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In recent years, plastic comb foundation has become widely used by beekeepers but it has not been studied to see if it hinders recruitment communication by reducing the transmission of the comb vibrations produced by bees performing waggle dances. We used laser vibrometry to compare combs built with beeswax foundation vs. plastic foundation in terms of transmission of dance vibrations. We also used behavioral experiments to compare the recruitment effectiveness of dances performed on combs built with beeswax foundation vs. plastic foundation. We found that combs built with plastic foundation are markedly poorer at transmitting the 250 Hz vibrations produced by dancing bees. Nevertheless, we found no evidence of reduced effectiveness of dances performed on combs built with plastic foundation vs. combs built with beeswax foundation. Evidently, a comb built with plastic foundation provides a fully suitable substrate for waggle dance communication.
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Apidologie 36 (2005) 513–521
© INRA/DIB-AGIB/ EDP Sciences, 2005
DOI: 10.1051/apido:2005037
Original article
Does plastic comb foundation hinder waggle dance
Thomas D. SEELEYa*, Adrian M. REICHa, Jürgen TAUTZb
a Department of Neurobiology and Behavior, Cornell University, Ithaca, NY 14853, USA
b bee group, Biocenter, Universität Würzburg, Am Hubland, 97074 Würzburg, Germany
Received 8 February 2005 – revised 16 March 2005 – accepted 16 March 2005
Published online 13 September 2005
Abstract In recent years, plastic comb foundation has become widely used by beekeepers but it has not
been studied to see if it hinders recruitment communication by reducing the transmission of the comb
vibrations produced by bees performing waggle dances. We used laser vibrometry to compare combs built
with beeswax foundation vs. plastic foundation in terms of transmission of dance vibrations. We also used
behavioral experiments to compare the recruitment effectiveness of dances performed on combs built with
beeswax foundation vs. plastic foundation. We found that combs built with plastic foundation are markedly
poorer at transmitting the 250 Hz vibrations produced by dancing bees. Nevertheless, we found no evidence
of reduced effectiveness of dances performed on combs built with plastic foundation vs. combs built with
beeswax foundation. Evidently, a comb built with plastic foundation provides a fully suitable substrate for
waggle dance communication.
Apis mellifera / comb vibration / plastic foundation / recruitment communication / waggle dance
Modern beekeeping is based on four key
inventions from the 1800s: movable frame
hive, bellows bee smoker, honey extractor, and
comb foundation (Crane, 1990). Comb foun-
dation – thin sheets of material embossed with
the hexagonal pattern of worker cells on which
the bees build their combs – ensures that the
bees build planar combs, saves them much wax
synthesis during comb building, and inhibits
them from rearing drones, by doing away with
most of a colony’s drone comb. Over the past
20 years, the technology of comb foundation
has changed markedly, expanding the types of
foundation available to beekeepers. Previ-
ously, comb foundation consisted simply of
sheets of beeswax that were fitted to wooden
frames. Now beekeepers have the options of
plastic foundation fitted to wooden frames, and
one-piece plastic frames and foundation. Under
favorable conditions, bees will build comb on
plastic foundation, and many beekeepers have
adopted this technology. To date, however, no
one has investigated whether frames of comb
built with plastic foundation, which is less flex-
ible and more massive than beeswax founda-
tion, provide the bees with a fully suitable
substrate for waggle dance communication. If
not, then such combs may have negative effects
on a colony’s recruitment communication and
on its honey production. The aim of this study
was to see if plastic comb foundation hinders
waggle dance communication.
Bees recruit nest mates to rich food sources
by performing waggle dances. Each circuit of
* Corresponding author:
1Manuscript editor: Stan Schneider
Article published by EDP Sciences and available at
514 T.D. Seeley et al.
the dance contains a waggle run during which
the bee makes lateral body vibrations at
approximately 15 Hz and thorax/wing vibra-
tions at approximately 250 Hz (Esch, 1961;
Wenner, 1962; Michelsen et al., 1986). The
thorax/wing vibrations generate airborne
sounds that dance followers detect when adja-
cent to a waggle dancer (Michelsen, 1993,
2003). Several recent studies indicate that the
vibrations produced by a dancing bee are also
transmitted via the comb and help followers
locate dancers. (1) Dances performed on open
empty cells recruit twice as many bees per wag-
gle run compared to dances performed on
capped brood cells (Tautz, 1996). (2) The dis-
tance from which a dancer attracts followers is
greater if the dance occurs on open empty cells
rather than capped brood cells (Tautz and
Rohrseitz, 1998). (3) Vibrations applied to the
rims of open cells are transmitted across the
comb and are even amplified through reso-
nance if their frequency is approximately 20 Hz
or 250 Hz (Sandeman et al., 1996). (4) The
200–300 Hz comb vibrations produced during
waggle dances have amplitudes above the
detection threshold of the subgenual organ
(Kilpinen and Storm, 1997; Nieh and Tautz,
2000). These findings leave little doubt that
dancing bees rely, at least in part, on substrate-
borne vibrations in their recruitment commu-
nication. They also suggest that if the transmis-
sion of these comb-borne vibrations is hindered,
then recruitment is weakened.
In this study, we used laser vibrometry to
investigate whether combs built with plastic
foundation, relative to combs built with
beeswax foundation, are inferior at transmit-
ting 250 Hz vibrations. In addition, we used
behavioral measurements to investigate whether
waggle dances produced on combs built with
plastic foundation, relative to those produced
on combs built with beeswax foundation, are
less effective in recruiting bees to a food
2.1. Laser vibrometry study
in the laboratory: April 2003
We used 3 frames that were fitted with foundation
and then filled (by bees) with beeswax comb:
(1) BW: beeswax foundation in a wooden frame,
(2) PW: plastic foundation (Snap-in Pierco) in a
wooden frame, and (3) PP: plastic foundation built
into a plastic frame (Pierco 1-Piece Frame/Founda-
tion unit). All were full-depth Langstroth frames. In
June 2002, the three frames were put in one strong,
comb-building colony. It completely filled the
frames with beeswax comb and used them for honey
storage. In mid September 2002, the honey was
extracted from the three frames of comb and in early
October, when no nectar was being collected, the
combs were returned to the comb-builder colony,
which cleaned and repaired them. In mid October,
the combs, empty of honey and perfect in form, were
retrieved for study.
When taking measurements, we fastened each
frame horizontally to a vibration-free table with a C-
clamp at each end. The bees built the cells in each
comb such that two walls of each hexagonal cell
were vertical when the frame was vertical. We dis-
placed the rims of vertical cell walls from the side
and in the plane of the comb face, thereby approxi-
mating the vibration pattern produced by a bee mak-
ing vertically-oriented waggle runs on the comb.
