Opinion
The honeybee waggle dance: can we
follow the steps?
Christoph Grüter1 and Walter M. Farina2
1
Laboratory of Apiculture and Social Insects, Department of Biological and Environmental Science, John Maynard-Smith Building,
University of Sussex, Falmer BN1 9QG, UK
2
Grupo de Estudio de Insectos Sociales, IFIBYNE-CONICET, Departamento de Biodiversidad y Biologı́a Experimental, Facultad de
Ciencias Exactas y Naturales, Universidad de Buenos Aires, Pabellón II, Ciudad Universitaria, C1428EHA Buenos Aires, Argentina
The honeybee (Apis mellifera) waggle dance, whereby
dancing bees communicate the location of profitable
food sources to other bees in the hive, is one of the
most celebrated communication behaviours in the
animal world. Dance followers, however, often appear
to ignore this location information, the so-called dance
language, after leaving the nest. Here we consider why
foragers follow dances and discuss the function of the
dance as a multicomponent signal. We argue that the
‘dance language’ is just one information component of
the waggle dance and that the two terms should not be
used synonymously. We propose that location information is often backup information to be used in case
information acquired directly from the environment
about food sources is not available, is of poor quality
or is associated with unprofitable food sources.
Introduction
The so-called ‘dance language’ of foraging honeybees (Apis
mellifera) was first decoded over 60 years ago by Karl von
Frisch, who trained foraging honeybees to artificial food
sources and discovered that foragers could communicate
the location of distant food sources to other foragers by
means of a stereotyped behaviour [1]. Foragers perform
this figure-of-eight-shaped dance on the vertical wax combs
inside the hive [1,2] (Figure 1), conveying information to
fellow nestmates about the distance of a food source, based
on the duration of the waggle phase of the dance circuit,
and information about the direction of the food, based on
the orientation of the body of the dancer relative to gravity
[1–5]. In this way, honeybees advertise nectar, pollen and
water sources or even potential nest sites. This discovery
highlighted the sophistication of insect communication and
remains one of the most remarkable discoveries in the
study of animal behaviour.
Given that foraging bees dance only if the recently
visited food source is profitable, other foragers following
these dances should be able to find out where the richest
food sources are [2,6]. If the food source becomes depleted
or reduces in quality, foragers cease dancing and recruitment stops, thus enabling a colony to exploit the best
resources in an ever-changing environment [6]. The regulation of foraging by means of dancing has been used as an
illustrative example of self-organisation of a collective
activity [6,7].
The ‘dance language’ (see Glossary) has often been
considered key to honeybee foraging success (e.g. [6,8]).
Thus, theoretical work has focussed mainly on this location
information provided by dancers to explain the ultimate
function of dancing and dance following [9–11]. By contrast, recently published studies [12,13], as well as experiments performed during the late 1960s [14], suggest that
the importance of location information for honeybee foraging success is overestimated. Here we discuss the importance of the dance language for honeybee foraging and
explore additional reasons why foragers follow dances.
How important is location information?
Although the location information encoded in the waggle
dance can be deciphered by dance followers [4,5,15], they
often ignore this information. Here we explore why these
foragers follow dances despite then appearing to ignore the
information that the dances contain.
Glossary
Dance language:: a term used by von Frisch [2] (‘Tanzsprache’) to denote a
series of repetitive waggle movements performed by successful foragers
conveying information about the direction and distance of food sources or nest
sites (Figure 1). Here, the word ‘language’ does not imply the use of a set of
symbols and a system of grammar.
Multicomponent (or composite) signal:: a signal comprising more than one
informational component. The components can be redundant (i.e. lead to the
same or an enhanced response) or nonredundant (i.e. provide multiple
messages that lead to particular responses each); examples are given in Refs
[47,48].
Multimodal (or multisensory) signal:: a multicomponent signal that transfers
information in more than one sensory modality, for example, sound and
olfaction [47,48].
Private information:: information acquired via direct interaction with the
environment, as opposed to social information.
