AMER. ZOOL., 29:1177-1197 (1989)
Concept-Centered versus Organism-Centered Biology 1
ADRIAN M. WENNER
Department of Biological Sciences,
University of California, Santa Barbara,
Santa Barbara, California 93106
SYNOPSIS. Controversies occur frequently, are unavoidable, and probably represent an
important component of biological research. Ironically, emerging empirical evidence appears
to contribute less to the development of controversy than the fierce loyalty with which
many biologists adhere to some traditional methodology and/or outlook in their scientific
discipline. Not infrequently, prevailing attitudes inhibit biologists from recognizing that
a new experimental protocol might provide an appropriate means for testing an established
hypothesis.
Lack of consensus about the suitability of different experimental approaches can also
help explain why scientific manuscripts and grant proposals which support existing theory
and which have employed traditional approaches encounter less difficulty during the
review process than do those manuscripts or proposals which contest established hypotheses or which offer alternative hypotheses and approaches. That is true even if the conclusions drawn in supportive manuscripts and proposals are not parsimonious. When a
biologist's allegiance to an established hypothesis and/or when a traditional methodology
becomes the overriding concern in research, new empirical evidence emerging from
careful study of living organisms becomes relatively unimportant.
I provide here some examples of biological controversies and disparities that stemmed
from adherence to prevailing hypotheses. The biologists involved either did not recognize
that favored hypotheses lacked confirming empirical evidence or that those hypotheses
could not account for important new evidence.
INTRODUCTION
We have to understand first how many
elements can be brought to bear on a
controversy; once this is understood, the
other problems will be easier to solve.
(Latour, 1987, p. 62)
Organisms are necessary; without them
there would be no biology. Beyond that,
organisms are often ultimately responsible
for the resolution of conflict when interpretations differ about our perception of
nature.
Differing interpretations, in turn, stem
from a diversity in our backgrounds. When
provided the same set of facts during a controversy, participants can form quite different sets of conclusions, as Weimer (1979,
p. 66) indicated: "Testing cannot separate
two paradigms—both will be supported in
places, refuted in others. . . ." That condition arises because there is always a difference in philosophical framework of par1
From the Symposium on Is the Organism Necessary?
presented at the annual meeting of the American
Society of Zoologists, 27-30 December 1987, at New
Orleans, Louisiana.
ticipants and concomitant disagreement
about appropriate methodology; and,
despite disclaimers, we all work within some
"paradigm" or other (Kuhn, 1970a) and
practice specific methodologies. Consequently, in concept-driven research on
organisms, biology may become secondary
when theory is threatened.
Just how much attention do biologists
generally pay to philosophical, methodological, and sociological considerations
during the course of their research? Many
would likely agree with Alan Chalmers'
(1982, p. 168) cynical assessment: ". . . philosophy or methodology of science is of no
help to scientists."
Others would align themselves more
closely with Thomas Kuhn's (1970a, p. 94)
position: " . . . this issue of paradigm choice
can never be unequivocally settled by logic
and experiment alone. . . ." Clearly, we
have a fundamental discrepancy before us.
Can these views be reconciled?
Despite the fact that new young scientists
generally lack any background in philosophy and methodology, some people consider the products of our graduate schools
to be well prepared for their research
1177
1178
ADRIAN M. WENNER
careers. Kneller (1978, p. 117) was confidently optimistic on that point:
. . . the scientist generally tries harder
than the layman to screen out personal
prejudice and check for possible error.
He seeks to make his assumptions explicit
and attends to the work of others in his
field. He reports his findings more accurately and makes predictions that can in
principle be tested precisely.
Instead of formal courses in philosophy,
sociology, or psychology of science, however, a graduate student "apprenticeship
training" program is usually the only preparation provided in scientific method. One
might insist that such undergraduate and
graduate training of those who become
biologists provides sufficient influence for
their later mental orientation and methodological "style" as professional scientists. Theocharis and Psimopoulos (1987,
p. 597) questioned that attitude, as follows:
One may wonder how many universities in the world give their science students compulsory formal courses of lectures on the rigours of the scientific
method. . . . [Instead] the hapless student is inevitably left to his or her own
devices to pick up casually and randomly,
from here and there, unorganized bits
of the scientific method, as well as bits
of wnscientific methods.
Scientists thus may have a reputation
(Kneller, 1978) and often may have a selfimage as objective searchers for truth.
Sociologists and psychologists of science
have gained quite a different perception;
according to them, scientists may react just
as subjectively as laypersons when interpretations are proposed which differ from
their own views (Mahoney, 1976). The
influence of "coherent social groups" (Griffith and Mullins, 1972) also controls the
rate at which new ideas may become
accepted in the scientific community.
As an example, Mahoney studied the
reaction of anomymous referees to pseudoscientific manuscripts provided them
(with the assistance of cooperating journal
editors). In two versions of those manuscripts the experimental results provided
referees were identical, but the conclusions
either meshed with the prevailing paradigm or countered that bias. Mahoney
found that referees overwhelmingly advocated publication for manuscripts which
supported established thought and generally recommended denial for manuscripts which had the opposite conclusions.
Philosophers of science also disagree
markedly with one another (see Lakatos
and Musgrave, 1970; Theocharis and Psimopoulos, 1987) when they propose reasons for the success enjoyed by science these
past few hundred years. It is no wonder
that any biologist delving into writings of
non-biologists on the alleged "process" of
science (i.e., "the scientific method") might
become confused when reading such
accounts.
Patrick Wells and I found that the diverse
viewpoints of various philosophers of science could, in fact, be compared to one
another (Wenner and Wells, 1990). We
devised a schematic diagram (Fig. 1) which
helped us perceive the relationship which
might exist among the several modern
analyses provided by philosophers and their
respective schools. After elaborating on
that diagram, I will present some examples
to illustrate how research on whole organisms can place the interplay between theory and research into perspective.
PHILOSOPHICAL ANALYSES:
THE NATURE OF SCIENTIFIC METHOD
There are apparently two major schools
in the philosophy of science, as exemplified
by Bernstein's terms (1983), "objectivism"
and "relativism." However, "realism" and
"anti-realism" are terms used by others for
the same two categories (Bernstein, 1983,
p. 1). Since "anti-realism" has confused
many people and since "objectivism" is not
the most common expression in the literature, we have juxtaposed realism and relativism in the left and right halves of the
diagram, respectively. Most of the different emphases described or advocated by
philosophers of science then fit in one of
the four corners.
The diagram itself can be explained by
proceeding around it in a clockwise direction, beginning in the lower left hand cor-
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
1179
THOMAS KUHN
(Paradigm Shift)
THOMAS CHAMBERLIN
JOHN PLATT
(Infer)
KARL POPPER
(Falsify)
REALISM VS. RELATIVISM
J
PAUL
FEYERABEND
RUDOLF CARNAP
(Verify)
MICHAEL POLANYI
(Crossing a Logical Gap)
JAMES ATKINSON
(Explore)
FIG. 1. Suggested scheme to illustrate the relationship among different philosophers of science with regard
to their views about "The Scientific Method." Both Popper and Carnap ("Realism" or "Objectivism School")
advocated an "objective" search for "truth" ("logical empiricism") but differed in the approach they advocated
to achieve that end.
Kuhn challenged whether science can proceed in the manner advocated by the logical impiricists and
described the neglected importance of sudden "paradigm shifts." An accumulation of incompatible evidence
(anomalies) permits someone in the community to experience such a paradigm shift; a crisis or period of
uncertainty results until a sufficient number of scientists can accommodate the new viewpoint.
