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Ryleigh Repass

Cognitive Ethology Final


A Critical Review
Honey bees have been an ever growing topic of discussion in many fields of study. Ever
since the discovery of the honey bee waggle dance in 1967 by Karl von. Frisch, researchers
have been working to determine what exactly the waggle dance is and how it works (Von
Frisch, 1967). However, it is generally known that bees portray direction and distance
information in their dances and they are able to recruit bees to the food location (Gould, 1975).
The overall importance of the waggle dance for foraging bees in collection of nectar is not quite
as well known. Therefore, Sherman and Visscher decide to test to see just how important the
waggle dance is in communication. Sherman and Visscher (2002) tests the hypothesis that the
nest is an information center where communication occurs to recruit foraging bees to the richest
food source, in turn increasing colony food collection.
Introduction:
Before they began their procedures, Sherman and Visscher (2002) wanted to see what
effect diffused light and uni-directional light had on orientation of the waggle dance. Diffused
light was from three 25-W bulbs diffused through white translucent Plexiglas suspended above
the colony and uni-directional light was from one 75-W bulb above a sheet of transparent
Plexiglas and a fluorescent black light providing short wave-lengths. They found that diffused
light caused bees to perform a completely disoriented dance and unidirectional light caused bees
to perform a well-oriented dance.
Methods:

Sherman and Visscher (2002) performed two types of procedures. Their first procedure
was to test to see if light treatment affected recruitment. They trained an equal number of
foragers to a feeder and tagged them for identification. The colonies each received opposite light
treatments for the first half of the observation and then were switched. Each unmarked recruit
was captured and recorded. Feeders were 250, 400, and 460 m from the colony for each trial
respectively. According to their hypothesis, Sherman and Visscher would hope to see less
recruits to the feeders when the bees were undergoing diffused light compared to when they were
exposed to directional light.
The second procedure compared foraging success at natural food sources for colonies
under alternating light treatments. One pair of colonies was given diffused light treatment and
the second pair of colonies was given oriented lights and they exchanged treatments
approximately every 11 days for 8 months. In order to test the success rate of the colonies, they
weighed the mass of the hives with an electronic scale. In order to support their hypothesis,
Sherman and Visscher would like to see a rise in mass during times of directional light and a
lowered mass during times of diffused light.
Results:
In their preliminary experiment to test the effects of the different types of lights, they
found that dances in the diffuse-light treatment did not differ from a circular random distribution
(p>0.5). Therefore, there was no directional orientation to the waggle runs when bees were
exposed to diffused light. The dances in the oriented light treatment and in the vertical comb
(control group) were highly directionally oriented (p<0.001 for oriented light). In other words,
diffused light treatment produced disoriented dances while oriented light treatment produced

oriented dances. Thus, Sherman and Visscher were able move onto testing their hypothesis with
support for the differing light treatments producing different dances.
In the test to see whether light treatments affected recruitment, they found significant
results. There were significantly more recruits that arrived at the feeders when the colony had
oriented light (thus, oriented dances) than diffused light (thus, disoriented dances) (p<0.02).
They found no time of day effects on recruit numbers (p<0.7) but found a colony effect in trial 1
(p<0.001) but not in trials two and three (p>0.25 and 0.15, respectively). Furthermore, to test
whether the light treatments affected the number of dances performed in each treatment, they
counted number of bees dancing in each colony. They found no significant differences in number
of waggle dances under the oriented or diffused light treatment (p>0.1, 0.7, and 0.2 for trials 1 to
3, respectively). They found no significant colony effects (p>0.2, 0.06, and 0.8, respectively) for
this test. However, they found a significant effect on time of day for dancing only in trial 1
(p<0.001) with more bees dancing at later hours but not in trials 2 and 3 (p>0.2 and 0.08
respectively). Finally, they found that the ratio of recruits that arrived at the feeders during
diffused light treatment decreased significantly with distance (p<0.02).
For their second phase, they found significant effects for season, period nested within
season, light treatment and the season X light treatment interactions. They found that p<0.001 for
season and period within season and p<0.005 for light treatment and p<0.05 for season X light
treatment. For the condition variation, they found that overall bees averaged greater food
collection when producing oriented dances as opposed to disoriented dances but this varied
significantly across seasons. During summer, colonies gained similar mass under either treatment
(p=0.22) and during autumn, colonies similarly lost mass regardless of treatment (p=0.85). The