Displacements were made with a bilayered piezo-
electric transducer (BM/ML 60/40/300, Piezo-
mechanik, München) generating 250 Hz vibrations.
Further technical details on the methods of loading
vibrations into the combs are given in Sandeman
et al. (1996).
The amplitude of cell rim displacement was
measured with a laser vibrometer, first at the rim
where the displacements were applied (input point)
and then at a rim, four cells away (output point, see
Fig. 1). Small reflective flags (0.5 × 0.5 mm) were
attached to the cell rims being measured. Input dis-
placement amplitude was set at 1 µm and the result-
ing output four cells away was measured. Measure-
ments were made only after the signal from the laser
vibrometer had stabilized for 30 s. Each measure-
ment was repeated 5 times.
Each frame of comb contained 45 rows of
cells. Measurements were made on cells in rows 2,
6, 10, …, 22 using cells one quarter of the horizontal
distance across the comb (see Fig. 1). Thus the six
measurement points were spread over the comb
region that corresponds to the “dance floor” (see
Fig. 5.4 in Seeley, 1995). A set of measurements was
made for each comb when the comb was 100%, 50%,
25%, and 0% attached to the frame along the sides
and bottom. Each comb started out 100% attached
to its frame, and was detached to increasing degrees
by melting away 4-mm strips of comb (see Fig. 1).
Even when 100% detached along the sides and bot-
tom of the frame, each comb remained fully attached
along the top.
Plastic foundation and recruitment communication 515
2.2. Behavioral study in the field: July
We set up an observation hive, trained 10 bees to
a feeder 350 m from the hive, and then video
recorded the dances of these bees and measured their
recruitment of other bees to the feeder. Data were
collected three times per day, each time with a dif-
ferent frame of comb as the dance substrate. We then
reviewed the video recordings to count the number
of waggle runs produced on each type of comb.
Using these waggle run counts and the recruit
counts, we calculated the waggle dance effective-
ness (measured as recruits per waggle run) for the
three types of dance substrate.
This experiment utilized six frames of comb, two
of each of the three types already mentioned, and
prepared as described above. During data collection,
one comb at a time was placed in the lower position
in a two-frame observation hive which contained
one frame of brood comb (upper) and one frame of
storage comb (lower) (shown in Seeley, 1995,
pp. 71–73). This hive housed a colony consisting of
approximately 4000 worker bees and one queen bee.
A sheet of queen excluder material between the two
frames prevented the queen from laying eggs in the
lower comb. A wedge just inside the hive’s entrance
forced all foragers to enter and exit the hive from one
side of the hive. Because returning foragers perform
their waggle dances shortly after entering the hive,
this wedge established a “dance floor” on one side
of the lower comb.
The observation hive colony was established at
the Cranberry Lake Biological Station (44°09’N,
74°48’W), in the Adirondack State Park, in northern
New York State. This study site is surrounded by for-
est and lakes, hence there are few natural food
sources for bees, which makes it easy to train bees
to forage at a feeder. The hive was mounted in a small
hut (see Fig. 4.3 in Seeley, 1995) and the dances were
recorded with a camera (Panasonic WV-F250B)
whose field of view covered the dance floor. Vide-
otapes were analyzed using a variable-speed video-
deck (JVC BR-S525U). Dances were illuminated by
the daylight that entered the hut through its translu-
cent roof. We used a feeder that provided scented
sucrose solution ad libitum, as described in Seeley
(1995). The sucrose solution was generally a
1.50 mol/L solution, but at times it was adjusted
higher or lower to get the desired level of dancing.
We collected data on 6 days (see Tab. I), using
one set of combs (BW, PW, and PP) on the first
Table I. The arrangement of the 6 trials of the experiment, with the two sets of combs. BW = beeswax
foundation in wooden frame. PW = plastic foundation in wooden frame. PP = plastic foundation in plastic
frame (“one-piece frame/foundation”).
set Date Time
Tes t
Number of
waggle runs
Number of
1 10 July 04 11:20–14:00 BW-PW-PP 2287 107
1 11 July 04 11:00–13:40 PW-PP-BW 2999 141
1 17 July 04 11:20–14:00 PP-BW-PW 3437 139
2 20 July 04 11:20–14:00 BW-PW-PP 663 24
2 23 July 04 11:00–13:40 PW-PP-BW 1714 69
2 24 July 04 11:40–14:20 PP-BW-PW 2261 111
Figure 1. Diagram of a test comb,
showing the locations of the vibration
input and output measurements, and
the way the comb was detached in three
stages from the sides and bottom of
the frame. The double-headed arrow
between the measurement points indi-
cates the displacement direction for the
input vibrations.
516 T.D. Seeley et al.
3 days and the other set on the second three days. On
each day of data collection, we followed a standard
protocol. We started with the feeder empty and a
standard comb (not a test comb) in the lower position
of the observation hive. At 9:00, we filled the feeder
and soon bees began visiting it and recruiting nest-
mates. (Previously, we had trained 10 labeled bees
to the feeder. Recruits to the feeder were easily
detected as unlabeled bees.) Once recruits began
arriving, we began capturing them in zip-lock
freezer bags, keeping tallies of the number of recruits
captured per 10 min period. This capturing and tal-
lying of recruits continued without interruption until
we ended the experiment in the afternoon. As soon
as our 10 labeled bees were regularly producing
dances for the feeder (by 11:00 or so), we took the
lower frame out of the observation and replaced it
with the first of the 3 test combs. We then shook the
bees off the removed frame onto a board mounted at
the hive entrance, whereupon they crawled back into
the hive. Next, we gave the colony 20 min to recover
and then we started video recording the dances of the
10 labeled bees. We continued this recording for
40 min, all the while capturing recruits at the feeder.
When the 40-min period was over, we repeated the
comb swapping procedure described above, replac-
ing test comb 1 with test comb 2, gave the colony a
20 min period for recovery, and then resumed video
recording the dances, now on a different substrate.
After 40 min of video recording for test comb 2, the
entire procedure was repeated for test comb 3. When
this was finished, we replaced test comb 3 with the
standard comb, and shut off the feeder to end the
experiment for the day. As is shown in Table I, this
protocol was performed on 3 days for each of the two
sets of test combs, but on each day we presented the
test combs in a different order, to control for possible
order effects. When we shifted to the second set of
combs, we switched to a new set of 10 recruiter bees.
Also, part way through the experiment, on 21 July,
we added approximately 1500 bees to the observa-
tion hive to restore its strength.