Those in the "Relativism School" do not advocate a search for "ultimate truth" but emphasize the need
to gain useful knowledge which will permit the design of new experiments. Chamberlin and Platt argued for
the application of "multiple inference" or "strong inference" in experimental design. Atkinson argued that
we actually only "create" phenomena in our minds as we explore nature and then hope to "convert" others
to our viewpoint when we feel we understand the new principle. Feyerabend militantly argued that the
application of many techniques is involved in our search for knowledge in science ("anything goes").
ner with Rudolf Carnap's contribution, out one explanation for observed events
verification of hypotheses. (However, the can be unsettling. Atkinson (1985, p. 732)
diagram should be viewed as a dynamic phrased the problem as follows:
summary. A scientist or philosopher locked
Image manipulation is essential for the
into one corner on a given issue at a given
highly
successful creative function of the
time may be in quite another corner on a
mind;
yet
a chaotic, totally free-flowing
different issue or at a different stage of
interaction
of images would make human
theory development.)
communication and cooperation impossible by creating as many worlds as there
Confirmation (verification): Analysis of
are people or [more] for each person than
Rudolf Carnap
they could possibly handle.
Every human being has a strong tendency to form a single hypothesis about an
As a consequence, we tend to focus on
observed event whenever a small amount a single explanation, seek confirmation
of evidence appears in support of a previ- (verification) of our belief in that hypothously held notion. In fact, a failure to single esis, and thereby further strengthen our
1180
ADRIAN M. WENNER
biases. Scientists within a subdiscipline of
biology, for example, may reach consensus
that a hypothesis, even though inadequately
tested, can be accepted as a close approximation to the truth. It is then that the
"necessity of the organism" may fade by
comparison.
Carnap, one of the logical empiricists,
justified and formalized the neoclassical
empiricism approach he advocated (i.e., the
value of searching for confirming evidence). Lakatos (1968, pp. 322-323) summarized Carnap's position as follows:
[Carnap's school] approached the
problem from the classical point of view
of the logic of justification. Since it was
clear that theories could not be classified
as probably true or false, they had . . .
to be classified as at least "partially
proved," or in other words, as "confirmed (by facts) to a certain degree." It
was thought that this "degree of evidential support" or "degree of confirmation" should somehow be equated with
probability in the sense of the probability
calculus.
Falsification: Analysis of Karl Popper
As early as 1919 Karl Popper had wondered, "When should a theory be ranked
as scientific?" That and other considerations eventually led him to a number of
conclusions which he reformulated in 1957
(Popper, 1957, pp. 159, 160). Among them
were: "Confirmations should count only if
they are the result of risky predictions. . . ."
and "Confirming evidence does not count
except when it is the result of a genuine test of
the theory. . . . " He also concluded (1957, p.
160):
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.
Thus it was that Popper, also a logical
empiricist, criticized Carnap's 1950 theory
of confirmation (Lakatos, 1968, p. 330) and
held fast to critical empiricism. Although
Rudolf Carnap and Karl Popper were
obviously at odds with one another with
regard to the emphasis they deemed most
successful (Michalos, 1971), both apparently felt that truth was still approachable.
Sufficient verification (Carnap) or an
inability to falsify (Popper) would eventually permit one no longer to need to question a theory or hypothesis (Fig. 2). (In
later years Popper altered his views somewhat. Imre Lakatos then referred to him
as Popper! and Popper2; see Chapter 6 in
Weimer, 1979.)
A little appreciated fact is that Popper
was pre-empted by others, notably Sir
Francis Bacon, who wrote (1620, p. 128):
In forming axioms, we must invent a
different form of induction from that
hitherto in use. . . . The induction which
proceeds by simple enumeration is puerile, leads to uncertain conclusions, and
is exposed to danger from one contradictory instance. . . . a really useful
induction . . . should separate nature by
proper rejections and exclusions, and
then conclude for the affirmative, after
collecting a sufficient number of negatives.
The analyses provided by Carnap and
Popper (and Bacon) fit what can be called
either the Realism (Leplin, 1984) or the
Objectivism (Bernstein, 1983) School (left
side of the diagram in Fig. 1). That is,
"objective t r u t h " exists and can be
approached by careful experimentation.
For many biologists, one or the other
approaches (verification or falsification)
apparently constitutes " T h e Scientific
Method" (as in Silvernale, 1965, pp. 4, 5).
Paradigm shift: Analysis of Thomas Kuhn
As indicated, both Carnap and Popper
analyzed scientific methodology largely
within the context of logical empiricism.
In his influential book, The Structure of Scientific Revolutions (1962, reprinted in 1970),
Kuhn (1970a) provided an alternative
interpretation about scientific methodology, an interpretation which continues to
appeal strongly to both scientists and many
non-scientists. Kuhn suggested that, when
a given hypothesis is entrenched in science
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
1181
100%
PARADIGM HOLD
I
Z)
cc
LL
O
_J
LU
LU
0%
EFFORT
7
FIG. 2. Proposed scheme for those who adhere to the philosophy of the Objectivism School. Under either
the falsification (null hypothesis) or the verification approaches, continued experimentation leads to an ever
greater appreciation of what might be "true" in the system under study. There exists a point during accumulation of knowledge where members of the scientific community may feel a given hypothesis may no longer
be doubted; Kuhn referred to activity above that point as a "Paradigm Hold."
("paradigm hold"—Fig. 2), resistance to
any new idea is almost certain to prevail,
even if the new interpretation has a solid
empirical basis. Only a fraction of scientists
within the scientific community is then willing to entertain the new interpretation; a
majority of the scientists resist because of
strong "paradigm holds." A crisis will
almost surely follow, as a rift develops on
that issue between those who feel the issue
had been resolved earlier (realists) and
those who recognize that alternatives
should still be considered (relativists).
Kuhn, however, did not use those terms in
his book.
To illustrate his thesis, Kuhn traced
examples of change in the scientific community during some familiar paradigm
shifts (his term). He indicated that those
transitions between earlier prevailing
hypotheses and new hypotheses had not
occurred in the calm, dispassionate
("objective") manner traditionally considered characteristic of scientists (Silvernale,
1965; Kneller, 1978; Theocharis and Psimopoulos, 1987).
Kuhn provided terms for the distinction
he perceived between non-controversial
and controversial science ("normal science" and "crisis resolution," respectively). In doing so, he was thus primarily
describing the actual practice of science, both
during day-to-day research and during
important paradigm shifts. His goal was
thereby different from that of analyzing
methodologies of successful scientists, as
had been done by objective empiricists such
as Carnap and Popper.
Kuhn's ideas swept through various scientific fields and have been accepted by
much of the scientific community, even
while apparently escaping the attention of
a great many biologists. He may be considered responsible, nonetheless, for institut-
1182
ADRIAN M. WENNER
ing a "paradigm shift" among philosophers of science and scientists—from a
consideration of how scientists ought to
employ method to how scientists do employ
method (see also Masterson, 1970).
Whereas our human nature meshes well
with the intuitive notion that a real world
exists out there and can be "discovered"
or "proved to be true" by the methods
advocated by those in the Realism School,
Kuhn stressed the overriding importance
of anomalies which constantly arise. Eventually some brave (or foolish) scientist pursues the anomalous result and perceives an
entirely different way by which that particular aspect of the world can be described.
Convincing others that the new interpretation fits the facts better than the old
("conversion") can be quite another matter.
The experience of Nobel Prize winner
Barbara McClintock, who studied corn
genetics provides a useful example of resistance to new ideas (see also below). Evelyn
Fox Keller graphically
described
McClintock's Kuhnian experience with the
scientific community. Keller (1983, p. xii)
wrote:
Scientific knowledge as a whole grows
out of the interaction . . . between individual creativity and communal validation. But sometimes that interaction miscarries, and an estrangement occurs
between individual and community.
