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10 June 2005
Once again we are treated
to a presumed final proof of bee "language."
The latest effort ("The flight paths of honeybees recruited
by the waggle dance") appears as a letter on pages 205-207
in Nature on 12 May 2005 by J. R. Riley, U. Greggers,
A. D. Smith, D. R. Reynolds, and R. Menzel. Is the question of
bee language now resolved? Hardly, because the nearly four-decade
old controversy does not revolve about evidence - as will be
documented in this treatise.
Ruth Rosin recognized that
element of the problem when she posted (6 June 2005) the following
comment (in part) on the honey bee e-mail network (BEE-L):
| "The Nature
(2005) radar-tracking study by Riley et al. has already created
considerable excitement in the popular scientific news-media.
The study, nonetheless, never did, nor could, salvage the dance
language hypothesis, because the whole study is simply totally
irrelevant to the dance language controversy" |
| "According
to the report only 2 bees, (out of the 19 radar-tracked bees
released near the hive, for which flight-tracks are provided)
"landed at the feeder." In response to questions about
various details concerning the study, Uwe Greggers (the scientist
who actually designed the study, however, informed me, among
others, (in e-mail exchanges), that those 2 bees did not actually
land at the feeder. Instead, they only landed on the stand on
which the feeder stood, but never found the food, or the feeder
(even when they were not more than 8 cm away from the feeder).
Anyone who questions that is free to personally check with him. |
Two days earlier she had also
written to me (in part):
| "There is,
however, another, much more basic problem with the Nature
(2005) study. The whole study is simply totally irrelevant to
the DL controversy. Before seeing the published report I assumed
that the experimenters did what they should have done first and
foremost, i.e. radar-track recruits recruited by foragers feeding
on scented food; which is what foragers invariably do in nature.
It turns out that the experimenters did exactly the opposite.
They strove very hard, and apparently succeeded in radar-tracking
only bees that never found any food." |
| "The honeybee
DL hypothesis was intended to provide an answer to the problem
how honeybee recruits find their foragers' food-source, and other
sources with the same major odor, in the field. In no way is
it possible to provide any answer to such a problem by studying
only bees that never found any food during the study. Since the
whole study is totally irrelevant to the problem, any results
obtained in the study cannot be relevant to the problem, either." |
A Broader Problem
During the past 35 years, the
dance language controversy has centered on the question, "Can
someone prove a hypothesis true and expect that hypothesis to
become a fact, a fact no longer open to question?" Bee language
advocates would have us believe so. Those who study scientific
process feel otherwise.
We published the results of
our double controlled and strong inference experiments in the
late 1960s and 1970s, results that did not mesh with predictions
of the dance language hypothesis (see below). Ever since then,
research on honey bee recruitment to food crops has remained
in a crisis state, with millions having been spent on efforts,
in various series of experiments, to "prove," once
more, that bee language is real. In each attempt, bee researchers
have admitted that previous attempts had not sufficed - but did
so only after they felt certain that their own efforts had succeeded.
The recent radar tracking studies
of bee flight paths that left the hive is just one more in such
a series of attempted "proofs." On 14 October 2003
James Fischer posted the following statement on BEE-L, one made
by those involved in that radar research:
| "We have used
harmonic radar to measure the flight trajectories of bees recruited
after observing the waggle dance, this has enabled us to settle
(hopefully once and for all) this controversy in favour of Von
Frisch." |
In the first few words of their
2005 Nature report, the radar tracking researchers
also used the phrase, "the dance language" of bees
instead of more objective phrases, such as: "the waggle
dance," or "the dance maneuver." It became clear
soon thereafter in their letter that they had started their experiments
with the assumption that bee language is real and then had tried
to prove it true (circular reasoning?).
The renowned science philosopher
Karl Popper would have been appalled at such a brazen and naïve
approach. For instance, in 1957 he stated: "It is easy to
obtain confirmations, or verifications, for nearly every theory
- if we look for confirmations." In scientific experimentation,
the easiest one to convince is oneself. That is why we need blind,
double controlled, and other rigorous designs in experimentation
- elements lacking in the present study.
