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A Honey Bee’s Waggle Dance and its Importance to the Colony
Mayleen Brown
Insect Behavior April 4, 2007
Abstract
Honeybees use the art of dancing to communicate the location, in direction and distance, of a food source to their nest mates. This discovery is the most sophisticated
example of non-primate communication that we know of. These dances are called the waggle dances and they depend on many different triggers. Properties of the actual nectar source are one trigger and stimuli from the foraging status of the colony, as a whole, is another trigger needed for a proper dance. A returning forager can sense the foraging status by how well the food transfer process goes. If the food receivers unload the crop swiftly and excitedly, then the forager will most likely dance because the colony’s nectar intake rate is low. The extent to which the worker bees obtain information from the waggle dance is very important. A majority of the new foraging honeybees rely on information obtained from a foragers waggle dance instead of actually just searching themselves. The waggle dance is also useful to the practiced foragers who use it for reactivation and confirmation of the food source. Therefore, the dance is useful to all foragers of the colony. Another point to this waggle dance is that a dance site is formed in the nest for the transformation of information. They are labeled in some way to encourage the congregation of dancers and dance receivers. It is very important for dance receivers to have the best opportunity to obtain information because they have to be positioned in certain ways with the forager to be able to get the maximum effect of the 2
dance. The follower bee has to stay behind the dancer at an angle for at least one dance
to perceive the maximum information needed. Time patterns and air flow from the
vibration of the dancer’s wings are the reason for this specific positioning. The waggle dance is a chief key to the honeybees and their ability to be major producers.
Introduction
The species Apis Mellifera, commonly known as the honeybee, has a simple but
remarkable way of communicating to its nest mates the specific location of food sources.
This profound discovery started with the work of Karl von Frisch who one the Nobel
Prize for the findings, experiments, and comprehension of the behavior. The honeybees
use the “waggle dance”, which is a dance that the foragers of the nest perform to pass on
information about the presence of desirable nectar sources. It is one of the most
outstanding forms of animal communication. Hundreds of studies have been conducted
to understand the precise techniques, mechanisms, ecological significance, and
evolutionary origins of this dance. (Biesmeijer and Seeley 2005) The dance is calculated
to illustrate the direction and distance of a nectar source that is a certain distance from the
hive. The main message of the waggle dance is that more foragers are needed at a
valuable and specific food location. (Michelsen 2003) The dance contributes to the
success of a honey bee colony when food sources are patchy and difficult to locate. The
way this dance works is quite straightforward. The bees use the vertical alignment of the
actual nest and the angle, in degrees, that the food source is from the sun, to calculate and
perform their dance. Then, rotating the vertical alignment of the nest to point to the sun
completes the direction by positioning the other part of the angle directly at the food 3
source. Bees used to dance in the exact direction of the food source when their hive was open. Now, they use the angles in relation to the sun and food because of the closed hive.
Components of the Dance
The waggle dance of the honeybees has a few different factors to its performance.
The purpose of the dance is to convey the presence of food sources to other forager bees
in a colony. One factor behind the regulation of the dance is the properties of the actual nectar source, like distance and direction. The other factor involves properties of the colony. There are certain stimuli that result from the foraging status of the entire colony.
Bees tune their dancing according to their colony’s nectar influx. For instance, if foragers are using a source that has steady profitability, they will mostly dance when the nectar intake rate is low. The method of discovering what the condition of the nectar intake rate is involves trophallaxis, which is the transfer of liquid food by mouth. This is a general behavior in social insects that is especially important here. The foragers of the colony transfer their crops to food-receivers when they return to the nest and this is where trophallaxis occurs. A forager may sense the eagerness and speed of the food-receivers to unload the crop, which then communicates the status of the nectar influx. (De Marco
2005) Trophallaxis is a very important interaction between dancer bees and workers because it is the transfer of not only food but of information about the colony. This exchange of food can change very quickly depending upon the food source profitability.
The numbers of trophallaxis offering contacts are increased when profitability is increased, but the length of these contacts is decreased showing a more hurried approach
to get as much food as possible while the probability is high. Also, when the probability
is high, these exchanges are performed more evenly throughout the visit of the dancers 4
instead of at the beginning of their arrival. Returning foragers also beg for food less
often and only do it before their departure, with high probability. This shows that the
bees can understand their colony’s need of food and the dancers will change their
routines to benefit the colony as a whole. The dancers perform more waggle runs and dance earlier in their visit when the food source profitability is increased, as to acquire the help of as many foragers as possible. (De Marco and Farina 2001) Also, discovered by an experiment on this specific topic, foragers can swiftly sense variations in the influx when there is no change in the time interval between their voyage return and the beginning of food transfer. This is because they use stimuli derived from the number of food-receivers. This causes them to set their dance thresholds in relation to the colony’s nectar influx. The probability of dance changes only after the foragers’ food-unloading.