Usually in such a case, the scientist loses
credibility. But should that not happen,
or, even better, should it happen and
then be reversed, we have a special
opportunity to understand the meaning
of dissent in science.
Kuhn's ideas are interesting but, alas,
they are much too vague to give rise to
anything but lots of hot air. . . . Never
before has the literature on the philosophy of science been invaded by so many
creeps and incompetents.
Kuhn, in his contribution, "Logic of discovery or psychology of research?" suggested that the disparity in thought between
traditional philosophers and himself might
itself be due to different starting points for
them and for him (a difference in paradigm
hold). He wrote (1970*, p. 3):
Sir Karl and I do appeal to the same
data; to an uncommon extent we are
seeing the same lines on the same paper;
asked about those lines and those data,
we often give virtually identical
responses, or at least responses that inevitably seem identical in the isolation
enforced by the question-and-answer
mode. . . . Though the lines are the same,
the figures which emerge from them are
not. That is why I call what separates us
a gestalt switch rather than a disagreement and also why I am at once perplexed and intrigued about how best to
explore the separation.
Kuhn continued with a cogent statement
summarizing the difference in frames of
reference between Popper and himself:
How am I to persuade Sir Karl. . . that
what he calls a duck can be seen as a
rabbit? How am I to show him what it
would be like to wear my spectacles when
he has already learned to look at everything I can point to through his own?
Perhaps the most important result of
Kuhn's
book has been the emergence of a
Kuhn's arguments incurred the wrath of
fuller
awareness
that Realism and Relativtraditional philosophers (see Lakatos and
ism
schools
exist
and have apparently
Musgrave, 1970; Feyerabend, 1981). By his
existed
throughout
centuries
under a varisuggestion that science proceeds in a mannames.
Bernstein
(1983,
p. 1) wrote:
ety
of
ner other than that specified by the traditionalists, Kuhn had inadvertently gen[There is an uneasiness which] affects
erated his own "revolution" of thought.
almost every discipline and every aspect
The philosophers themselves did not
of our lives. This uneasiness is expressed
behave in a calm, dispassionate manner
by the opposition between objectivism
toward Kuhn's analysis. Feyerabend (1981,
and relativism, but there are a variety of
p. 160), whose objections were perhaps the
other contrasts that indicate the same
most pointed, wrote:
underlying anxiety: rationality versus
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
irrationality, objectivity versus subjectivity, realism versus antirealism. . . . Even
the attempts that some have made to
break out of this framework of thinking
have all too frequently been assimilated
to these standard oppositions.
Michael Polanyi (1958) actually preempted Kuhn with respect to a part of
Kuhn's interpretation by his use of a phrase,
"crossing a logical gap," which apparently
had much the same meaning as Kuhn's
"paradigm shift." Polanyi's contribution,
however, did not have the same impact on
the scientific community as Kuhn's "paradigm shift" or "gestalt switch." Once
again, we recognize the difficulty one may
encounter in "converting" others to a new
point of view.
Polanyi's name has been placed on the
diagram (Fig. 1) in a position approximately equivalent to the location of Kuhn's
name, in recognition of Polanyi's priority
in the concept of change in mind set when
evidence is suddenly perceived in a different manner.
Multiple inference: Thomas Chamberlin's
analysis
Thomas Crowder Chamberlin, the
"Dean of American Geologists" at the turn
of the century, pre-empted many contemporary philosophical considerations in a
now-classic paper, "The method of multiple working hypotheses" (1890; reprinted
in 1965). He distinguished between 3 classes
of experimental approach: 1) The Method
of the Ruling Theory, 2) The Method of
the Working Hypothesis, and 3) The
Method of Multiple [Working] Hypotheses.
John R. Platt, writing on "Strong Inference" in 1964, was responsible for the
reprinting of Chamberlin's paper in the
journal Science (1965). Chamberlin's contribution has since been experiencing an
ever-widening impact within the scientific
community. Some of Chamberlin's words
are particularly important to the theme of
the present symposium.
Let us consider each of the three kinds
of scientific investigation process, as classified by Chamberlin. In turn, let us reflect
1183
on how biological scientists may thereby
view the "necessity" of the organism.
1) The ruling theory. When a ruling theory becomes dominant within a research
field, especially during the "normal science" activity of Kuhn, experimental
results may be accepted only within the
limits permitted by a particular paradigm.
Confirming evidence (verification) counts
for much in that approach, and experimental results at variance with the prevailing paradigm may not even be noticed
(see Bruner and Postman, 1949). Chamberlin wrote (1965, p. 755):
Briefly summed up, the evolution is
this: a premature explanation passes into
a tentative theory, then into an adopted
theory, and then into a ruling theory.
When the last stage has been reached,
unless the theory happens, perchance, to
be the true one, all hope of the best
results is gone.
2) The method of the working hypothesis.
Chamberlin had recognized a distinction
(between verification and falsification,
respectively) which was noted considerably
later by Carnap and Popper. Chamberlin
(1965, p. 755) wrote:
Under the method of the ruling theory, the stimulus was directed to the finding of facts for the support of the theory.
Under the working hypothesis, the facts
are sought for the purpose of ultimate
induction and demonstration, the
hypothesis being but a means. . . .
A fundamental weakness of that "Working Hypothesis" approach did not escape
Chamberlin. He wrote(1965,pp. 755,756):
It will be observed that the distinction
is not a sharp one, and that a working
hypothesis may with the utmost ease
degenerate into a ruling theory. Affection may as easily cling about an hypothesis as about a theory, and the demonstration of the one may become a ruling
passion as much as of the other.
3) Method of the multiple hypotheses.
Chamberlin's important contribution was
recognized by Platt (1964), while the latter
puzzled over the great differences he
1184
ADRIAN M. WENNER
observed in rate of progress in different not have a "language" (1901, pp. 162-167;
fields of science. After asking, "Why should see also below).
there be such rapid advances in some fields
and not in others?" Platt continued (1964, Exploration: Analysis ofJames Atkinson
p. 347):
Atkinson (1985) proposed that the process of science is a creative enterprise, in
I think the usual explanations that we which we mentally manipulate images
tend to think of—such as the tractability about what may happen on the basis of what
of the subject, or the quality or educa- we have seen. After "creating an image"
tion of the men drawn into it, or the size or interpretation (explanation or hypothof research contracts—are important but esis), we attempt to "convert" others to our
inadequate. I have begun to believe that point of view. If we succeed in convincing
. . . rapidly moving fields are fields where others, that success reinforces our own
a particular method of doing scientific commitment to the hypothesis we have
research is systematically used and generated. In the initial stages of this image
taught, an accumulative method of creation, it is most often (but not always)
inductive inference. . . .
the biology of the organism which provides
As indicated above, Platt advocated what the necessary input during generation of
he called "Strong Inference," whereby two our hypothesis.
hypotheses are pitted against one another
In some ways, Atkinson's deliberations
during experimentation. A repetition of resemble the concept of "personal knowlthat process when new results are obtained edge" as emphasized by Polanyi (1958), in
provides a "logical tree" approach and an which "objectivity" plays little part. Polanyi
exponential progress in the search for distinguished between "heuristic passion"
understanding.
and "persuasive passion." The former term
By contrast, Chamberlin had advocated applies to the practice of basic research for
pitting several hypotheses against one personal satisfaction and the thrill of
another (multiple inference) in order to understanding; the latter term applies to
avoid forming a "parental attachment" to one's attempt to "convert" others (Atkinany one explanation. As many sociological son, 1985) to the notion that understandand psychological studies of science reveal, ing has been achieved.