Scout–recruit concept:: traditionally, the terms used to characterise two
different types of forager; the differentiation can be based on different tasks
(searching for food versus collecting food), on information-use strategies
(searching independently versus using social information) or on whether bees
follow dances before leaving the hive (discussed in Ref. [49]). The two main
problems with this concept in honeybees are (i) the different definitions of the
terms used, and (ii) the limited informational content of a dichotomous
concept given the variety of decision-making strategies of foragers.
Signal:: information-bearing actions or structures that have been shaped by
natural selection specifically to convey information [6,50].
Social information:: any information acquired through the actions (e.g. waggle
dance), body structures (e.g. shapes or colours) or products (e.g. pheromones)
of other individuals.
Sugar response threshold:: this threshold is usually determined by applying
solutions of increasing sucrose concentrations to the antennae of retained
bees. When the antenna of a bee is touched with a sucrose solution of
sufficient concentration, the bee reflexively extends its proboscis (e.g. [21]).
Corresponding author: Grüter, C. (cg213@sussex.ac.uk).
242
0169-5347/$ – see front matter ß 2008 Elsevier Ltd. All rights reserved. doi:10.1016/j.tree.2008.12.007 Available online 21 March 2009
Opinion
Trends in Ecology and Evolution
Vol.24 No.5
reward, learning performance and several other behavioural traits (reviewed in Ref. [21]).
Figure 1. The honeybee waggle dance and follower bees. The dancer performs a
waggle run (1) followed by a turn to one side (2) to circle back to the starting point
of the waggle run (3), then she starts another waggle run (4) and usually turns to
the other side (5) and so on. The better the food source, the more waggle runs she
performs. During the waggle run, the dancer produces airborne sounds, airflows
and vibrations in the comb, as well as releasing chemicals [19,33,52]. Followers
touch the dancer directly with their antennae during the dance. However, the
importance of the different stimuli for location information transfer from dancer to
follower is not yet well understood. As well as providing location information,
these components might have an important psychological function for
surrounding bees. Multicomponent signalling can enhance the detection of the
signal and improve learning of the information [48].
The low efficiency of the dance language
Several researchers have noted that the dance language
appears to be less efficient at recruiting new foragers than
was originally proposed by von Frisch [4,16–18]. Theoretical and empirical works suggest that followers only need 5–
6 waggle runs to decode the location information [3,16,17].
However, many recruits that are successful in locating the
food source do so after following many more waggle runs of
several dances and subsequent search trips [16,17,19,20].
Some foragers follow >50 waggle runs without ever reaching the indicated food source [17]. Seeley [6] reported an
experiment in which dancers performed a total of 8722
waggle runs and recruited 153 bees. This corresponds to 58
waggle runs per successful recruit, indicating that the
transfer of location information is either inefficient or that
the dance language is being decoded only by a small
proportion of the bees interacting with a dancer.
It is still unclear why some recruits locate the food
source successfully after following only a few waggle runs,
whereas others follow dozens of waggle runs without being
successful. We speculate that some bees might need more
arousal and stimulation via dancing until they are motivated to start foraging, whereas others might simply be bad
learners. It would be interesting to test whether the
responsiveness of followers to rewards, such as sugar, is
related to the number of waggle runs that they follow and
the probability that they would discover the advertised
food source. In honeybees, the sugar response threshold
(see Glossary) is related to the individual evaluation of a
Olfactory information as an additional factor
Recruits can learn the odours of the collected food during
their interactions with dancing bees [2,22]. There is, however, disagreement about how important odours are for
honeybee recruitment (e.g. [14,23,24]). In an experiment
performed by Wenner et al. [23], most of the recruits were
observed at a location 140 m away from those indicated by
the dancers (a ratio of 6.8:1, respectively). Although the
indicated food sources where the dancers had been collecting food were scentless, the food source that was not
advertised contained a scent that had been collected by
foragers from the study colony in the days before the
experiment. The authors suggested that, although the
dancers successfully recruited bees to forage, recruits
relied mainly on olfactory information to locate the food,
rather than on the location information of the waggle dance
(Box 1). Lindauer [24] later repeated this experiment with
a few alterations, without finding the same exclusive effect
of odour (1:1.9 ratio of recruits at nonindicated food sources
versus indicated food sources). First, the colony in Lindauer’s experiment had less experience with the food odour
(i.e. a shorter exposure to scented food than in Ref. [23])
than was later used during the recruitment experiment;
and second, the food sources advertised by dancers were
also scented. Thus, although the dance language affected
the recruitment pattern in Lindauer’s experiment, many
bees were found at the food source that was not indicated
by the foragers but that did contain the same scent. The
discrepancy between the results of the studies has not yet
been resolved, but suggests that even small changes in
experimental design can lead to very different recruitment
patterns (discussed in Refs [14,24]). Thus, differences in
recruiting behaviour, depending on whether the food
sources are scented, require further study.