On each day, and for each of the three frames of
comb, we obtained 4 values of dance effectiveness,
one for each 10-min block of the 40 min of data col-
lection for each type of comb. We used the average
of these 4 values as our estimate of dance effective-
ness for a particular type of comb on a given day. We
tested the three frames of comb in each comb set on
three days, using a different presentation order each
day (see Tab. I), hence we used a complete block
design to look for effects of comb type and presen-
tation order. We analyzed our results for each comb
set with a two-way analysis of variance (ANOVA)
without replication, and so tested for significant dif-
ferences in dance effectiveness in relation to two fac-
tors: comb type and presentation order.
3.1. Laser vibrometry study
Figure 2 shows the results of the vibration
amplitude measurements made with the BW
Figure 2. Results of measure-
ments of the amplitudes of cell
wall vibrations. Each value is
the mean vibration amplitude
measured at a point 4 cells from
the input point. The input vibra-
tion was always a 250 Hz vibra-
tion with 1.0 µm amplitude.
Measurements were made when
comb was 100%, then 50%,
then 25%, and finally 0%
attached to the sides and bottom
of the frame (see Fig. 1).
Plastic foundation and recruitment communication 517
and PW combs. In the BW comb, there was
good transmission of the 250 Hz vibrations
from input point to output point 4 cells away,
though the degree of comb attachment to the
frame strongly influenced the pattern of vibra-
tion transmission. When the BW comb was
100% attached to the wooden frame, we found
a detectable level of vibration only in the cell
rows in the middle of the comb. In contrast,
when the BW comb was 0% or 25% attached
to the wooden frame, we found a detectable and
essentially uniform level of vibration in all the
cell rows, from bottom to middle of the comb.
Curiously, when the comb was 50% attached to
the frame, we found a considerable level of
vibration in bottom and middle cell rows, but
not in intermediate cell rows.
The results differ markedly for the PW
comb, which showed poor transmission of the
250 Hz vibrations. We found a detectable level
of vibration only when the PW comb was little
(0 or 25%) attached to the frame, and even then
only in the bottom cell row.
We do not show the results for the PP comb
in a graph because they are easily reported in
words: for all cell rows, and for all degrees of
attachment of comb to frame, we detected no
vibrations in cell walls located 4 cells from the
input point.
3.2. Behavioral study
Table II summarizes our results on the effec-
tiveness of waggle dances performed on differ-
ent types of comb. We found no effect of comb
type on dance effectiveness. The mean dance
effectiveness was similar for the three types of
comb: 0.05 recruits per waggle run (see Fig. 3).
ANOVA tests show that the mean values of
dance effectiveness do not differ significantly
among comb types, both for comb set 1 (F2, 4 =
0.16, P > 0.90) and for comb set 2 (F2, 4 = 2.94,
P > 0.25).
We did find, however, a strong effect of
position in the test series on dance effectiveness
(see Fig. 4). Regardless of comb type, the comb
presented at the start had a lower mean dance
effectiveness (approx. 0.025 recruits per wag-
gle run) than the comb presented in the middle
(approx. 0.050 recruits per waggle run), which
in turn had a lower mean dance effectiveness
than the comb presented at the end (approx.
0.068 recruits per waggle run). ANOVA tests
show that the mean values of dance effective-
ness differ significantly among test positions,
both for comb set 1 (F2, 4 = 21.95, P < 0.01) and
for comb set 2 (F2, 4 = 8.85, P < 0.04).
Table II. Mean values of dance effectiveness (recruits per waggle run) for two sets of combs. For both sets,
we show the measured value of dance effectiveness for each combination of comb type and test position.
Comb set 1 results Comb set 2 results
Position in test series Position in test series
1st 2nd 3rd Row means 1st 2nd 3rd Row means
Comb type BW 0.027 0.042 0.071 0.047 0.023 0.048 0.066 0.046
PW 0.022 0.054 0.060 0.045 0.022 0.050 0.088 0.053
PP 0.027 0.058 0.061 0.049 0.032 0.062 0.063 0.052
Column means 0.025 0.051 0.064 0.026 0.053 0.072
Figure 3. Comparison of the three types of comb
with respect to mean dance effectiveness.
518 T.D. Seeley et al.
The results described so far are based on
bees performing dances that were illuminated
by daylight for video recording. Having found
no effect of comb type on dance effectiveness,
we wondered if this was because the dances
were performed in daylight. Perhaps dance fol-
lowers had easily found dancing bees by seeing
their movements rather than by detecting their
comb vibrations. If so, then differences among
the comb types in vibration transmission would
not matter.
To see if we had obscured an effect of comb
type on dance effectiveness by studying dances
performed under lighted conditions, we per-
formed another test with the combs of comb
set 1. For each comb, we checked whether
dances were less effective when performed on
a darkened vs. a lighted dance floor. We rea-
soned as follows. Hypothesis: dances on combs
with plastic foundation were as effective as
dances on combs with beeswax foundation
because dances produced on a lighted dance
floor are unusually easy to find. Prediction:
dances on combs with plastic foundation will
be less effective on a darkened dance floor than
on a lighted one. To check this prediction, we
made recruitment measurements under con-
stant conditions except that part of the time the
observation hive’s cover was off (dance floor
lighted) and part of the time the cover was on
(dance floor darkened). The results are shown
in Figure 5. For all three types of comb, we
found no sign of reduced dance effectiveness
with darkened dance floor.
Do combs built with plastic foundation
hinder the bees’ ability to communicate with
waggle dances? Evidently, they do not. In our
Figure 4. Comparison of the three positions in the
test series with respect to dance effectiveness.
Figure 5. Results of tests for effect of light on
dance effectiveness. The number of bees recruited
every 10 min by 10 dancers visiting a feeder was
measured for a 150 min period. For the first 50 min
and last 50 min, the observation hive was not
covered, so the dances were performed in daylight.
For the middle 50 min, the observation hive was
covered, so the dances were performed in darkness.
The same result was found for all three types of
comb: no difference in the mean number of recruits
per 10 min between the cover-off and cover-on
Plastic foundation and recruitment communication 519
behavioral tests, we found no sign whatsoever
of reduced dance effectiveness when bees
danced on combs built with plastic foundation
relative to combs built with beeswax founda-
tion. Because we measured the effectiveness of
dances performed in daylight in an observation
hive, we must ask whether our conclusion (no
hindrance of dance communication by plastic
foundation) also holds true for dances per-
formed in darkness in a normal hive.
Two lines of evidence indicate that we did
not get falsely negative results. The first is that
previous investigators of substrate effects on
waggle dance communication (Tautz, 1996;
Tautz and Rohrseitz, 1998) also studied dances
performed in daylight and they found strong
substrate effects, e.g., differences in recruit-
ment success between dances performed on
open cells vs. sealed cells. Clearly, substrate
differences can influence dance effectiveness
even when dances are performed in daylight.