The lower the colony’s sugar solution intake rate is the higher the dance probability is.
(De Marco 2005)
The extent that honeybees acquire information from the waggle dance throughout their foraging careers is very diverse. The dance that is performed in the colony is a useful tool for all ages of foragers. It would normally be thought that the waggle dance is mainly used to help young and first time foragers to obtain information about their first food source, but in actuality, it is used for all foragers including the most experienced.
The waggle dance is utilized to start work at new and foreign food sources, to reactivate a food location after the interruption of foraging, and to confirm that the food location is still yielding forage. Dancing is followed by the experienced much more often after an unsuccessful foraging trip than after a successful one. An experiment was done to support these findings and it further explains the results. For first time foragers in a 5
colony, they relied on the waggle dance, the majority of the time, to obtain information
on food sources. They were recruited rather than scouting and searching on their own.
With a good percentage of experienced foragers who have been interrupted, they used the
waggle dance for reactivation instead of examining the food source themselves. For an
experienced forager that is still engaged, they would follow dances just briefly for
confirmation. (Biesmeijer and Seeley 2005)
Now, there is a differentiation in the dances that these honeybees do. Duration is
not the only factor in how the dances can be set apart from one another. Karl von Frisch came up with the theory that when foragers found a profitable food patch within a 20 meter radius from the hive, they would perform round dances with no given distance information. When studied further, it was found that distance information is given during waggle phases which have no direction orientation up to 15 meters in distance. At 80 meters though, dancers execute a true waggle dance in which the direction of the waggle indicates the angle of direction of the food source. The waggle dances steadily increase with longer distances. (Sarma et al. 2003)
A waggle dance also depends on how substantial the food is at the food source
location. When feeders that were created with different concentrations of sugar were
used in an experiment, they brought about very understandable conclusions. If a feeder
had a low concentration of sugar, then every step in the forager’s duties is changed and
performed differently. Foragers do not dance in the hives when there is a low
concentration of sugar at the feeder. Their flights to the feeder are often carried out alone
instead of with the usual many recruits from the dance. Also, instead of circling around
the feeder they land immediately and no recruits follow or land. When the sugar 6
concentration is raised though, it changes the performance in every aspect. Strong
waggle dances are performed and the foragers and recruits fly together in groups and circle the feeder before landing around the same time. Or, the recruits will land immediately after the forager because they have never done it before. Recruits will circle around a feeder if it is complemented by the release of geraniol scent, a colorless scent found in many oils of prominent flowers. It is proposed that recruits who have witnessed a waggle dance are not likely to find a non-scented feeder unless the foragers provide visual cues like circling around the feeder. This is a strategy for the recruits to overcome gaps in the process. For example a dance showing only a general area and not an exact position would lower the probability of some recruits ever making it to the feeder. (Tautz and Sandeman 2003)
The duration of a waggle dance can be summed up to have many components.
The overall quality of a food source is what increases the duration of the dance. The
“quality” is defined by factors like the distance to the target, the conditions of the weather, and the season of the year. The repetitions of the dance sound are also increased in accordance with quality. Dancers are more “lively” when the quality of a food source is high. This liveliness can be measured by the time interval between repeated waggles.
Motivated bees are quicker to return to the hive, as if excited to pass on the information.