Private information as an additional factor
Active foragers rely on their own memories to find particular locations when visiting food patches repeatedly. After
an interruption in foraging (e.g. after nightfall), these
memories can be triggered by encountering the odour of
the corresponding food source inside the hive [2,14,25–27].
This so-called private location information also affects the
way in which foragers use the waggle dance.
Two recent studies suggest that, once they have left the
hive, dance followers with foraging experience (which,
during the spring and summer, includes most foragers
[28]) often do not rely on social location information
[20,28]. Biesmeijer and Seeley [28] reported that 12–25%
of all interactions with dancers accounted for attempts to
discover food. In the remaining interactions (75–88%),
dance following was observed in the context of reactivation
and confirmation of experienced foragers that might have
already known the food source that they later visited. Most
of these bees followed dances only briefly (<5 waggle runs)
[28].
Biesmeijer and Seeley [28] also assumed that bees
following <5–10 waggle runs did so for reactivation and
otherwise followed dances to discover new food sources.
243
Opinion
Box 1. How foragers can find a food source
Honeybee foragers can use many different strategies to find food.
They can rely on private information and different kinds of social
information, with and without following dances. If a forager is
initially unsuccessful in finding a food source, she could begin a
random search in the field, search for food sources with a known
odour or fly back to the nest to get more information about food
sources. Below we detail possible foraging strategies; however,
there is little information about the relative importance of these
different strategies for food source discoveries in honeybees under
natural conditions, mainly because it is difficult to track the flight
paths of most foragers.
Without following dances
Without using private information
By random search
By searching for plants with odours that were learned socially
inside the nest
Using private information initially
Inspecting a food source that has reappeared
Inspecting an empty food source, subsequently searching another
food source of known scent nearby
Inspecting an empty food source, searching randomly for another
food source nearby
Reactivation by olfaction, flying to a familiar food patch
Reactivation by olfaction to a still-empty food source, then
searching another food source of familiar scent nearby
Reactivation by olfaction to a still-empty food source, then
searching another food source of novel characteristics nearby
After following dances
Without using private information
Using location information plus olfactory information and searching the indicated food patch
Using location information plus olfactory information, but then
finding another food patch with the same scent
Using location information plus olfactory information, but then
finding another food patch by random search
Ignoring location information, searching for plants with known
scent
Ignoring location information and olfactory information, searching randomly for good food sources in the field
Using private information initially
Reactivation by dance, flying to a familiar food source
Reactivation by dance, flying to an empty food patch, then finding
another food patch with a familiar scent nearby
Reactivation by dance, flying to an empty food patch and then
finding another food patch nearby by random search
However, in a second study [20], foragers following as
many as 17 waggle runs were reactivated to resume foraging at already visited food sources. Overall, 93% of all
foragers with private information about the location of a
good food source ignored the dance language [20]. This
suggests that the importance of the waggle dance for
reactivation under natural circumstances was underestimated in Ref. [28]. The waggle dance often triggers navigational memories about food sources known to the dance
follower, irrespective of the location indicated by the dancer [20]. This could also partly explain why the dance
language appears to be inefficient, in that many foragers
do not find the indicated food source after following a dance
[4,16–18]. In studies of dance efficiency, it is often unclear
whether ‘unsuccessful’ recruits made any attempt to fly to
the indicated location or, instead, inspected known food
sources.