The second line of evidence is the finding in the
present study that when we darkened the dance
floor, we found no drop in recruitment by bees
dancing on combs built with plastic foundation.
If these combs were inferior substrates for
dances performed in darkness, then we should
have seen reduced recruitment when we dark-
ened the dance floor, but we did not. We
believe, therefore, that our main conclusion –
combs built with plastic foundation do not
hinder waggle dance communication – is a true
negative, not a false one.
The finding that combs built with plastic
foundation provide a fully suitable substrate for
waggle dances was, at first, surprising because
our laser vibrometry study revealed much
poorer transmission of 250 Hz vibrations in
plastic-based vs. beeswax-based combs. How
can we resolve this seeming conflict between
the two parts of our study? The most likely
explanation is that dance followers used fre-
quencies other than the 250 Hz that we used in
our laser vibrometry study. When performing
a waggle dance, a bee produces not only 250 Hz
vibrations by activating her thoracic muscula-
ture, but also 15 Hz vibrations by swinging her
body side to side. We suggest that dance fol-
lowers use mainly the 15 Hz vibrations to orient
to dancers. We chose 250 Hz displacements for
our laser vibrometry study because prior inves-
tigations found that unframed combs have a
striking peak in transmission effectiveness
(Fig. 7 in Sandeman et al., 1996) and a sharp
drop in impedance (Fig. 25 in Rohrseitz, 1998)
for 250 Hz vibrations. These same studies also
found, however, that framed combs have poor
transmission and high impedance of 250 Hz
vibrations, but have good transmission and low
impedance of 15–20 Hz vibrations (Fig. 10 and
12 in Sandeman et al., 1996, and Fig. 26 in
Rohrseitz, 1998). If a framed comb is freed
from the frame on bottom and sides, then there
is improved transmission effectiveness of
250 Hz vibrations (a finding confirmed by our
results, see Fig. 2), but even so it remains below
the transmission effectiveness of 15–20 Hz
vibrations. In hindsight, it is regrettable that we
studied the effects of plastic foundation on
vibration transmission using only 250 Hz stim-
uli. This work needs to be extended using 15 Hz
stimuli. We predict that the transmission of
15 Hz vibrations will not differ between
framed combs built with beeswax vs. plastic
Besides testing the effects of plastic founda-
tion on dance communication, our behavioral
study also yielded a curious finding about the
waggle dance system of bee communication:
its effectiveness changed markedly over the
course of a day. As is shown in Figure 4, we
found that dances performed early in the day
were only one third as effective in getting
recruits to the feeder compared to dances per-
formed later in the day. This change in dance
effectiveness appears robust, for we found the
same pattern on all 6 days of data collection
(Tab. II). How did this change in dance effec-
tiveness arise? Perhaps it reflects a gradual acti-
vation of the study colony’s foragers over the
course of a day, so that more bees were ready
to follow dances later in the day. Another pos-
sibility, however, is that bees that had followed
a dance for the feeder and left the hive to search
for it somehow found it easier to find the feeder
later in the day. Future studies are needed to see
if the daily increase in dance effectiveness that
we found is a typical, and if so, to determine
whether it reflects a gradual “waking up” of the
foragers in a honey bee colony.
This project was supported by the National
Research Initiative of the USDA Cooperative State
Research, Education and Extension Service, grant
520 T.D. Seeley et al.
number 2004-35302-14838, and by the Alexander
von Humboldt Foundation, which has provided
TDS with a Research Award. We are also grateful
to Dr. Stephen A. Teale, director of the Cranberry
Lake Biological Station, for continuing to provide
us with space and facilities at this lovely field sta-
RésuméLes fondations en plastique gênent-
elles la communication dansée ? Ces dernières
années les fondations en plastique se sont largement
répandues chez les apiculteurs, mais aucune étude
n’a été faite pour savoir si elles gênaient le recrute-
ment en réduisant la transmission des vibrations du
rayon produites par les abeilles qui exécutent la
danse frétillante. On sait que les danseuses se fient,
au moins partiellement, aux vibrations du rayon pour
communiquer les sources de nourriture, probable-
ment parce que ces vibrations aident les abeilles à
localiser les danseuses dans l’obscurité de la ruche.
Nous avons étudié trois types de rayon : (i) rayon
construit sur une fondation en cire d’abeilles fixée à
un cadre en bois (BW), (ii) rayon construit sur une
fondation en plastique fixée à un cadre en bois (PW),
(iii) rayon construit sur une pièce unique fondation/
cadre en plastique (PP).
Nous avons utilisé la vibrométrie laser pour compa-
rer les trois types de rayons en termes de
transmission des vibrations produites par les danses.
Les rayons construits sur une fondation en plastique
transmettaient mal les vibrations de 250 Hz. Par con-
tre ceux construits sur une fondation en cire d’abeille
les transmettaient très bien, en particulier lorsque le
rayon était partiellement détaché du cadre en bois
(Fig. 2).
Nous avons réalisé une étude comportementale pour
comparer l’efficacité du recrutement (ER) des dan-
ses sur les différents types de rayons. L’ER a été
mesurée en termes de nombre d’abeilles recrutées
sur un nourrisseur à 350 m par un trajet frétillant pro-
duit pour le nourrisseur. Nous n’avons trouvé aucun
effet (Fig. 3). Les trois types de rayon ont donné
la même valeur moyenne d’efficacité : environ
0,05 recrue par trajet frétillant. Quel que soit le type
de rayon, l’ER différait notablement selon que
c’était le premier, le second ou le troisième rayon uti-
lisé dans la journée. Les danses exécutées en fin de
journée avaient une plus grande efficacité que celles
exécutées plus tôt (Fig. 4). Nous suggérons que
l’augmentation de l’efficacité au cours de la journée
reflète une activation graduelle des butineuses de la
colonie, si bien que les ouvrières sont plus nombreu-
ses à suivre les danses plus tard dans la journée.
Que nous n’ayons pas trouvé un effet du type de
rayon sur l’ER peut être dû au fait que notre expéri-
mentation a été réalisée en lumière du jour. Cela a
pu aider les abeilles à trouver les danseuses en
voyant leurs mouvements plutôt qu’en sentant leurs
vibrations. Nous avons testé cette hypothèse en cher-
chant si les danses exécutées sur les rayons PW et
PP, qui ont une faible transmission des vibrations de
250 Hz, voyaient leur efficacité diminuer lorsque le
plancher des danses était assombri. Nous n’avons
trouvé aucun signe montrant une réduction de l’ER
(Fig. 5). Nous en concluons que l’absence d’effet du
type de rayon sur l’efficacité des danses est un résul-
tat négatif vrai et non un faux. Il est clair qu’un rayon
construit sur une fondation en plastique constitue un
substrat tout à fait adapté à la communication par la
danse frétillante.