Bees who are less motivated tend to lollygag and may even perform silent dances as not to recruit others. (Michelsen 2003)
Importance for Honeybees and Humans
The waggle dance of honeybees is very useful for the interpretation of experimental results involving other aspects of bee’s structure. For instance the operation 7
of their visual odometer can be understood because of the duration of the waggle dance
of experimental bees. Their dances are equivalent only to the distance traveled to a feeding location, which is the amount the environment moves in their eyes, concluding that the visual odometer is independent of a third dimension. So, by comparing different
waggle lengths and different feeding distances of many diverse configurations,
experimenters could draw this conclusion. (Dacke and Srinivasan 2007)
The waggle dance is also very important to the honeybees because it is the connecting piece to their orientation flights. Orientation flights are flights performed by the foragers of the colony that help honeybees to obtain information about the surrounding terrain of their hives. These flights contain all of the flights performed by the bee before it makes its first flight to and from a known forage location. These orientation flights are successful to contributing to the knowledge of the bee’s surroundings because honeybees exchange the obtained information on the location of forage patches through their waggle dance. They perform the dance to other foragers of the colony. (Reynolds 2006)
Further more Complex Innovations
The transfer of information from the dancer to the follower bee is very important and rather complicated in some ways. There is a lot of flexibility in the dances. The dances are a combination of touch and the perception of air flows. Follower bees must stay behind the dancer within the angle of wagging for at least one full waggle run to obtain the correct information. The antenna of the follower bee is the air-flow receiver and it experiences the flow when the dancer’s abdomen comes near. The vibrations of the wings produce air flows alternating with the vibration frequency of the wings and 8
may also create jet air flows. The jet air flows are very useful in that they help to align
the body of the follower-bee to match the dancer. They are also signals for the direction
to the target. The time spent in between consecutive waggle runs depends upon the motivation of the dancer bee. This means that they can switch on and off the narrow and broad jet air flows which gives them a lot of flexibility. Patterns of the dances that follow
have flexibility, which relies on the presence of other targets. The patterns performed by
the follower bee when moving with the dancer also show a lot of flexibility. (Michelsen
2003) This shows how many intricate devices there are involving the transfer of information which is the basis of this behavior.
It seems that genetics have a place among the variability of the waggle dance as
well. A colony that has many subfamilies can increase its efficiency in the dance
language when members of a specific family understand and realize the dance instead of random workers. An experiment done with genetic markers per family show that there is a component associated with the dance language that is linked to genetic makeup. There are a couple of specific subfamilies that perform most of the waggle dances which shows that some families have nothing to do with foraging and probably perform other duties.
The subfamilies all must have different thresholds for dancing and it is these thresholds that motivate the dance of a certain forager after its return. The threshold is the level of nectar source profitability. Therefore, the waggle dance is a tool in suggesting that subfamilies of one colony differ from other subfamilies in their performance of tasks because of different distributions of response thresholds for task stimuli. (Arnold et al.
2002) 9
Honeybees use the combination of their dance and the construction of the actual honeycomb to attract their dance-followers when they are at a distance too far away for visual signals to play a role. The honeycomb has a unique property that helps dance- followers to find this dance. When the vibration of a dancer bee is around 200 hertz, the honeycomb represents an amplification of the vibrations as a signal to the followers. The walls of a single comb reverse their displacement and move in opposite directions to one another which cause the message to be sent. The behaviors of the dance-followers show that their position when recruited corresponds to the location of the reversal of the comb movement relative to the dancing bee. This amplification arises when a bee straddles a cell and its legs move more towards and away from one another instead of in their usual same direction. This is the indication to a dance-follower that a dance is being performed in the area. (Tautz, J. et al. 2003) Not only can honeybees sense the dance from a distance, but they can become dance followers very easily depending on a few different factors. They can become a follower after touching the dancer or touching another dance follower with its antenna. Without antennal contact, the strength of the attraction depends entirely on the floor, on which the dancer is dancing, on whether the follower is on the same substrata as the dancer, the direction of the follower relative to the dancing bee, the size of the dance group, and finally the light conditions over the dancer. The floor plays a big part because it depends on whether the cells are open or capped. As already noted above, the opened cells help with the traveling vibrations. A lateral direction to the dancer bee works best for communication and attraction for the follower bees. Also, infrared light is not as appealing as the illumination of artificial visible light. 10
These factors are the determinants of how well follower bees obtain information and actually become followers. (Tautz and Rohrseitz 1998)
Conclusion
The waggle dance has become a discovery of many dimensions in which it helps not only the honeybees but the environment and human beings. If it was not for the waggle dance and the expert communication, it would take honeybees much longer to discover food which would mean that their hive would always be weak. A weak hive and overworked honeybees would not have made it this far in evolution, it is there strength that ability to work with each other that has brought them this far. The waggle dance, like shown in one of the experiments, helps us to understand other behaviors of the honeybee which broadens our education and movement towards more discoveries. Also, honeybees are a key pollinator for almost all of the crops that human beings and other animals eat. If it were not for the success of their waggle dance, which makes them so dynamic, our world would not work in the way that it does today.
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References
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