what we observe in studies of biology often
Scientific progress occurring as a result
depends on what our background permits of our new image (feasible or infeasible)
us to see (concept-dominated observation). may not be rapid, however. That is particIs the organism then necessary? Chamber- ularly likely to be the case if the image we
lin would have believed so; he wrote (1965, have created conflicts with other well
p. 759):
established images or if our image falls too
far outside the ability of others to perceive
Under the method of multiple hypoth- what we mean. One of Atkinson's exameses, it is the first effort of the mind to ples of the latter case was the experience
see truly what the act is, unbeclouded by of Barbara McClintock mentioned above
the presumption that this or that has been (Keller, 1983; see also below).
done because it accords with our ruling
If no complications exist or if no serious
theory or our working hypothesis.
objections are raised to the new image creChamberlin's philosophy actually may ated, Atkinson suggested, nearly all within
have reflected a common thought process the entire scientific community can accept
in his time. Sir Arthur Conan Doyle's the validity of the new hypothesis through
famous detective, Sherlock Holmes, fre- a "gestalt switch" or "conversion experiquently applied the multiple inference ence." Atkinson thus recognized the "tenapproach. Another example is that of tative and uncertain" nature of scientific
Nobel Prize winner Maurice Maeterlinck; knowledge. We do not necessarily get closer
he conducted a strong inference experi- to the truth as we proceed in our research;
ment and concluded that honey bees did we merely explore reasonable alternatives.
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
If we succumb too readily to "persuasive
passion" (ours or that of others) we may
accept a hypothesis which lacks sufficient
basis.
"Anything goes": Analysis of Paul Feyerabend
Feyerabend (1970, 1975, 1978) is a special case and is consequently enclosed within
an ellipse in the diagram (Fig. 1). He militantly insisted that no one approach should
be advocated in scientific research and
argued that adherence to the deliberations
of traditional philosophers of science could
actually dampen free scientific inquiry.
Furthermore, he considered the advocacy
of one method over another by other philosophers (such as the logical empiricists)
as intolerable incursions on that inquiry
and recognized the socio-political forces at
work within science. As an example, Feyerabend (1975, p. 43) wrote:
Next, [a scientific] development
becomes known to the public. Popular
science books . . . spread the basic postulates of the theory; applications are
made in distant fields, money is given to
the orthodox, and is withheld from the
rebels. More than ever the theory seems
to possess tremendous empirical support. . . . this appearance of success can-
1185
"antithesis" of Theocharis and Psimopoulos(1987).
Thus it is that we have come full circle
in the schematic diagram (Fig. 1). Quite
disparate approaches are used by biologists
in their research programs. In that diagrammatic scheme, the most rapid advances
occur for those who work within the Relativism School. Commitment to "Ruling
Theories" can most readily blind those who
belong to the Realism School. The breach
between philosophies adhered to within
biology appears to be so great that communication may not even be possible
between scientists in the different schools.
Controversies, which erupt when communication between realists and relativists
becomes no longer possible, begin to break
the stranglehold of theory and most often
surface when the organism has ceased to
be the most important factor under consideration. The value of controversies then
becomes apparent, just as stated by Popper
(1963, p. 316):
Criticism is, in a very important sense,
the main motive force of any intellectual
development. Without contradictions,
without criticism, there would be no
rational motive for changing our theories; there would be no intellectual progress.
not in the least be regarded as a sign of truth
Let us now turn to some specific examand correspondence with nature. . . . the suspicion arises that this alleged success is due ples of research and see how various
to the fact that the theory, when extended approaches have been applied during the
beyond its starting point, was turned into rigid course of investigation.
ideology. . . . Its "success" is entirely man-
made, [emphasis Feyerabend's]
(In this last connection, see Latour, 1987,
pp. 153-155.)
According to Feyerabend, then, a prevailing theory can dominate a field, even
when it no longer accords with nature. The
organism may no longer be necessary when
a theory is "turned into rigid ideology."
One can also recognize in Feyerabend's
writings both the "creation of an image"
and the "conversion" elements described
by Atkinson. Other comments in Feyerabend's writings appear to place him quite
firmly in an extreme position within the
Relativism School, an embodiment of the
INTRADISCIPLINARY CONTROVERSY:
THE INTERPRETATION OF
NATURAL PHENOMENA
As we have seen, philosophical analysts
have characterized modern biologists as
scientists, who ought to or who do do science
from within several different frameworks
("verification," "falsification," "strong
inference," "exploration," etc.). It is not
surprising that intradisciplinary problems
can arise when a particular field of biological research is dominated by scientists who
are strongly imbued with only one orientation.
Let us now consider several instances in
which a prevailing attitude toward the way
1186
ADRIAN M. WENNER
sented by the genus Cephenemyia [deer
in which evidence is sought and utilized
bot flies] of North America and Europe
within a discipline can actually distort the
. . . evidently hold the world's record for
scientific enterprise and can interfere with
high speed in flight. This is the more
efforts to understand the living organism.
strange since these flies take no nourishAs will be evident, some of the controverment whatever in the [adult] stage.
sial examples I have chosen exemplify the
interference one can encounter. A great
many other instances have occurred in In that same year the editor of The Scientific
biology and could have been used except Monthly indicated that "Several corresponfor the fact that they have not appeared in dents have written concerning the artiprint. While some examples may seem triv- cle. . .," but that journal published only one
ial now, they did not appear so at the time. paper, which was highly supportive of
Conversely, while some seem non-trivial Townsend's article (Haight, 1926).
A year later (1927) Townsend responded
now, they may well appear to be trivial at
to a comment by a Mr. Howard S. Rapsome time in the future.
pleye, of the U.S. Coast and Geodetic SurIt is with studies of whole organisms that vey. Rappleye indicated that Townsend and
the relationship between theory and Haight's calculations had been in error, and
research can become most apparent. How- that a speed less than 818 miles per hour
ever, lest some of those examples appear at the 40th parallel would be sufficient to
to cast aspersions on any particular subfield circle the world in a day. Townsend
of biology, I hasten to add that such exam- exploited this indirect support of his
ples are legion in other areas of science. hypothesis "to call attention to the peculiar
For example, physical chemistry had its structure of the fly wing" and how that
"polywater" (Franks, 1981), physics its "N- wing structure could make such fast speed
rays" (Nye, 1980), cancer research its possible.
"kinase cascade" (Broad and Wade, 1982),
The fly's "world record" for animal
plant ecology its 'individualistic and community-unit concepts" (Shipley and Keddy, speed persisted in the literature for more
1987), and bioenergetics its long-term than a decade until Nobel Prize-winning
resistance to the notion of chemiosmotic physicist, Irving Langmuir, published a
coupling in oxidative and photosynthetic short technical comment in Science (a AAAS
phosphorylation (Gilbert and Mulkay, companion publication to The Scientific
Monthly). Townsend's reported flight speed
1984).
of deer bot flies had seemed to Langmuir
(1938) to be: ". . . so fantastically high . . .
"Around the world in a daylight day"
that the fly would have to consume his own
At times, a scientific community can eas- weight of fuel every few minutes or so to
ily be "converted" to belief in an exotic supply the necessary power." Langmuir had
notion which even laymen may question. made "some rough mental calculations"
That seems to be the case for the "world's and concluded that Townsend was in error
record for high speed in flight." Published but pursued the topic no further at that
charts in the 1930s and 1940s indicated time.
that male "deer flies" (Cephenemyia), with
Langmuir, however, did act when he saw
their speed of 818 mph, outperformed all
other animals; female flies were only slightly a "2-page diagram in the Illustrated London
News, January 1, 1938." (This was apparslower, at 614 mph.
ently the same diagram I saw posted in our
That claim had an early inception. In campus vertebrate museum in 1960.)