only time significance is noted was during the winter months in which colonies gained mass
when exposed to oriented light and lost mass when exposed to disoriented light (p=0.0005).
Discussion:
Sherman and Visscher (2002) completed what they set out to do. They hypothesized that
the colony was a center for information exchange and communication to recruit bees to a rich
food source, in turn increasing food collection. First, they broke the hypothesis down into two
parts: communication occurs in the hive to recruit bees, and this increases food collection. In
doing so, they were able to accurately test both parts of their hypothesis. However, before even
going into testing their hypothesis, they performed preliminary experiments to assure that the
data they collected supported their hypothesis.
The preliminary testing stage was an important step that the authors completed. In this
testing, they found that diffused light produces disoriented dances and directional light produced
oriented dances. This stage is important to the overall testing because their experiments rest on
the idea that diffused light effects the dancing abilities of the bees.
From here, they went on to test the effects of disoriented versus oriented dances in the
ability to recruit bees. As expected, bees that were exposed to diffused light recruited less bees to
the feeders than bees exposed to oriented light. In other words, bees use the encoded information
in waggle dances to orient themselves to the correct feeder. Without correct information in the
waggle dance, recruiters are unable to follow the information to the feeder.
However, some recruits were still recorded at the feeding sites - raising the question as to
why this is. One aspect of bees that Sherman and Visscher did not touch on is the ability to
smell. One argument against the communicative ability of the waggle dance is the
communication by scent. Honeybees have scent glands on their bellies and are able to open and

close these glands based on the environment (Ribbands, 1955). Honeybees often times leave
their scent on flowers for one of two reasons: marking the flower as already foraged upon, or to
draw more bees to their scent presumably due to more pollen availability. This understanding of
honeybees might go to explain why the honeybees in Sherman and Visschers experiment were
less likely to find the feeder the further it got away. The scent argument, however, does not
explain why bees, under oriented light, have significantly more recruits than bees under
disorienting light - no matter the distance the feeder. Therefore, perhaps the bees are using a
mixture of the two - olfactory and waggle dance - depending on what is better in the situation as
suggested by Kirchner and Grasser (1997).
In the next experiment, seasons proved to play a large role in the foraging abilities of
honeybees. Primarily, the first response to seeing that bees increase colony size in the summer
under both light conditions is that flowers are readily available and therefore neither waggle
dances nor smell is necessary to the honeybees prosperity. Similarly, as food resources decrease
in autumn months, so does the colonies mass - no matter whether there is orienting light or
disorienting light. The most interesting and not well addressed issue is in the winter months
where honeybees increase colony mass under orienting light and decrease colony mass under
disorienting light.
The results Sherman and Visscher document for winter months are what we would expect
to see in all months where waggle dance is important to finding nectar rich sources. The issue is
that only in the winter months do we see this significant change in colony mass between light
conditions. If their hypothesis were true, we would expect to see a significant difference in any
months where resources are not quite as readily available but we do not. A baseline colony
would have been beneficial to their experiment. If they had a colony that was only exposed to

natural light or orienting light and observed for their mass changes, Sherman and Visscher would
have had something to better compare their results to. The idea they put forward was to expose
the colonies to different lights every 8 to 11 days so that each colony was ultimately under a
control group (i.e. orienting light) half the time but this switching of lights seems like it would be
pretty disorienting on its own. They did not provide sufficient argument as to why they decided
to do this and ultimately, I think it degraded from their original argument.
A way to try to figure out why they have significant results for winter but not summer or
autumn might be to perform the test in different locations. This specific test was performed in
California where bees forage profitably throughout the year which might have skewed their
results (Sherman and Visscher, 2002). Had they performed the testing in a location where there
was a noticeable drop in nectar availability during specific months, then we might see more of a
difference in colony collection rate with the differing lights. In fact, Dornhaus and Chittka (2004)
test bees foraging ability in different environmental situations and found that bees do not use the
waggle dance for communicative reasons when there is a wide abundance of resources and only
use the waggle dance when resources are clustered. The study by Dornhaus and Chittka (2004)
suggests that the results from Sherman and Visscher (2002) were indeed skewed.
An interesting addition to this study would be to see if the bees still followed these
disoriented dances at all or realized they were disoriented and did not attempt to gain information
from them. If the bees used the dances regardless of orientation, one would expect to see bees
flying to many different locations with no preference towards a specific location. Although
Sherman and Visscher might not have been interested in this, it could have gone to further
support their argument - that bees use the waggle dance for nectar collection. Also, this addition

could have shown a level of cognitive awareness if the bees did not follow disoriented dances
because it would not be just an involuntary response to waggle dances but a choice.
Overall Sherman and Visscher (2002) did an excellent job of testing their hypothesis .
With that said, they failed to address certain issues that may have either supported or refuted
their results. Similar to almost all science, more research needs to be done in order to come to a
better understanding of honey bees and how they work.
References:
Dornhaus, A., & Chittka, L. (2004). Why do honey bees dance?. Behavioral Ecology and
Sociobiology, 55(4), 395-401.
Gould, J. L. (1975). Honey bee recruitment: the dance-language controversy.Science, 189(4204),
685-693.
Kirchner, W. H., & Grasser, A. (1998). The significance of odor cues and dance language
information
for the food search behavior of honeybees (Hymenoptera: Apidae). Journal of insect
behavior, 11(2), 169-178.
Ribbands, C. R. (1955). The scent perception of the honeybee. Proceedings of the Royal
Society of London B: Biological Sciences, 143(912), 367-379.
Sherman, G., & Visscher, P. K. (2002). Honeybee colonies achieve fitness through dancing.
Nature,
419(6910), 920-922.
Von Frisch, K. (1967). The dance language and orientation of bees.