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Trends in Ecology and Evolution Vol.24 No.5
Little effect of waggle orientation on overall colony
foraging success
When looking at the overall importance of location information for colony foraging success, two studies found that
there is often no positive effect [12,13]. These studies
compared the foraging success of colonies having correctly
dancing bees with colonies having only disoriented dances,
created by tilting the combs into a horizontal position.
Unless the bees have a direct view of the sun or a polarised
skylight, they perform dances in random direction, because
they lose gravity as a reference. The studies found that, in
temperate habitats, there was no difference between colonies during the spring and summer, when most nectar and
pollen are harvested. The dance language did have a
positive effect, however, when food sources were scarce
[10,12] or appeared clumped, as in some tropical habitats
[13]. Yet, even relatively small benefits of location information could be important for colony survival in such
ecological situations.
In summary, studies suggest that the spatial information encoded in the dance affects the behaviour of
individual bees and the overall foraging success of a colony
only under certain circumstances. However, as we discuss
below, the waggle dance does more than just provide
spatial information.
The waggle dance: a multicomponent signal
The waggle dance is a multicomponent signal in that it (i)
attracts surrounding bees so that they can receive other
types of information; (ii) informs bees of the presence of
good food sources; (iii) activates private navigational information (if present) in followers; (iv) facilitates the acquisition of information about food odours; and (v) indicates
the location of the food source. To understand how the
waggle dance affects collective foraging patterns and,
therefore, what its ultimate function might be, the behavioural response to the different components of this multicomponent signal should be considered.
Dancing modulates the readiness to respond to
information
Before von Frisch discovered the location coding of the
waggle dance, he reported that foragers could be recruited
to food sources by what he called the ‘round’ dance [25],
which is performed by foragers returning from nearby food
sources [2,25] and is now considered to be a special version
of the waggle dance [29,30]. Bees following round dances
leave the hive in a random direction and search nearby for
food sources [2]. von Frisch reported that dancing attracts
the attention of bees in the immediate vicinity of the
dancer [25]; thus, the dancer primes the readiness of
surrounding bees to respond to other components of the
multicomponent signal.
Chemosensory information
During their interaction with the dancer, followers can
learn the odour of the food source [2,22,25], which helps
them to locate other food sources with the same odour
[2,14] (see also Box 2). Followers position themselves
around dancing bees in ways that optimise olfactory
information acquisition, for example, they preferentially
Opinion
Box 2. The evolutionary origin of the waggle dance
Comparisons of recruitment systems of different social bee species
have led to the suggestion that successful ancestral foragers
showed some kind of excitatory movements (e.g. zig-zag runs,
shaking movements, running into other foragers and/or produced
buzzing noises) [51–53] that attracted the attention of other
nestmates and made them leave the nest in search of food. Such
recruitment behaviours can still be observed in bumblebees,
stingless bees and ants [54–56]. In addition, foragers would have
learned the odour of the food source in the nest during food
offerings that were performed during interruptions in the excitatory
movements. Thereby, these movements might have facilitated the
acquisition of olfactory information. Such nest-based food odour
learning can still be observed in stingless bees, bumblebees, ants
and wasps [54,57–59].
These excitatory behaviours are assumed to be the origin of a
dance that conveyed information about the location of the food (e.g.