Apis mellifera / vibration du rayon / fondation en
plastique / communication / recrutement / danse
ZusammenfassungBehindert eine Kunststoffmit-
telwand der Bienewabe die Tanzkommunikation?
In den letzten Jahren werden von Imkern zunehmend
Mittelwände aus Kunststoff für den Ausbau von
Waben eingesetzt. Es war bisher nicht untersucht, ob
solche Mittelwände die Rekrutierung zu Futterplät-
zen beeinträchtigen, etwa indem sie die von den
Bienen beim Schwänzeltanz erzeugten Schwingun-
gen reduzieren. Es ist bekannt, dass tanzende Bienen
für ihre Futterkommunikation zumindest teilweise
auf über die Wabe weitergeleiteten Schwingungen
angewiesen sind. Vermutlich helfen diese Schwin-
gungen tanzenden Bienen in der Dunkelheit des
Nestes zu lokalisieren. Wir untersuchten die drei fol-
genden Wabenarten: Auf Mittelwänden oder auf
Plastikmittelwänden gebaute und in ein Holzrähm-
chen eingepasste Waben (BW bzw. PW), und eine
aus einem Stück bestehend Einheit aus Mittelwand
und Rähmchen ausgebaute Wabe (PP).
Wir benutzten Laservibrometrie, um die drei
Wabentypen auf ihre Weiterleitungseigenschaften
für die beim Tanz entstehenden Vibrationen zu
untersuchen. Auf Plastikmittelwänden gebaute
Waben leiteten Vibrationen mit einer Frequenz von
250 Hz schlecht weiter. Waben mit Wachsmit-
telwänden leiteten diese Frequenz dagegen gut
weiter, besonders wenn sie teilweise nicht mit dem
umgebenden Rähmchen verbunden waren (Abb. 2).
Anhand einer Verhaltensuntersuchung verglichen
wir die Effizienz der Rekrutierung von Bienen durch
die Tanzkommunikation auf den drei Wabentypen.
Als Rekrutierungseffizienz wurde die Anzahl der zu
einer Futterstelle angeworbenen Bienen pro gezeig-
tem Schwänzeltanz bestimmt. Wir fanden keinen
Einfluss des Wabentypes auf die Rekrutierungseffi-
zienz (Fig. 3); auf allen drei Wabentypen war diese
ähnlich und betrug ungefähr 0.05 rekrutierte Bienen
pro Schwänzellauf. Wir fanden allerdings einen
Positionseffekt in der Testserie. Unabhängig von
dem Wabentyp gab es deutliche Unterschiede in der
Effizienz der Tänze je nachdem ob diese auf der ers-
ten, zweiten oder dritten der im Verlauf des Tages
genutzten Waben stattfanden.
Später am Tag durchgeführte Tänze waren effektiver
als früh am Tag durchgeführte (Fig. 4). Wir nehmen
an, dass dieser Anstieg in der Tanzeffektivität im
Tagesverlauf auf die allmähliche Aktivierung der
Sammlerinnen in einem Volk zurückzuführen ist,
Plastic foundation and recruitment communication 521
wonach später am Tag mehr Sammlerinnen bereit
wären den Tänzen zu folgen. Es hätte sein können,
dass ein Wabeneffekt deshalb nicht gefunden wurde,
weil die Experimente unter Tageslichtbedingungen
durchgeführt wurden unter denen es für die Bienen
leicht gewesen sein könnte, Tänzerinnen auch ohne
Wabenvibrationen anhand ihrer Bewegungen zu fin-
den. Wir untersuchten daher bei verdunkeltem
Tanzboden, ob die Tanzeffektivität auf PP Waben
mit ihrer schlechteren Weiterleitung für Vibrationen
von 250 Hz geringer war als bei PW Waben. Hierbei
ergab sich kein Anzeichen einer verringerten Tanz-
effektivität (Fig. 5) woraus sich schließen lässt, dass
das negative Ergebnis über einen Wabeneffekt ein
wahres, nicht falsches Negativergebnis darstellt.
Offensichtlich stellt eine auf einer Plastikmittelwand
gebaute Wabe einen vollständig geeigneten Unter-
grund für die Schwänzeltanzkommunikation dar.
Apis mellifera / Wabenvibration / Plastikmit-
telwände / Futterkommunikation / Schwänzel-
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To access this journal online:
... A number of "waggle dances" are used to convey information about foraging, food supplies, and nesting. The dances contain vibrating "steps" at various frequencies, mostly clustered from 15 to 20 Hz [549] or from 200 to 300 Hz. [550][551][552][553] The open cells of a honeycomb resonate around 20 or 250 Hz, thereby amplifying vibrations at these frequencies and turning the honeycomb into a mechanism for wide signal broadcasting. [554] Bees that dance on empty, uncapped cells are able to recruit around twice as many bees than those that dance on capped brood cells; [555] these followers are also recruited from a greater distance. ...
... [554] Bees that dance on empty, uncapped cells are able to recruit around twice as many bees than those that dance on capped brood cells; [555] these followers are also recruited from a greater distance. [550,556] Another vibration-mediated behavior known as "shimmering" arises in the giant honeybee, Apis dorsata, upon the emergence of a threat such as a predator. Shimmering behavior is a social motion, similar to "the wave" in football stadiums, [557] in which the bees on the surface of a nest all periodically (<1 Hz) raise their abdomens in a manner that propagates across the surface, often emanating from a central locus. ...
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Over the course of their wildly successful proliferation across the earth, the insects as a taxon have evolved enviable adaptations to their diverse habitats, which include adhesives, locomotor systems, hydrophobic surfaces, and sensors and actuators that transduce mechanical, acoustic, optical, thermal, and chemical signals. Insect-inspired designs currently appear in a range of contexts, including antireflective coatings, optical displays, and computing algorithms. However, as over one million distinct and highly specialized species of insects have colonized nearly all habitable regions on the planet, they still provide a largely untapped pool of unique problem-solving strategies. With the intent of providing materials scientists and engineers with a muse for the next generation of bioinspired materials, here, a selection of some of the most spectacular adaptations that insects have evolved is assembled and organized by function. The insects presented display dazzling optical properties as a result of natural photonic crystals, precise hierarchical patterns that span length scales from nanometers to millimeters, and formidable defense mechanisms that deploy an arsenal of chemical weaponry. Successful mimicry of these adaptations may facilitate technological solutions to as wide a range of problems as they solve in the insects that originated them.
