2684: 2681: 2679: 2676: 2674: 2671: 2669: 2666: 2664: 2661: 2659: 2656: 2654: 2651: 2649: 2646: 2645: 2643: 2639: 2633: 2632:Zoomusicology 2630: 2628: 2625: 2623: 2620: 2618: 2615: 2611: 2608: 2606: 2603: 2601: 2598: 2596: 2593: 2592: 2591: 2588: 2586: 2583: 2581: 2578: 2576: 2573: 2571: 2568: 2566: 2565:Neuroethology 2563: 2561: 2558: 2556: 2553: 2551: 2548: 2546: 2543: 2541: 2538: 2536: 2533: 2531: 2528: 2526: 2523: 2521: 2518: 2514: 2511: 2510: 2509: 2506: 2504: 2501: 2499: 2496: 2494: 2491: 2489: 2486: 2484: 2481: 2479: 2476: 2474: 2473:Anthrozoology 2471: 2469: 2466: 2464: 2461: 2459: 2456: 2454: 2451: 2449: 2446: 2444: 2441: 2440: 2438: 2434: 2430: 2423: 2418: 2416: 2411: 2409: 2404: 2403: 2400: 2388: 2385: 2383: 2380: 2378: 2375: 2373: 2370: 2369: 2367: 2363: 2357: 2356:United States 2354: 2352: 2349: 2347: 2344: 2342: 2339: 2337: 2334: 2332: 2329: 2327: 2324: 2322: 2319: 2317: 2314: 2313: 2311: 2305: 2299: 2296: 2294: 2291: 2290: 2288: 2284: 2278: 2275: 2273: 2270: 2268: 2265: 2263: 2260: 2258: 2255: 2253: 2250: 2248: 2247: 2243: 2242: 2240: 2236: 2230: 2227: 2225: 2222: 2220: 2217: 2215: 2212: 2210: 2207: 2205: 2202: 2200: 2197: 2195: 2192: 2190: 2187: 2184: 2180: 2176: 2172: 2168: 2164: 2160: 2157: 2155: 2152: 2151: 2149: 2145: 2139: 2136: 2134: 2131: 2129: 2126: 2124: 2121: 2119: 2116: 2114: 2111: 2109: 2106: 2104: 2101: 2100: 2098: 2094: 2088: 2085: 2083: 2082: 2078: 2076: 2075: 2071: 2069: 2068: 2064: 2062: 2061: 2057: 2055: 2054: 2050: 2048: 2047: 2043: 2041: 2040: 2036: 2034: 2033: 2029: 2027: 2026: 2022: 2020: 2019: 2015: 2013: 2012: 2008: 2006: 2005: 2001: 1999: 1998: 1994: 1992: 1991: 1987: 1985: 1984: 1980: 1978: 1977: 1973: 1971: 1968: 1966: 1963: 1961: 1958: 1956: 1953: 1951: 1948: 1946: 1943: 1941: 1938: 1936: 1933: 1931: 1928: 1927: 1925: 1921: 1915: 1912: 1910: 1907: 1905: 1902: 1900: 1897: 1895: 1892: 1891: 1889: 1885: 1879: 1876: 1874: 1871: 1869: 1866: 1864: 1861: 1860: 1858: 1854: 1849: 1842: 1837: 1835: 1830: 1828: 1823: 1822: 1819: 1811: 1807: 1803: 1798: 1795: 1793: 1786: 1782: 1777: 1774: 1768: 1764: 1760: 1756: 1752: 1748: 1744: 1740: 1736: 1732: 1728: 1724: 1719: 1715: 1711: 1707: 1703: 1699: 1695: 1691: 1687: 1682: 1680: 1676: 1672: 1668: 1664: 1662: 1658: 1654: 1649: 1638: 1634: 1630: 1626: 1622: 1617: 1612: 1608: 1604: 1600: 1596: 1592: 1588: 1587: 1582: 1578: 1573: 1572: 1571: 1569: 1556: 1552: 1548: 1544: 1540: 1536: 1532: 1527: 1523: 1519: 1514: 1509: 1504: 1499: 1495: 1491: 1487: 1483: 1479: 1474: 1473: 1461: 1457: 1453: 1449: 1445: 1441: 1434: 1432: 1430: 1420: 1411: 1402: 1394: 1390: 1386: 1382: 1377: 1372: 1368: 1364: 1360: 1353: 1351: 1342: 1338: 1334: 1330: 1325: 1320: 1316: 1312: 1308: 1301: 1299: 1292: 1288: 1284: 1279: 1268: 1264: 1263: 1262:New Scientist 1258: 1251: 1243: 1237: 1229: 1225: 1221: 1217: 1213: 1209: 1205: 1201: 1197: 1193: 1186: 1171: 1167: 1163: 1159: 1155: 1151: 1147: 1143: 1136: 1129: 1121: 1120:beesource.com 1117: 1111: 1103: 1099: 1095: 1091: 1087: 1083: 1079: 1075: 1071: 