[51]). Interestingly, it has been suggested that stingless bee
Melipona panamica foragers transmit spatial information to other
foragers by sounds inside the nest [53]. However, the intermediate
stages toward a dance that provided the direction and distance to a
food source are still poorly understood [52]. There is now growing
consensus that selection initially favoured the transfer of location
information about new nest sites, rather than about food sources
(e.g. [54,60]).
contact body parts where food odours are most intensive
like the mouthparts of nectar foragers or the hind legs of
pollen foragers [31]. Small nectar offerings from dancers to
surrounding bees during interruptions in dancing also
appear to have an important role as reinforcement for
olfactory conditioning [2,22,32].
Additionally, waggle-dancing bees release chemicals
(two alkanes and two alkenes of long carbon-chain length),
which increase the propensity of bees to leave the hive [33].
The effect of these chemicals was confirmed by blowing a
volatile mixture of the chemicals on the comb surface,
where bees usually perform dances. This could partly
explain von Frisch’s observation that many bees following
dances subsequently show an elevated probability to leave
the hive and search for food [25]. In this way, the dance
informs other bees that profitable food sources can be found
in the surroundings of the hive. The fact that artificially
injecting these chemicals into the hive encourages bees to
leave the hive suggests that foragers might be recruited by
dances without ever physically contacting a dancer (Box 1).
It might be that bees acquire information about the
amount of dancing and, therefore, the foraging conditions,
by the amount of these chemicals that they perceive on the
hive floor [33]. However, this needs to be tested experimentally. If it is the case, then we would expect a positive
relationship between the concentration of the chemicals on
the hive floor or in the hive atmosphere and the number of
foragers leaving the hive.
What kind of information do dance followers use?
Researchers tend to interpret dance following as an
attempt of the follower bee to decode spatial information
about a food source. However, there are other reasons why
dances are followed [20,28]. For example, the dance often
functions as a means of reactivating inactive foragers to fly
to food patches that they have already visited, but does not
necessarily have to indicate a location or plant species that
Trends in Ecology and Evolution
Vol.24 No.5
is already known to the follower [20,34]. Hence, some
inactive foragers seem to use the dance as an indication
of an increased chance of finding food at places where food
has been found previously without decoding the location
information of the dance. By contrast, other bees appear to
rely strongly on the food odour information gained during
dance following, when they search for food in the field [23].
Because dance followers are often foragers that are
already engaged in foraging, it has been suggested that
active foragers follow dances to confirm the continued
availability of the type of food that they had been collecting
[28]. Most of these bees follow only a few waggles (<5
waggle runs) and, therefore, are not interested in detailed
location information [28]. This kind of dance following was
found to be particularly frequent in bees that had had an
unsuccessful foraging trip immediately before dance following [28]. However, in such a situation, it might be
beneficial to follow more waggle runs and decode the
location instruction of the dance because the advertised
food patch is apparently profitable. Experiments are
needed to test whether dance following of active foragers
does serve to confirm the availability of particular food
types.
Leaving the hive without following dances
Foragers sometimes leave the hive without having followed dances (e.g. [28,35–37]). Such bees are often called
‘scouts,’ and it is assumed that they do not use acquired
information. However, under natural conditions, bees leaving without having followed a dancer could (i) use private
information after reactivation (e.g. via chemosensory cues;
Box 1); (ii) use private information and inspect previously
visited food sources [2]; (iii) use social information (e.g.
chemosensory information acquired previously [38], or in
the present [33]); or (iv) use neither private nor social
information and rely on innate search behaviours. To
separate the strategies, information is required about
the foraging experiences of a bee, about whether she
was reactivated and, most importantly, what she does
after leaving the hive. However, to our knowledge, little
is known about the relative frequency and importance of
these strategies under natural conditions.
This array of behavioural strategies of bees either following dances or leaving the hive without doing so (Box 1)
makes the classic dichotomous scout–recruit concept an
arbitrary classification of little informative value. Considering more strategies of information use (Box 1) instead
will lead to a better understanding of collective foraging in
honeybees.
Private versus social information
Many plant species only bloom (and offer nectar or pollen)
at limited periods during the day [2,39]. Honeybees can
learn these times of food abundance after a few days [40]
and often specialise to collect one or two different food
types at particular times of the day at particular locations
[41,42]. A foraging honeybee will return to the same flower
patch at the same time period for many days (even her
entire life) if the patch remains rewarding [42].