1926, C. H. T. Townsend published a paper Because this claim was "going the rounds
in The Scientific Monthly (a AAAS publica- over the whole country," Langmuir felt
tion) with the title, "Around the world in compelled to publish his technical coma daylight day: A problem in flight." In his ment. In that comment he provided the
article he wrote (p. 309):
reasons behind his conclusion that "a speed
The flies of a muscoid group, repre- of 25 miles per hour is a reasonable one
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
for the deer fly [Langmuir's designation],
while 800 miles per hour is utterly impossible."
The editor of Science twice refused
Townsend's attempts to reply to Langmuir's comments, but Townsend managed
to get his case into print in the Journal of
the New York Entomological Society (Town-
send, 1939). Townsend provided his own
mathematical calculations and arguments
in support of his earlier hypothesis. While
those arguments might seem fanciful
(although interesting reading) under current standards, Townsend's comments
seem to be what might follow from one
working within the verification orientation
described above.
Was the organism necessary? Townsend
had "created the image" of fly speed on
the basis of field observations and had
"converted" (Atkinson, 1985) many other
scientists (and much of the public) to his
point of view. Langmuir, with the aid of
string, weights, and calculations, concluded that flies could not travel that fast;
and, if they did, they would have been
invisible to Townsend. Apparently neither
Townsend nor Langmuir had any data on
the actual flight speed of those flies.
1187
so far as to suggest that the zoeae
Thompson saw hatch from crabs confined in the laboratory were parasites. . . .
Westwood's persuasive argument, including his use of the above ad hoc "parasite"
hypothesis, resulted in his later being
awarded a gold medal from the Royal Society of London for his complete refutation
of Thompson's "heresy" (Coffin, I960).
Was the organism necessary? In winning
the gold medal, Westwood had not thoroughly studied the organism. He apparently did not conduct a single experiment
with zoeae (rhetoric or "persuasive passion" substituted for science). For his part,
however, Thompson had not raised a single zoea through metamorphosis into the
juvenile form.
Use of landmarks or sun compass by
orienting ants?
Among ants, individual foraging workers (in species of ants which do not follow
trails) are considered capable of using "sun
compass orientation" as they travel
between their nest and food sources (prevailing theory). However, it is also possible
that they could use landmarks or other
stimuli not immediately perceptible to the
The crustacean zoea: A larva or a parasite?
scientists studying the ants' orientation.
John V. Thompson (1831) was the first Supporting evidence for either hypothesis
person known to recognize that transpar- has been obtained by scientists using a realent larval forms (zoeae), which were found ist, "verification" procedure.
in the ocean, could later metamorphose
Wehner and Menzel (1969) studied the
into familiar crabs (Coffin, 1960). He did
paths
of individual ants which had been
not actually see that happen; he concluded
collected
at one point remote from their
it was true by watching zoeae hatch from
nest
and
released
from a point at a particcrab eggs (exploration approach). Thompular
distance
in
an
opposite direction from
son's conclusion, however, did not mesh
their
nest.
However,
a re-analysis of the
with prevailing opinion, since zoeae were
"known" to comprise their own zoological Wehner and Menzel data shows that the
direction travelled by the captured ants
group (the "ruling theory").
from their point of release was actually ranAfter the appearance of Thompson's dom. As an example, one can combine the
papers, J. O. Westwood (1835) "rescued" solid point data, a single release event, from
the prevailing paradigm. As Coffin (1960) their figs. 2d and 2e (Fig. 3 here) into a
wrote:
single figure (Fig. 4.). Statistical analysis
[Westwood] reviewed Thompson's then reveals no difference from random in
work point by point and showed how his the resultant orientation of that group of
observations were contrary to the known ants.
Wehner and Menzel, on the other hand,
methods of metamorphosis and ecdysis
and therefore impossible. He even went worked within the sun compass orientation
1188
ADRIAN M. WENNER
NEST
DIRECTION
NEST
DIRECTION
FIG. 3. The solid dots in the above two figures actually represent performances for a single group of ants
released at one time from a point 10 m from their nest. Those which returned to their nest (left circle) were
a posteriori assumed to have used landmarks as the stimulus during orientation. Those which did not return
to their nest (right hand circle) were assumed, again a posteriori to have used a sun compass orientation
capability as they traveled away from the nest. (Data extracted from figures in Wehner and Menzel, 1969.)
paradigm (Chamberlin's "Ruling Theory") and partitioned their data. There
were ants which returned to their nest (e.g.,
their fig. 2d in Fig. 3 here) and those ants
which did not return directly (e.g., their fig.
2e in Fig. 3 here). Wehner and Menzel's
interpretation thus rested on the assumption that "successful" ants arriving back at
the nest had used "their memory of landmarks." Since the other released ants did
not return directly to the nest, Wehner and
Menzel concluded that those ants had continued on in that direction because they
had used their sun compass ability (as if the
ants were still in a search mode which Wehner and Menzel assumed would have
accounted for ants moving away from the
nest).
Was the organism necessary in their
experiments? Whenever one partitions the
data in a circular distribution into two subcomponents, each sub-component will yield
a statistically significant orientation (Fig.
3). In fact, any random set of data after
such partitioning would yield statistically
"meaningful" results, regardless of the
behavior of the animals involved.
The editors of Science would not publish
a short note in which I pointed out this
weakness in the Wehner and Menzel data
analysis. (One referee insisted I should
repeat the experiments, done in Israel,
before objecting.) In another manuscript,
where one of my students challenged a similar interpretation regarding sun compass
orientation of amphipods (see below), a
referee wrote: "The sun compass orientation hypothesis is not open to question."
The Wehner and Menzel paper thus
continues to be cited in support of a sun
compass orientation capability in ants (e.g.,
Holldobler, 1971; Adler and Phillips,
1985), despite the serious flaw in their data
analysis. Adler and Phillips, for example,
wrote (p. 548):
The work of Wehner and his associates (Wehner and Menzel 1969; Wehner
and Duelli 1971; Wehner 1976) has
demonstrated that the desert ant is an
excellent model system for learning how
invertebrates use celestial (including
polarized light) and other cues in orientation.
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
NEST
DIRECTION
I
FIG. 4. When the data of Figure 3 are recombined
(above), it is apparent that a hypothesis of random
dispersal from the release point was not negated by
the experiment. (An arbitrary separation of data in a
circular distribution will yield statistically significant
"orientation" results for the residuals.)
1189
observer contribute to the mortality of
the males.
Roeder's disclaimer became lost, but not
his conclusion that extirpation of the
suboesophageal ganglion (or decapitation)
removed inhibition of the male's mating
behavior. Decapitation eventually came to
be interpreted as a requisite act for mating,
as expressed by Alcock (1979, p. 155):
"Sooner or later the female reaches back
over her shoulder and begins munching on
the head of the male."
Davis and Liske questioned the veracity
of that tale but went further. They conducted a series of experiments {exploration
approach), with many individuals from several species of mantids (Davis and Liske,
1985; see also Liske and Davis, 1987). While
doing so, they used video cameras to record
behavior (no disturbance by investigators).
In only one case out of dozens of pairings
did the female cannibalize the male. When
that particular female attacked the male,
however, she actually consumed the entire
male and was therefore unable to mate with
him.
Navigation or range adjustment in
monarch butterflies?
Cannibalism of male mantids: Necessary for
mating by females?
When first teaching my animal behavior
course (1964), one example of instinctive
behavior which I used was that a female
mantid apparently could not complete
mating unless she had first bitten the head
off her male partner. After fruitless
attempts to find the experimental results
in support of that notion, I stopped teaching that "fact" (prevailing theory).