Background Information
Karl von Frisch abandoned an
earlier odor-search hypothesis and proposed his dance language
hypothesis in 1946, a conclusion that rapidly became accepted
nearly universally (including by me for my doctoral studies in
the 1950s). That hypothesis provided a rational (though not exclusive)
explanation about how newly recruited bees might manage to find
a food source exploited by regular foragers. His interpretation
morphed into the status of fact ("proved" or "discovered"),
rather than remaining in people's minds the hypothesis that it
was.
What few appreciate today is
that von Frisch's language conclusion was at odds with his earlier
published results and conclusion; that is, recruited bees use
odor, and only odor, after leaving their parent colony in search
of the crops exploited by experienced foragers. In 1937, for
instance, he wrote:
| "I succeeded
with all kinds of flowers, with the exception of flowers without
any scent. When the collecting bee alights on the scented flowers
to suck up the food, the scent of the flower is taken up by its
body-surface and hairs, and when it dances after homing the interested
bees perceive the specific scent on its body and know what kind
of scent must be sought" |
For more information on that point, please access:
http://www.beesource.com/pov/wenner/bw1993.htm
The exotic notion of bee "language,"
though unproven and lacking rigorous experimental tests, eclipsed
von Frisch's earlier odor-search hypothesis. One prominent bee
researcher, H. Kalmus, held out. He wrote in 1960 (Simple
Experiments with Insects. Doubleday & Company, Inc.
Garden City, NY. p. 96):
| "The explanation
[of how a forager communicates this information], however, is
really quite simple; and any fairy tales about one bee telling
the others, or leading the others to a locality, can be discounted.
When a bee returns to the hive she starts dancing on the combs
beating her wings and thus spreading the smell of the flower
which clings to her body. The other bees become interested by
the dance and go searching for that particular smell." |
That comment by Kalmus thus
matched the firm conclusion von Frisch had published in 1937.
Although I conducted my doctoral
research in the 1950s with the belief in bee language, later
experiments by my co-workers and myself yielded results clearly
at odds with that hypothesis. A series of unfortunate and unexpected
events (for example, see http://www.beesource.com/pov/wenner/aoac.htm)
led us to execute rigorous experimental tests of the language
hypothesis during the 1960s, the first such real tests ever conducted.
(For clarification, experiments designed to gain support for
a hypothesis do not count as tests.)
The first set of experiments
involved a double control design. Bees from one hive foraged
at only one station in a set of four; bees of another color foraged
at all four stations. In contrast to predictions of the dance
language hypothesis, recruits from both hives ended up in equal
proportions at the four stations, despite the wide disparity
of information presumably provided by dancing bees in the two
hives. For details, see:
http://www.beesource.com/pov/wenner/scifeb1967.htm
http://www.beesource.com/pov/wenner/scifeb1967b.htm
The second set of experiments
involved only one hive. In that case, a constant set of marked
foragers imbibed scented sugar solution at two stations for 3
hours each day during a 24-day period. All unmarked bees were
caught and tallied during that period. On some of the days, we
switched to unscented food at the target stations but provided
the scented food at a third station.
On those subsequent days, recruited
bees ignored the dance maneuver information in their parent colonies
and arrived, instead, at the third station where we had used
scented food that had been provided on the previous day. For
details see:
http://www.beesource.com/pov/wenner/sci1969.htm
Those sets of experiments are
straightforward (with mutually exclusive designs) and the results
should have been heeded. That did not happen. Instead, bee language
proponents (by then, their numbers were legion) erred most seriously
in two ways, errors repeated up to the present time:
1) Language advocates ignored
Karl Popper's caution: "Some genuinely testable theories,
when found to be false, are still upheld by their admirers -
for example, by introducing ad hoc some auxiliary assumption,
or by re-interpreting the theory ad hoc in such a way
that it escapes refutation."
2) Language advocates apparently accept as valid only evidence
that supports the von Frisch hypothesis. It appears that they
consider counter evidence just "noise" in the system.
Back in 1865 Claude Bernard
admonished against falling into such error: "[The experimenter]
must never answer for [Nature] nor listen partially to her answers
by taking, from the results of an experiment, only those which
support or confirm his hypothesis."
In short, during the past few
decades, bee language advocates have repeatedly committed the
error of only focusing on supportive evidence, while ignoring
or discounting other evidence that does not fit within their
"belief system." That is, only positive evidence counts.