Private information is especially important for the
organisation of collective foraging in times of good foraging
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Opinion
conditions when food patches offer food for several days.
With increasing foraging experience and, therefore, more
precise private information about the location and flowering time of food sources, foragers show a decreasing interest in following dances [28,36].
Social information use is often assumed to be adaptive
because the costs of individual exploration can be avoided
[43]. In changing or heterogeneous environments, however, erroneous or outdated information could be obtained
by relying on others. Hence, social information use is not
always an adaptive strategy [43,44]. There is evidence from
various taxonomic groups, including birds, fish, insects and
mammals, that private information is used if it is available
and reliable, whereas social information is used as a
backup if private information has proven to be unreliable
[43,45,46] or outdated [45] or if individuals evaluate this
option as unprofitable [43]. Honeybees also appear to rely
on private rather than social location information if the
former is both available and reliable [20,28,36].
The impression that dance followers often use social
location information could be a consequence of the fact that
experiments investigating the waggle dance are normally
performed after the end of the flowering season or at places
where there are few alternative food sources because it is
otherwise difficult to train bees to artificial feeders [6].
Under these circumstances, one would expect private information to be outdated or not available and that foragers
would rely more on social location information. To explore
the use of private and social information in honeybees
further, controlled feeder experiments should experimentally manipulate the reliability and outdatedness of private and social information, as well as the costs and
benefits associated with using information.
Conclusions and future directions
It has been convincingly shown that honeybees following a
waggle dance can decipher the spatial information encoded
by the dance [4,5,15]. However, followers often do not use
this information. Experienced foragers usually rely on
private information about food source locations after following a waggle dance, whereas other foragers appear to
rely mainly on food odours learned during dance following
to find food sources with the same odour [23]. This shows
that the dance language is only one component of the
waggle dance and that the two terms should no longer
be used synonymously.
For a better understanding of the adaptive value of the
waggle dance, various questions still need to be addressed,
including: why do engaged foragers frequently follow
dances? Which circumstances favour the use of social
location information versus private location information
in experienced foragers? What explains the large variation
between individuals with respect to dance-following behaviour? Future theoretical and experimental work should
explore communication and recruitment in honeybees as a
more complex process where multiple kinds of information
are available and can act together. Harmonic radar technology, which enables one to follow the flight paths of bees
over distances of hundreds of meters, might help us to
understand how common different foraging strategies are
(e.g. [5]). Adopting an approach that does not reduce the
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Trends in Ecology and Evolution Vol.24 No.5
waggle dance to a mechanism to communicate spatial
information but also considers alternative strategies of
information use (Box 1) will help us understand why bees
follow its steps.
Acknowledgements
We thank Heikki Helanterä, Francis Ratnieks, Roger Schürch and
Francisca Segers for comments on earlier versions of the article, and
Nadja Stadelmann from Descience for Figure 1. C.G. was supported by a
postdoctoral fellowship from the Swiss National Science Foundation
(grant PBBEP3–123648). W.M.F. was supported by the University of
Buenos Aires (grant X 077), the National Agency for Scientific and
Technological Promotion (grant PICT 2006–1155) and the National
Council for Scientific and Technological Research from Argentina.
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Forthcoming Conferences
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5–10 July 2009
International Congress of Systematic and Evolutionary Biology (ICSEB VII), Veracruz, Mexico
http://www.botanik.univie.ac.at/ICSEB7/
11–16 July 2009
International Congress for Conservation Biology; 23rd Annual Meeting of the Society of
Conservation Biology 2009, Beijing, China
http://scb2009.ioz.ac.cn/index.asp?CFID=10602810&CFTOKEN=24167527
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28–31 July 2009
First International Entomophagous Insects Conference, Minneapolis, MN, USA
http://www.cce.umn.edu/conferences/entomophagous/index.html
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