This episode apparently represents a case
of embellishment through the retelling of
a tale (Brown, 1986). By "freezing" when
noticed by females during their approach,
males normally manage to get into position, copulate, and escape without being
eaten (Liske and Davis, 1987). In his original account Roeder (1935, p. 205) had
included a disclaimer:
. . . it is felt that the evidence presented will show that artificial conditions
and even the mere presence of an
The mass movement of monarch butterflies in North America has been studied
primarily by biologists who have relied on
the verification approach. Virtually all such
studies have begun with the assumption that
monarchs exhibit a bird-like migration
south in the fall and north in the spring.
Experiments have thus been designed to
provide support for that hypothesis, but
apparently no experiments have been
designed as attempts to falsify the migration hypothesis or to explore alternate
hypotheses by means of the inference
approach. The verification approach in that
instance was epitomized by Williams (1958,
p. 201):
. . . we must continue to try [to mark
and recapture individual butterflies],
whatever the difficulties; for only by
recovery at a distance can we prove that
a single individual really makes the whole
journey. Apart from such evidence, long
distance migration of insects is an infer-
1190
ADRIAN M. WENNER
ential theory, perhaps the only one that
will explain the known fact, but not
"proved" to the unbeliever!
In the western United States, the results
of field studies of monarch butterflies
revealed that an alternate hypothesis might
suffice, one which does not require any long
distance navigational capabilities on the
part on monarchs (Wenner and Harris,
1989). That is, if monarchs expand their
range generation by generation while flying
in all directions in the spring (after leaving
their overwintering aggregation sites along
the California coastline), they will eventually repopulate the western states. Conversely, if monarch adults produced in California fly primarily against the wind in the
fall, they will again almost all end up overwintering somewhere along the California
coastline, due to the direction of prevailing
winds there during the fall.
The above alternative hypothesis proposed for monarch butterfly movements in
the western states encountered strong
opposition during the Second International Conference on Monarch Butterflies
held in Los Angeles (September 1986),
even though a similar hypothesis had been
independently proposed earlier for monarch population movements in Australia
(Smithers, 1977).
Moon compass or slope orientation in
beachhoppers?
When released on the sand, beachhoppers (Family Talitridae) may hop toward
the ocean. On the other hand they may
not, as the students in my animal behavior
classes observed directly. Since a community of researchers (reviewed in Hartwick,
1976) had previously "demonstrated" that
beachhoppers would hop toward the ocean
"in order to avoid desiccation," my students were unprepared for the results
observed.
Is the organism necessary? The typical
student reaction when confronted with
their own "anomalous" results (the beachhoppers hopped away from the ocean for
them rather than toward it as the prevailing theory indicated they should have done)
was: "What have we done wrong?" The
students could not perceive that the prevailing hypothesis could be in error.
As a result of a series of experiments,
Craig (1971, 1973a, b) suggested an alternate hypothesis: "Beachhoppers primarily
react to slope rather than to sun compass
or moon compass cues as they move up and
down the beach." That set of results leads
one to ask, "Which of the approaches
shown in Figure 1 had been used in the
original experiments?" The answer to that
appears in published comments, some of
which follow.
1) Ercolini and Scapini (1972, p. 75):
"Solar and lunar astronomical orientation
of littoral Amphipods . . . has been demonstrated in an extensive series of experiments [by Papi and Pardi]. . . ."
2) After repeating Craig's experiments
on slope orientation of beachhoppers,
however, Ercolini and Scapini (1974, p.
108) altered their conclusions, as follows:
"Our data on the number of Talitri which
go in opposite directions at the various test
gradients essentially agree with Craig's on
Orchestoidea."
3) Hartwick (1976) later wrote: "When
attempting to demonstrate that an animal
orients by means of specific stimuli. . . ."
One can recognize a willingness to
change an experimental approach in comments by Ercolini and Scapini (from verification to falsification) and a steadfast
adherence to the verification approach in
Hartwick's choice of words. Hartwick's
subsequent experimental results led him to
conclude that beachhoppers could likely
use any of a variety of orientation mechanisms as they move about on the beach.
According to him, beachhoppers may use
either landforms, slope angle, or compass
orientation, according to the circumstances. Hartwick (1976, p. 457) concluded: "Further research may, hopefully,
flesh out this basic skeleton of the complex
hierarchy of orientation strategies in talitrid amphipods."
Hartwick's compound interpretation
(instead of multiple inference approach)
permitted him to retain the earlier prevailing compass orientation hypothesis
("ruling theory"), as well as to accommodate, ad hoc, those experimental results not
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
in agreement with that hypothesis. Is the
organism then any longer necessary? In
reply to that question, consider the specific
case at hand. Under Hartwick's composite
interpretation, what can one expect beachhoppers to do when they are released on
the beach?
Honey bee recruitment to food sources:
Special or commonplace?
Are honey bees really capable of humanlike communication (language)? Those who
still accept the honey bee dance language
hypothesis believe so, but that question has
had a long history (Wenner and Wells,
1990). Aristotle concluded that recruited
bees simply followed foragers as they flew
back out to the food source (see Wenner
and Wells, 1990). In the centuries which
followed, one hypothesis or the other prevailed at different times; some naturalists
agreed with Aristotle, others pressed the
case for a language.
Maeterlinck (1901) ended the debate for
a time. He devised and conducted a series
of experiments with an approach and attitude resembling that of strong inference
(Chamberlin, 1890; Platt, 1964). In his
experiments Maeterlinck found that
recruited bees failed to find the food source
unless the forager bee was allowed to leave
the hive at the same time as the recruits.
His results apparently dampened any serious consideration of a "language" among
honey bees for more than four decades.
What did more recent proponents of the
dance language hypothesis do about Maeterlinck's results? They either ignored them
(Francon, 1938), dismissed them (von
Frisch, 1954), trivialized them (Ardrey,
1963; Ardrey in Marais, 1969), or misrepresented them (Gould, 1976).
My colleagues and I, in a series of experiments (Wenner, 1974; Wenner and Wells,
1990), tested the bee language hypothesis.
When we did so, that hypothesis failed to
explain the results we obtained. In addition
to conducting double controlled experiments (Johnson, 1967; Wenner, 1967), we
used a strong inference design (Platt, 1964;
Wenner et al., 1969), in which recruited
bees could use either the "language" information provided by foragers or they could
1191
use the food odor which had not been used
since the previous day. In either of those
types of experiments, when additional controls were inserted into von Frisch's original experimental design, the results
obtained no longer agreed with expectations of the von Frisch hypothesis.
Our results were rejected at the time
largely, as we view it now, because of an
overriding commitment by others to the
verification approach ("Ruling Theory" of
Chamberlin). Our use of a strong inference
approach impressed few biologists; it simply led to a breakdown of communication.
Proponents of the language hypothesis
could not fathom the need for such experiments, since sufficient evidence supportive
of the language hypothesis was already
available.
In starting a research project, there is a
strong likelihood that the scientist will
adopt a particular attitude (perhaps unwittingly). When Gould et al. (1970) set about
to "resolve" the honey bee dance language
controversy, they again resorted to the verification approach, as indicated in their
opening statement (p. 544):
Simply demonstrating that olfactory
cues are sufficient in a particular situation does not mean that the dance language is not used under other conditions,
[emphasis added]
Within the same sentence one can recognize the rejection of Popper's "falsification" approach and commitment to the
"Ruling Theory" approach even before
they proceeded with their experiments.
Richard Dawkins (1969) and Edward O.
Wilson typified that same attitude ("Ruling
Theory" or verification approach) toward
our experiments and results. For example,
Dawkins (1969, p. 751) resorted both to
an appeal to authority and to an ad hoc
dismissal of our results, as follows (1969,
p. 751):
Wenner and his colleagues presume to
challenge findings of a great biologist. . .
and . . . In brief, bees are easily distracted.