Flaws in "Proof"
Experiments
During the past few decades
we have seen the same mistakes occur repeatedly; language advocates
have sought and embraced supportive evidence and dismissed negative
evidence in an almost pathological manner. Any new supportive
evidence always received wide acclaim, with great fanfare in
the media (the exotic sells). Manuscripts submitted in support
of the language hypothesis became published readily. Grant proposals
that might yield positive evidence received funding. In fact,
during the past several decades, millions have been spent on
efforts to "prove" the language hypothesis true, once
and for all.
Time in Flight and Success
Rate for Searching Recruits
First came the results of some
experiments by three undergraduates (Gould, Henerey, and MacLeod)
that actually yielded results that did not mesh with predictions
of the language hypothesis.
For example, they had marked
with individual numbers a couple of thousand bees in the hive.
They recorded 277 that left the hive after contacting a dancer.
Of those, only 37 found either one of the two stations located
in opposite directions from their hive. However, a third of those
ended up at a station in the exact opposite direction from which
the dance maneuver had presumably indicated the food location.
Nevertheless, they concluded that attendant bees had used direction
information obtained from the dancing bee. Despite that serious
discrepancy, the journal (Science) would not permit
us to publish a rebuttal (see: http://www.beesource.com/pov/wenner/EXC.htm).
"Misdirection"
Experiments
Next came James Gould's "misdirection"
experiments. By shining a bright light at an angle to dancers
in the hive, he reportedly changed the direction of the dance
maneuver. For his first publication in that series (Nature
1974), in support of bee language, he included the results of
only three of the 33 half-hour runs he had run. While first hailed
as "elegant," it later became apparent that the small
sample sizes in his experiment and repeated publication of the
results of his experiments in different display formats grossly
exaggerated the importance of his contribution. Neither has anyone
successfully repeated his experiments.
Mechanical Bee Experiments
After people finally became
disenchanted with Gould's results, researchers in Germany and
Denmark published (with much fanfare) the results they had obtained
by use of a mechanical bee ("robot" bee). They also
claimed that they had obtained the final resolution of the controversy.
Once again, though, it became apparent that they had not resolved
the matter after all. They had to use odor in their experiments
in order to obtain recruitment. (Without odor, there is no recruitment,
just as von Frisch observed.) In one experiment, two real foragers
plying between hive and station resulted in 50 recruits showing
up at the test station. By contrast, only two recruits finally
arrived at the feeding station after numerous bees had attended
a robot bee. Support for their conclusions gradually faded.
It is no wonder that the honey
bee dance language controversy has now persisted for nearly half
a century. Others conducted less exotic studies during the past
few decades and claimed that they had successfully resolved the
issue of bee "language." As usual, though, they tailored
their experimental designs toward gaining supportive evidence
for the hypothesis - rather than conducting true tests of that
hypothesis.
Radar Tracking of Recruited
Honey Bees
The latest "high tech
confirmation" experiments involved radar tracking of recruited
honey bees. Once again we see the usual pattern: a small sample
size, biased researchers, much media splash, and dogmatic conclusions.
Never mind that the evidence doesn't fit the original hypothesis;
they now considered that they had obtained "proof"
of bee "language." In an interview with a science reporter
for the San Francisco Chronicle, for example, the
lead scientist of the radar tracking project was quoted as insisting:
"'no really sensible person can come to any other conclusion'
than that von Frisch was right."
In their experiments the radar
tracking researchers made some serious errors in approach. Consider
some background. The researchers had some bees trained to fly
out to a feeding station and saw them later dance. They followed
a bee that attended such a dance, caught it as it left the hive,
attached a transponder onto its body, released it, and then followed
the path of that bee with radar. To conduct their study they
embraced a number of assumptions:
1) That a bee attending a dancer
has the neurological/physiological equipment to obtain abstract
physical information from another bee.
2) That such a particular bee leaving a dancer "intended"
to travel to the "target" site (that experimental bees
had gained a route memory of the feeding station relative to
the hive).
3) The capture of a bee and fastening a transponder onto it doesn't
interfere with its presumably "programmed" behavior.
4) That the released bee flight path has only one explanation
(in favor of "language," in this case).
5) That they worked in an "odor free" area (odor free
to bees) for their experiments.