... set danceFollowersNectar danceCircuits * 0.05 based on the assumption that 0.05 foragers can be recruited per dance circuit (Seeley et al. 2005). DanceFollowersNectar is patch specific and updated every foraging round, as nectar collection by the foragers may affect the handling time and hence the energetic efficiency. ...
... ;*************************************************************************** ; ************** PARAMETERIZATION FLOWER PATCH ********************** ;*************************************************************************** to CreateFlowerPatchesProc ; creates 2 flower patches ("red" & "green"), ;*************************************************************************** ; ************** PARAMETERIZATION FLOWER PATCHES FROM FILES ******** ;*************************************************************************** to Create_Read-in_FlowerPatchesProc ; copy of CreateFlowerPatchesProc but data are read from input file ; calculates derived values (e.g. EEF, flight costs etc) ; ********************************************************************************************************************** ; in this case, foragers always dance for their patch, ; irrespective of its quality set danceFollowersNectar danceCircuits * 0.05 ; Seeley, Reich, Tautz (2005): "0.05 recruits per waggle run (see Fig. 3)" ] ; ask flowerPatches end ; ********************************************************************************************************************** to ForagingRoundProc ; CALLED BY Start_IBM_ForagingProc calls Procedures involved in each foraging trip ; and does foraging related plots set ColonyTripDurationSum 0 set ColonyTripForagersSum 0 ; used to calculated duration of this foraging round set DecentHoneyEnergyStore (TotalIHbees + TotalForagers ) * 1.5 * ENERGY_HONEY_per_g ; DecentHoneyEnergyStore reflects the amount of energy a colony should store ; to survive the winter, based on the assumption that a bee consumes ca. let plotname (word "Generic plot " i) ; e.g. ...
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(JPEbecherSA1_ODD.pdf): Detailed model description, following the ODD (Overview, Design concepts, Details) protocol (Grimm et al. 2006, 2010).
... The number of dance circuits of returning foragers is multiplied by 0.05 to determine the number of foragers that are following to the patch. Reference is made to Seeley et al. (2005) in order to justify the value of 0.05 followers per dance circuit. ...
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The Panel has interpreted the Terms of Reference by carrying out a stepwise evaluation of the BEEHAVE simulation model with a view to assessing its suitability for use in a regulatory context and for risk assessment of multiple stressors at the landscape level. The EFSA opinion on good modelling practice was used to evaluate the model and its documentation systematically. The overall conclusion is that BEEHAVE performs well in modelling honeybee colony dynamics, and the supporting documentation is generally good but does not fully meet the criteria of the good modelling opinion. BEEHAVE is not yet usable in a regulatory context primarily because it needs a pesticide module. BEEHAVE has a Varroa/virus module, although this seems to underestimate the impact of Varroa/virus on colony survival, and additional stressors (chemical and biological) would need to be added to allow investigation of the effects of interactions of pesticides with multiple stressors. BEEHAVE currently uses a very simple representation of a landscape and this should be extended. There is only one environmental scenario in the present version of BEEHAVE (European central zone-weather scenarios for Germany and the UK) and extension to other European zones would be needed. The supporting data and default parameter values should be further evaluated and justified. The modelling environment used by BEEHAVE (NetLogo) has an excellent user interface but provides limited opportunities for extending the model. The Panel recommends that BEEHAVE should be adopted as the basis for modelling the impact on honeybee colonies of pesticides and other stressors, but that further development should use a standard, object-oriented language rather than NetLogo.
... The lowest frame (hereafter, Bthe focal comb^) contained an area of comb that was built naturally by the bees before the experiment (no wires, wax foundation, etc.), and so was more representative of combs found in natural nests. Most importantly, the focal comb had no wires to artificially stabilize the comb in the wooden frame, because unnatural materials can affect comb vibrations (Seeley et al. 2005). To measure comb vibrations on the focal comb, we pressed the Bgreen board^(3.2 ...
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Communication is impossible if the sender’s signal cannot overcome background noise to reach the receiver. This obstacle is present in all communication modalities, forcing organisms to develop diverse mechanisms to overcome noise. Honey bees will modify combs to improve signal efficiency of substrate-borne vibrations, but it is unknown whether, and if so, how, bees compensate for the largest potential source of noise: the bees themselves. The number of bees in a colony changes markedly throughout the year, but the size of the nest cavity does not, forcing workers into high densities on the combs. How, then, do bees communicate via substrate-borne vibrations on combs that are covered in bees? We used accelerometers to measure comb vibrations, while varying the number of workers on the comb. Surprisingly, comb vibrations decreased with increased worker number. Furthermore, inserting freshly killed bees to the comb demonstrated that it is not simply the bees’ collective mass that damps vibrations, but is probably their behavior. We propose that their posture damps vibrations, with each bee linking up to six neighboring cells with her legs. This collective damping reduces background noise and improves the landscape for communication. These results demonstrate how living systems, including superorganisms, can overcome physical obstacles with curiously simple and elegant solutions. Significance statement Background noise is a pervasive problem in communication. Honey bees must address this problem because thousands of individuals occupy and communicate within a single nest made of beeswax combs. While it is known that bees use beeswax comb vibrations to communicate, it is unknown how they overcome background noise when the combs become covered in bees. We show that comb vibrations decrease, not increase, as the number of bees on the comb increases. This unexpected result is not due to bees’ mass, but rather their interactions with the comb that damps vibrations. By reducing background vibrations, workers make the comb “quieter” and improve the substrate for communication. Therefore, we show that the communication landscape for sending signals within the superorganism is improved, not hindered, as the colony grows.
... Substrates also influence the transmission of honeybee waggle dance vibrations. Transmission of artificial vibrations of 250 Hz had greater amplitude in honeybee combs made of beeswax than in artificial plastic combs (Seeley et al., 2005). ...
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The influences of artificial and natural rearing substrates on mating success were investigated for the parasitoid wasp Cotesia marginiventris (Cresson) (Hymenoptera: Braconidae), a candidate for augmentative biological control of various lepidopteran pests. Five rearing substrates were tested: plastic, glass, chiffon fabric, and leaves of two host plants, bean [Vigna unguiculata (L.) (Fabaceae)] and maize [Zea mays L. (Poaceae)]. Mating success was highest on chiffon, lowest on plastic and glass, and intermediate on maize and bean. The transmission characteristics of one component (buzz 1) of the courtship vibrations produced by male wing fanning were investigated using laser vibrometry. The duration of buzz 1 was longer on maize, bean, and chiffon than on plastic and glass. The fundamental frequency of buzz 1 (~300 Hz) was lowest on bean and highest on glass, and intermediate among other substrates. The relative amplitude of buzz 1 was higher on chiffon than on plastic, glass, or bean, and intermediate on maize. The relative importance of airborne sound and substrate vibration as courtship signals was also investigated with experiments that manipulated the production of courtship vibrations and the mating substrates. The amplitude of courtship vibrations on chiffon was significantly higher for winged males than for dealated males. The mating success of males was impacted by both the presence of wings and the mating substrate. These findings suggest that mating success and transmission of courtship vibrations are influenced by the rearing substrate, and that courtship vibrations are critical to mating success in C. marginiventris. Future efforts to mass rear this parasitoid and other insects should consider the potential influences of rearing substrates on mating.