Wilson took a different approach when
he wrote (1972, p. 6):
1192
ADRIAN M. WENNER
The evidence . . . is overwhelmingly
in favor of a communicative function for
the waggle dance. A long series is now
published of remarkable experimental
results that have not been reasonably
explained in any other way.
I have by no means taken a poll of
experts on the subject but my impression
is that Professor Wenner's rather intricate interpretations have been increasingly questioned and are now accepted
by very few.
Only 4 years later, however, James Gould
wrote (1976, p. 241):
Von Frisch's controls do not exclude
the possibility of olfactory recruitment
alone, and Wenner is certainly correct
in saying that an endless repetition of
ambiguous experiments does not add
anything to the evidence.
Gould, however, admitted the inadequacy
of earlier results only after he felt that he
had "verified" the language hypothesis,
again with the additional acquisition of
supportive evidence.
What attitude toward the language
hypothesis can we expect from someone
who adheres not to the "verification" but
to the "falsification" approach? One of the
best examples comes from the writings of
Karl Popper himself (Popper, 1977, in Miller, 1985, p. 271). Popper treated the honey
bee language hypothesis as "fact" when he
wished to compare that "language" with
other animal "languages." He did so even
though the bee language hypothesis had
been experimentally "falsified" 10 years
earlier (reviewed in Wells and Wenner,
1973), in the very manner he had advocated earlier (Popper, 1957). (Popper was
perhaps closer to Carnap's views than he
himself realized. On the other hand, he
may not have been aware of our experimental results.)
Rosin (1978, 1980a, b, 1984, 1988) wrote
a series of articles summarizing the evidence and arguing against acceptance of
the language hypothesis, but those arguments have so far fallen on deaf ears (e.g.,
Getz, 1988). The stimulus for Rosin's articles was the fundamental nature of prob-
lems occasioned by generation of the honey
bee "language" hypothesis, as first proposed by von Frisch in 1946 (von Frisch,
1947, 1950). For example, for Rosin, the
bee language hypothesis violated an insectlevel, human-level barrier with respect to
behavioral patterns.
Are the organism and future studies of
the organism necessary for resolution of
the bee language controversy? Without a
return to such exploratory studies and by
conveniently using ad hoc and other arguments (e.g., Dawkins, 1969; Gould, 1976),
that hypothesis has now been "rescued" in
the minds of many; at the same time it has
been reduced to a state where it no longer
has predictive value (Wells and Wenner,
1973; Rosin, 1978, 1988; Wenner and
Wells, 1987; Wenner and Wells, 1990).
Future application of the verification
approach in studies of honey bee recruitment can provide more supportive evidence for the language hypothesis; a true
test of the hypothesis or multiple inference
approach will likely yield additional evidence refuting that hypothesis (see Weimer, 1979, p. 66). Resolution of that controversy thus lies in the future (Rosin,
1988).
An exemplary case: Barbara McClintock
Evelyn Fox Keller's (1983) account of
the life and contributions of Barbara
McClintock (^4 Feeling for the Organism)
appears to mesh quite well with the observation that creativity in science is best realized by an unfettered attitude on the part
of the scientist. McClintock "created
images" which even today are ill-understood by her peers. Keller (1983, p. xiv)
wrote:
Gradually I came to understand that
the importance of [Keller's] story lies
precisely in her independence. Because
of the simplicity with which she pursued
what was "obvious" to her, almost independent of response from her peers. . . .
Because of her often peripheral relation
to the community in which she worked,
her story provides a view that would normally not be possible [within] the powerful, shifting currents of group interest.
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
Keller also specified several other reasons
for McClintock's success, always within the
context that the organism 15 necessary. Of
McClintock's attitude she (1983, p. 198)
wrote:
[McClintock's] answer is simple. Over
and over again, she tells us one must have
the time to look, the patience to "hear
what the material has to say to you," the
openness to "let it come to you." Above
all, one must have a "feeling for the
organism."
Keller (1983, p. 201) continued:
A deep reverence for nature, a capacity for union with that which is to be
known—these reflect a different image
of science from that of a purely rational
enterprise. Yet the two images have
coexisted throughout history.
McClintock was somehow able to perceive what others could not see; however,
she could not "convert" a sufficient number of others to the same view within a
reasonable time for her work to have a full
and immediate impact. She apparently
lacked the "persuasive passion" described
by Polanyi (1958). To McClintock's credit,
she continued her basic research for the
personal satisfaction and thrill of understanding, exhibiting the "heuristic passion" of Polanyi.
PERSPECTIVE
1193
methodological attitudes is not requisite for
scientific research. Great discrepancies in
emphasis can thus prevail both within and
between the different subfields of biology,
as well as between the approaches advocated by different philosophers of science.
For example, tradition within a particular
community of scientists may dictate the
manner by which one attempts to falsify
working hypotheses, but evidence for a
competing hypothesis can be overlooked if
only the falsification approach is used.
Alternatively, tradition in one sub-field
of biology may favor use of the verification
approach. One who attempts to falsify a
prevailing hypothesis in that field and thus
works outside the prescribed traditional
approach may not be able to communicate
effectively with colleagues. Negative evidence obtained relative to a prevailing
hypothesis, which may be gathered by such
a person, can then be easily overlooked or
dismissed by the community.
There are some realists who take an
extreme stand and want nothing at all to
do with philosophical analysis. Theocharis
and Psimopoulos (1987), among others,
exhibited concern about the uncertainty
that can arise if one wonders too much
about how science progresses. In their
paper, one can recognize the now familiar
confrontation between Realism and Relativism (Objectivism and Relativism of
Bernstein, 1983, p. 1), labelled in their article as "true theses" and "antitheses" (1987,
p. 595):
How a scientist uses scientific method may
vary widely, from the mental orientation
We shall refer to these erroneous and
suggested by Atkinson to those proposed
harmful ideas as the epistemological
by Carnap, Popper, Chamberlin, and Platt
antitheses. . . . Articles and programmes
(Fig. 1). Within the scientific community
attacking
the scientific theses and chamthere are staunch advocates and practipioning
the
antitheses are published and
tioners of each of the methods, scientists
broadcast
regularly by the British
who view one orientation as the appropriate
media.
.
.
.
This
state of affairs is bad
orientation for all. Few seem to realize that
enough.
But
things
are even worse: pereach experimental orientation has its valversely,
many
individual
scientists and
ues and its limitations.
philosophers seem bent on questioning
Commitment to a single orientation
and rejecting the true theses, and supseems to be especially prevalent for those
porting the antitheses.
who work within the Realism School; they
apparently feel that some use of methodTraditional philosophers (logical empiological analysis is permissible, but undue ricists such as Carnap or Popper) have usuattention to divergent philosophical or ally advocated only a single approach (e.g.,
1194
ADRIAN M. WENNER
verification, falsification), reminding one
of Atkinson's "Views of the Elephant." He
wrote (1985, p. 727):
. . . the various historians of science,
philosophers of science, sociologists of
science, and psychologists of science seem
to have grasped some part of the intellectual pachyderm we call science yet
none of the various schools or "isms"
seems entirely satisfactory.
Those philosophers of science who were
scientists first and philosophers second (e.g.,
Polanyi, Kuhn, Platt, and Boehm) recognized the unnecessary restrictions imposed
by the singular attitudes insisted upon by
the more traditional philosophers of science (e.g., Carnap, Popper, and Lakatos).
For the practicing scientist, scientific
method could well include implementation
of all four of the emphases discussed above.
Exploration can be followed initially by Verification until one feels somewhat comfortable with the understanding gained. Immediately thereafter (before one gains the
"parental attachment" of Chamberlin or
before such an explanation becomes a
"Ruling Theory"), one can apply the Falsification or "Null Hypothesis" approach
and truly test an emergent hypothesis.