From the account published
in Nature, it appears that some critical controls
were missing, including:
1) Flight paths of bees that
had been similarly treated but had not attended a waggle dance.
2) Flight paths of departing bees that had attended a waggle
dance that had visited food located in a different direction.
3) Flight paths of bees searching for a scented food source (as
pointed out by Rosin, above).
That is, the reasoning seems
to be: if the escaping bees fly off in the "right"
direction (the only direction arranged for), then they
have used their language.
All of the above reminds me
of a 1971 statement by Nobel Laureate Albert Szent-Gyorgyi (biochemistry):
"If you know in advance what you are going to do, or even
to find there, then it is not research at all: then it is only
a kind of honorable occupation..."
What do beekeepers think? What
percentage of them accept the assumption that one can grab a
bee, attach something to its body, release it, and then expect
it to go on its way as if nothing had happened? In my experience,
such a bee would fly an escape path and would not likely continue
a presumably "programmed" behavior. Those interested
in what recruited bees normally might do after leaving the hive,
including search for an odor source can access:
http://www.beesource.com/pov/wenner/az1991.htm
That publication of ours contains
several quotations by von Frisch that stand in sharp contrast
to the observations and claims made in this latest Nature
publication.
Some other questions might
have occurred by now to beekeepers and bee researchers:
1) What about the very small
sample size - can the behavior of 19 disturbed bees undermine
the results obtained from hundreds of unrestrained bees searching
for odor sources in our double controlled and strong inference
experiments? (And, note, only two bees showed up near the target
station in the radar tracking experiment after some unspecified
time delay.)
2) What about the behavior of the many other bees they must have
experimented upon. Did they only gather data from fewer than
two dozen bees and then quit when they had supportive evidence?
Tracking recruited bees with
radar to study their flight paths after leaving the hive is a
noble goal, provided the bees have not been disturbed.
Neither do results obtained without the use of odor have relevance
to the language hypothesis (as pointed out by Rosin, above).
Also, the experimenters should not bother to conduct research
unless they can set aside any vested interest in the outcome;
that is, that they should instead execute a true test
of the language hypothesis and not attempt only to seek confirmation
of their prior entrenched beliefs.
As is usual in such a case,
some reporters contacted me for my impressions of the Nature
report. David Perlman of the San Francisco Chronicle
recognized that I had not had time to study the publication and
came out with a fairly accurate assessment in his article. Other
reporters would not accept my explanation that I would need a
few days to study the original publication, with a rather common
comment: "I have a deadline to meet."
No, I don't think that the
behavior of 19 bees will resolve this controversy, one that has
run now for well more than three decades. Nor will beekeepers
benefit from the claims made in this latest effort. They can
go back to worrying about varroa mites, small hive beetles, and
other real problems with, hopefully, some real solutions appearing
in the near future. Consider the money and time spent on this
supposedly final solution to the bee language controversy!
Wouldn't that time and effort have been better spent on breeding
a varroa resistant bee, for example?
Despite assertions by the radar
tracking crew, the exotic bee language controversy will remain
with us for quite some time to come. As stated at the outset,
the controversy does not revolve about evidence with respect
to validity of the hypothesis. The dance language hypothesis
fails to account for all the available evidence with respect
to the fundamental question: "How does a naïve bee
find a target food source?
Bee language advocates should
now address the really important question (how do they find food)
instead of attempting to prove a hypothesis. The favored hypothesis
continues to be elusive. Until they address the full role of
odor as first elucidated by von Frisch, we will continue at a
stalemate in the controversy (see: http://www.beesource.com/pov/wenner/jib2002.htm).
In summary, did radar tracking of bee flight paths resolve the
bee language controversy? No, it did not; indeed, in the protocol
recently reported in Nature, it could not. Because now, just
as for the past thirty plus years, a statement that Patrick Wells
and I published in Nature back in 1973 (pages 171-175)
still holds true ". . . the honey bee forager recruitment
controversy is not about the nature of evidence but rather about
the nature of hypotheses. It is not what investigators observe
(the data) but what they believe (infer) that is at the heart
of the controversy." See: http://www.beesource.com/pov/wenner/natjan1973.htm
Adrian M. Wenner
967 Garcia Road
Santa Barbara CA 93103
wenner@lifesci.ucsb.edu
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