... There are a number of selective contexts that could account for changes in acoustic courtship signals. For example, artificial and natural rearing substrates influence vibrational communication and mating in other arthropods including parasitoids wasps (Miklas et al. 2001; Elias et al. 2004; Seeley et al. 2005; Joyce et al. 2008). The courtship acoustics produced by male wing fanning in parasitoids have both an airborne and substrate-borne component (Leonard and Ringo 1978; van den Assem and Putters 1980; Sivinski and Webb 1989; Field and Keller 1993; Joyce et al. 2008), and both types of signals are known to be detected by Hymenoptera (Greenfield 2002). ...
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The courtship acoustics of five species of parasitoid wasps (Hymenoptera: Braconidae), potential candidates for augmentative biological control of Anastrepha (Schiner) species (Diptera: Tephritidae), were compared between recently colonized individuals and those continuously reared 70–148 generations. During courtship, males of these parasitoid species fan their wings and produce a series of low amplitude pulses. The first series of 15 or more continuous courtship pulses was used to measure the pulse duration, frequency, and interpulse interval (IPI) from the beginning, middle, and end of the pulse series. Each parameter was compared between young and old colonies, and among species. Several differences in courtship acoustics were detected in colonies that had been continuously reared. The pulse duration at the end of the pulse series was longer in old colonies for Doryctobracon crawfordi (Viereck) (Hymenoptera: Braconidae), but shorter for old colonies of Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae). The IPI of the middle pulse was shorter in old colonies of Opius hirtus (Fischer) (Hymenoptera: Braconidae), and was also shorter at the last pulse for old colonies of both Utetes anastrephae (Viereck) (Hymenoptera: Braconidae) and D. longicaudata. The duration of the middle pulse distinguished the three native species, and separated the two introduced species from each other. We discuss our findings in light of their biological and applied implications, particularly those dealing with quality control of mass-reared parasitoids. Keywords Anastrepha ludens - Doryctobracon crawfordi - Opius hirtus - Utetes anastrephae - Diachasmimorpha longicaudata - Diachasmimorpha tryoni -Mass rearing-Laboratory selection-Quality-Rearing substrate-Courtship vibration
Communication on honeybee combs includes both distance and direction in the case of waggle dances. Potential recruits attending a dancer emit vibrations which elicit a response from the dancer to give the emitter a sample of nectar. Tooting and quacking by queens are both airborne sounds and substrate vibrations which are carried mainly by the fundamental frequency component. Bees recognize these signals mainly by their temporal structure and comparisons of the threshold, emission level, and attenuation with distance, which suggests that they are used only within a restricted area of the comb. When waggle-dancing honeybees move on comb, they produce vibratory movements that indicate the location of the waggle dancer and the pulsed vibrations are increased during waggle phases, so amplifying the signals for remote dance followers. Because sound intensity decreases with the density of the medium and with distance, beeswax is a medium for sound transmission. Pheromones in comb serve as slow-release systems with long time constants and include transmissions of colony odour, queenrightness, cell capping, colony odour, kin recognition, footprint pheromones, wax-salvaging behaviour etc. The specific dance sites that occur on combs are due to chemical tagging. Masking colony odour occurs when receiver bees are conditioned to the same comb source as introduced bees, which are accepted. A series of only a few methyl esters produced by queens and workers are sufficient to induce capping of mature brood; but capping worker brood may depend on the depth of larvae in comb cells and not just ratios of ester emissions. Nonetheless, these results are not mutually exclusive in principle.
An experiment was performed to see if plastic comb foundation affects a colony's comb building and honey production. Small colonies were installed in hives equipped with one of three types of frame with foundation - wooden frame with beeswax foundation, wooden frame with plastic foundation, and plastic frame with plastic foundation - and their patterns of comb building and weight gain were compared. After 12 weeks, the colonies given beeswax foundation had their hives nearly 100% filled with drawn comb, whereas the colonies given plastic foundation had their hives only 70% filled with comb. Also, the colonies with beeswax foundation had gained weight by 9.6 kg, on average, whereas those with plastic foundation had gained weight by only 4.8 kg, on average. These results confirm previous claims that bees can be reluctant to accept plastic foundation and that plastic foundation can depress a colony's honey production.
This work, a sequel to Honeybees and Wax published nearly 30 years ago, starts with a brief introduction and discussion of nesting sites, their spaces and densities, self-organization of nest contents, and interspecific utilization of beeswax. The following chapters cover communication by vibrations and scents and wax secretion, and discuss the queen in relation to the combs. Discussions on completed nests include the significance of brood, the roles of pollen and nectar flow, and comb-building, and are followed by a triad of related chapters on the construction of cells and combs and their energetic costs. An in-depth examination of the conversion of wax scales into combs, the material properties of scale and comb waxes, and the wax gland complex are presented. The next chapters are devoted to a comprehensive analysis of the literature on the chemistry and synthesis of beeswax, and, finally, the material properties of honeybee silk are highlighted. Content Level » Research Keywords » Beeswax - Biomimetics - Brood - Comb - Comb waxes - Comb-building - Honeybee nests - Honeybee silk - Honeybees - Materials sciences - Nectar flow - Nesting sites - Pollen - Wax gland complex Related subjects » Biomaterials - Biophysics & Biological Physics - Entomology - Organic Chemistry
Honey bees live in groups of approximately 40,000 individuals and go through their reproductive cycle by the swarming process, during which the old queen leaves the nest with numerous workers and drones to form a new colony. In the spring time, many clues can be seen in the hive, which sometimes demonstrate the proximity to swarming, such as the presence of more or less mature queen cells. In spite of this the actual date and time of swarming cannot be predicted accurately, as we still need to better understand this important physiological event. Here we show that, by means of a simple transducer secured to the outside wall of a hive, a set of statistically independent instantaneous vibration signals of honey bees can be identified and monitored in time using a fully automated and non-invasive method. The amplitudes of the independent signals form a multi-dimensional time-varying vector which was logged continuously for eight months. We found that combined with specifically tailored weighting factors, this vector provides a signature highly specific to the swarming process and its build up in time, thereby shedding new light on it and allowing its prediction several days in advance. The output of our monitoring method could be used to provide other signatures highly specific to other physiological processes in honey bees, and applied to better understand health issues recently encountered by pollinators.