1067: 1060: 1053: 1047: 1039: 1035: 1031: 1027: 1023: 1019: 1015: 1011: 1004: 996: 992: 988: 984: 980: 976: 969: 962: 956: 950: 945: 937: 933: 927: 919: 912: 908: 904: 900: 896: 892: 885: 877: 873: 869: 865: 861: 857: 850: 841: 836: 832: 828: 824: 817: 809: 805: 800: 795: 790: 785: 781: 777: 773: 766: 757: 748: 746: 744: 742: 732: 730: 728: 719: 713: 709: 702: 700: 698: 696: 694: 692: 690: 688: 679: 675: 670: 665: 660: 655: 651: 647: 643: 639: 635: 631: 630:Srinivasan MV 624: 620: 611: 608: 606: 603: 601: 600:Tremble dance 598: 596: 593: 591: 588: 586: 583: 581: 578: 577: 571: 569: 564: 561: 552: 543: 539: 536: 535:cognitive map 532: 522: 519: 518: 512: 509: 505: 500: 496: 492: 488: 487:Zachary Huang 481: 471: 469: 459: 457: 453: 452: 447: 442: 440: 434: 432: 428: 423: 415: 412: 411: 410: 407: 405: 404:nectar source 395: 391: 389: 385: 384: 379: 378: 373: 368: 366: 362: 357: 353: 349: 345: 341: 340: 335: 327: 322: 313: 310: 305: 303: 299: 295: 289: 282:Communication 279: 276: 272: 268: 264: 260: 256: 255: 250: 240: 236: 227: 223: 219: 215: 206: 202: 194: 190: 181: 179: 169: 160: 158: 154: 153: 142: 139: 129: 126: 122: 118: 114: 110: 101: 92: 90: 86: 81: 77: 73: 69: 65: 61: 59: 55: 54: 49: 45: 41: 37: 33: 29: 21: 2817: 2810: 2803: 2796: 2789: 2748:E. O. Wilson 2698:Jane Goodall 2658:Donald Broom 2627:Zoosemiotics 2580:Sociobiology 2477: 2382:Honey Museum 2309:by countries 2245: 2079: 2072: 2065: 2058: 2051: 2044: 2037: 2030: 2023: 2016: 2009: 2002: 1995: 1988: 1981: 1974: 1935:Buckfast bee 1908: 1810:the original 1805: 1791: 1784: 1773:Researchgate 1734: 1730: 1714:the original 1693: 1689: 1671:the original 1651:The work of 1647: 1640:. Retrieved 1636: 1590: 1584: 1565: 1534: 1531:Science News 1530: 1485: 1481: 1470:Bibliography 1443: 1439: 1419: 1410: 1401: 1366: 1362: 1314: 1310: 1278: 1267:the original 1260: 1250: 1236: 1195: 1191: 1185: 1173:. Retrieved 1145: 1141: 1128: 1119: 1110: 1069: 1065: 1059: 1051: 1046: 1013: 1009: 1003: 978: 974: 968: 960: 955: 944: 936:virginia.edu 935: 926: 894: 890: 884: 859: 855: 849: 830: 826: 816: 779: 775: 765: 756: 707: 641: 637: 623: 610:Zoosemiotics 605:Waggle dance 565: 557: 540: 528: 515: 513: 508:ethyl oleate 484: 468:trophallaxis 465: 462:Trophallaxis 449: 443: 435: 424: 420: 408: 401: 392: 381: 375: 369: 360: 351: 339:waggle dance 337: 331: 306: 304:brought in. 291: 252: 246: 237: 233: 224: 220: 216: 212: 209:Color memory 203: 199: 187: 178:color vision 175: 150: 148: 135: 123:. 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Index


cognitive
sensory
bees
families
clade
Apis mellifera
bumblebees
Honey bees
pheromones
ultraviolet
classical
operant conditioning
cognitive maps
waggle dance

associative learning
operant
classical conditioning
vertebrates
foraging
working memory
Apis mellifera
Karl von Frisch
experimental design for testing color vision in honey bees.
color vision
bee collecting pollen.
bumblebees
Bombus impatiens
anthers

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