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The subgenual organ of the honeybee (Apis mellifera) is suspended in a haemolymph channel in the tibia of each leg. When the leg is accelerated, inertia causes the haemolymph (and the subgenual organ) to lag behind the movement of the rest of the leg. The magnitude of this phase lag determines the displacement of the subgenual organ relative to the leg and to the proximal end of the organ, which is connected to the cuticle. Oscillations of the subgenual organ are visualised during vibration stimulation of the leg, by means of stroboscopic light. Video analysis provides fairly accurate values of the amplitude and phase of the oscillations, which are compared with the predictions of a model.   The model comparison shows that the haemolymph channel can be described as an oscillating fluid-filled tube occluded by an elastic structure (probably the subgenual organ). The mechanical properties of the subgenual organ and haemolymph channel resemble those of an overdamped mass-spring system. A comparison of the threshold curve of the subgenual organ determined using electrophysiology with that predicted by the oscillating tube model suggests that the sensory cells respond to displacements of the organ relative to the leg.
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Vibration of the rims of open cells in a honeycomb, applied in the plane of the comb face, is transmitted across the comb. Attenuation or amplification of the vibratory signal depends on its frequency and on the type of comb. In general, framed combs, both large and small, strongly attenuate higher frequencies, whereas these are amplified in small open combs. The very poor transmission properties of the large framed combs used in commercial hives may explain the bees' habit of freeing an area of comb from the frame in those areas used for dancing. Extracellular electrical recordings from the leg of a honeybee detect large action potentials from receptors that monitor extension of the tibia on the femur. Measurements of threshold displacement amplitudes show these receptors to be sensitive to low frequencies. The amplification properties of unframed combs extend the range of these receptor systems to include frequencies that are emitted by the bee during its dance, namely the 15 Hz abdomen waggle and 250 Hz thorax vibration.
Apis mellifica erzeugt in der Schwänzelphase des Werbetanzes ein Geräusch — die Vibrationsbewegung — dessen Grundfrequenz durchschnittlich 250 Hz beträgt. Das Geräusch hat in 1 cm Abstand über den Flügeln eine Lautstärke von 70–80 Phon. Es wurde elektromagnetisch und akustisch registriert. Die Vibrationsbewegung besteht aus kurzen Impulsen der 250 Hz-Bewegung (Vibrationsstößen). Die Vibrationsstöße dauern ungefähr 15 msec. Ihnen folgen Pausen ungefähr gleicher Dauer. Die Vibrationsstöße werden kontinuierlich während der ganzen Schwänzelphase ausgeführt. Ihr Vorkommen ist auf die Schwänzelphase beschränkt. Eine direkte Beziehung zwischen dem Aufbau der Vibrationsbewegung und der Futterplatzentfernung besteht nicht. Vielleicht dient die Vibrationsbewegung indirekt der Entfernungsmeldung, indem sie die Schwänzelzeit als Signal der Entfernung auffällig für die Nachtänzerinnen markiert.
The dance language of honeybees will always be associated with the name of Karl von Frisch, who was one of the two founders of Zeitschrift fr Vergleichende Physiologie, now the Journal of Comparative Physiology. The discovery of the dance language has already led to a great number of investigations of physiological mechanisms, and more studies can be expected in the future. It therefore seems most appropriate to let this King Solomon Lecture deal with the progress and problems in our efforts to understand the transfer of information in the dance language of honeybees.
Sound and vibrational signals exchanged by honeybees during the performance of wagging dances were simultaneously recorded by means of a microphone and a laser vibrometer. Previous descriptions of the 280-Hz sounds emitted by the dancing bee were confirmed, and no vibrational (substrate-borne) component could be detected. In contrast, the 320-Hz begging signals (emitted by bees following a dancer and used as a request for food samples from the dancer) do vibrate the comb with peak-peak displacement amplitudes up to 1.5 m. Artificially-generated comb vibrations of sufficient amplitude cause bees standing on the comb to freeze. The threshold for obtaining a detectable freezing response was measured for frequencies between 100 Hz and 3 kHz. At 320 Hz it is just below the amplitude of the natural begging signals. Thus it seems likely that these signals are received by the bees as vibrations of the comb. The propagation velocity of waves, damping, and mechanical input impedance of honeybee combs were studied. These results, combined with the observed amplitudes of the begging signals, support the assumption that the begging signals are generated with the flight muscles. The begging signal propagates as a bending wave. The attenuation of the begging signal with distance is relatively small, so the amplitude of the signal probably needs to be carefully adjusted in order to restrict the range of the communication.
In the vicinity of a dancer, a honeybee can become a dance follower after touching the dancer or a dance follower with an antenna. If the attraction occurs without such antennal contact, the strength of the attraction over distance depends on several factors: the kind of dance floor (empty open cells versus capped brood cells); whether dancers and dance followers stand on the same substratum or on separate substrata; the position and direction of the attracted bee relative to the dancer bee; the size of the dance group (the dancer plus follower bees); and the light conditions under which the dance takes place. Dances on open cells are significantly more attractive than dances on sealed cells. Dancers on open cells attracted 90% of all followers from within 27 mm (about five to six cell diameters). Dancers on sealed cells attracted 90% of all followers within 18 mm (about three cell diameters). The majority of bees that were attracted by the dancer were standing laterally to the dancer. Dances illuminated by artificial visible light are significantly more attractive than dances illuminated by infrared light. As a group, “glassplate bees” (bees standing mechanically isolated from the dancer bee) were least attracted.
1. A pulsed sound of approximately 200 cps. which is produced during the straight run of the honey bee waggle dance is described. The presence of this sound provides a new possibility for explaining the method whereby information about the distance of the food source from the hive is communicated among bees. Besides the elements considered by von Frisch & Jander (1957), sound production time during the straight run and number of pulses present in the straight run are each shown to be also capable of carrying distance information. Since the ratio of the sound pulse rate to waggle rate is approximately 2·5 to 1, the sound is not an incidental result of waggling of the abdomen by the dancing bee.
The waggle dance of the honeybee Apis mellifera, used to recruit nestmates to a food source, takes place on the surface of the combs in the dark hive. The mechanism of information transfer between dancer and follower bees is not entirely understood. The results presented here reveal a novel factor that must be brought into any consideration of this mechanism, namely that the nature of the floor on which the bees dance has a considerable influence on the recruitment of nestmates to a food source. Dancers on combs with open empty cells recruit three times as many nestmates to a food source as dancers on capped brood cells.