Better still, one can compile a list of all
feasible competing hypotheses, as in the
Multiple Inference approach, and design
experiments to eliminate as many of those
hypotheses as possible. Once that is done,
one can go back again to the Exploration
approach and search for other images. Perhaps the above composite approach is what
Feyerabend (1975) meant when he wrote,
"anything goes." It may also be akin to the
meaning of Polanyi's expression, "heuristic passion" (Polanyi, 1958), the true desire
to gain further understanding for its own
sake.
What do biologists do today? Within any
one subdiscipline of biology, scientists may
limit themselves to one prevalent orientation, a particular orientation that has
become characteristic of the manner in
which science is done in that subdiscipline.
Ecologists, for example, appear to have
placed an unduly strong emphasis on the
"null hypothesis" (falsification) approach of
Karl Popper. Specific projects in ecology,
if they are to be supported by the community, apparently must fall within precise
guidelines and must conform to that theoretical framework which has already been
verified (e.g., competition theory, predatorprey theory) before they are taken seriously. (See Loehle, 1987 and Mclntosh,
1987, for an examination of this circumstance.)
Sociobiologists appear to emphasize
assumptions implicit in the verification
approach. There is much of Polanyi's "persuasive passion" involved in that approach
(Polanyi, 1958), and exotic hypotheses can
gain ready acceptance. Microbiologists and
geneticists (fast moving fields of biology),
by contrast, work within a different tradition and apparently nearly automatically
emphasize multiple inference or strong inference (Platt, 1964) in the design of their
experiments and, in doing so, move rapidly
between exploration and inference.
In any case, research in all fields of biology has been conducted without much conscious concern about suitability of approach
or without thoughtful attention to the possible pitfalls of personal involvement in
outcomes. Consequently, projects which
may have started out within a "working
hypothesis" format (Chamberlin, 1890)
may later fall into a "ruling theory" confinement. When that happens, as Chamberlin put it, "all hope of the best results
is gone." Yet, papers continue to be
accepted for publication and large federal
grants continue to be awarded even when
it is clear that the most applicable scientific
methods are no longer being used. (See
Mahoney, 1976; Peters and Ceci, 1982;
Jackson and Prados, 1983; Loehle, 1987.)
As indicated earlier, communication
among scientists may not even be possible
within a subdiscipline if different methodological approaches are used by participants; controversy may then emerge.
Susanne K. Langer recognized that there
could be a breakdown of communication
between scientists (or between laymen for
that matter) who have different mental orientation to their disciplines. She (1967, p.
xxii) wrote:
To convince the contrary-minded is a
hopeless task and an arrogant undertak-
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
ing; they have their reasons for thinking
as I have for my way. The foundations
of a theory cannot be factually proven
right or wrong; they are the terms in
which facts are expressed, essentially
ways of saying things, that make for special ways of seeing things.
What scientists fail to realize is that
emergent controversies can and do provide the "self-correcting mechanism"
which keeps science on track and leads to
intellectual advancement (Popper, 1963).
We should calmly leave the security of our
former beliefs and welcome the insight
provided by controversy. As biologists, we
should turn again to the organism. Doing
that, we might minimize the duration of
those controversies and thereby speed our
search for understanding of nature.
Confusion could be altered by an institution of properly designed courses in the
history, sociology, psychology, and philosophy of science. Such courses should not
be merely traditional courses in methodology or philosophy but should contain
examples of experience in logic, ethics, and
epistemology. Suitable examples could be
drawn from research on the biology of
whole organisms (because of their intuitive
nature), and illustration of applicability of
the different scientific orientations (methods) could be stressed.
If these fundamentally important courses
existed in biological sciences curricula in
colleges and universities throughout the
nation, we would be far ahead in the production of a more enlightened student at
entrance to graduate school. Some of that
enlightenment might even persist into later
research programs.
ACKNOWLEDGMENTS
I thank A. Alldredge, J. W. Atkinson, K.
Cooper.J. E. Dugan, W. J. Davis, C. Loehle,
M. Mahoney, H. M. Page, and E. Schultz
for their comments on earlier versions of
the manuscript. Special thanks go to W.
Bock, L. Russert-Kraemer, and P. H. Wells
for the extra effort they expended during
the final stages of writing, as well as to the
ASZ and the Department of Biological Sciences at the University of California, Santa
Barbara for their support.
1195
REFERENCES
Adler, K. andj. B. Phillips. 1985. Orientation in a
desert lizard (Uma notata): Time-compensated
compass movement and polarotaxis. J. Comp.
Physiol. A 156:547-552.
Alcock, J.
1979. Animal behavior: An evolutionary
approach. Sinauer, Sunderland, Massachusetts.
Ardrey, R. 1963. African genesis. Readers Union, Collins, London.
Atkinson, J. W. 1985. Models and myths of science:
Views of the elephant. Amer. Zool. 25:727-736.
Bacon, Sir Francis. 1620 (1952 reprint). Novum
Organum. In R. M. Hutchins (ed.), Great books of
the Western World, Vol. 30, pp. 103-195. Encyclopedia Britannica, Inc., Chicago.
Bernstein, R.J. 1983. Beyond objectivism and relativism:
Science, hermeneutics, and praxis. Univ. of Penn-
sylvania Press, Philadelphia.
Broad, W. and N. Wade. 1982. Betrayers of the truth.
Simon and Schuster, New York.
Brown, S. M. 1986. Of mantises and myths. BioScience 36:421-423.
Bruner,J. S. and L. Postman. 1949. On the perception of incongruity: A paradigm. J. Personality
18:206-223.
Chalmers, A. 1982. What is this thing called science?
An assessment of the nature and the status of science
and its methods. 2nd ed. Univ. of Queensland Press.
St. Lucia. Queensland, Australia.
Chamberlin, T. C. 1890(1965 reprint). The method
of multiple working hypotheses. Science 148:754759.
Coffin, H. G. 1960. The ovulation, embryology, and
developmental stages of the hermit crab Pagurus
samuelis (Stimpson). Walla Walla Coll. Publs. Dep.
Biol. Sci. 25:1-28.
Craig, P. 1971. An analysis of the concept of lunar
orientation in Orchestoidea corniculata (Amphip-
oda). Anim. Behav. 19:368-374.
Craig, P. 1973a. Behavior and distribution of the
sand-beach amphipod Orchestoidea corniculata.
Mar. Biol. 23:101-109.
Craig, P. 19736. Orientation of the sand-beach
amphipod Orchestoidea corniculata. Anim. Behav.
21:699-706.
Davis, W. J. and E. Liske. 1985. Der Balztanz der
Gottesanbeterin. Naturwiss. Rundsch., Stuttgart
38:223-230.
Dawkins, R. 1969. Bees are easily distracted. Science
165:751. (Letter)
Ercolini, A. and F. Scapini. 1972. On the non-visual
orientation of littoral amphipods. Monit. Zool.
Ital. (n.s.) 6:75-84.
Ercolini, A. and F. Scapini. 1974. Sun compass and
shore slope in the orientation of littoral amphipods (Talitrus saltator Montagu). Monit. Zool. Ital.
(n.s.) 8:85-115.
Feyerabend, P. K. 1970. Consolations for the specialist. In I. Lakatos and A. Musgrave (eds.), Criticism and the growth of knowledge, pp. 197-230.
Cambridge Univ. Press, London.
Feyerabend, P. K. 1975. Against method: Outline of an
Anarchistic theory of knowledge. New Left Books,
London.
1196
ADRIAN M. WENNER
Feyerabend, P. K. 1978. Science in a free society. New Langmuir, 1. 1938. Thespeedof the deer fly. Science
Left Books, London.
87:233-234.
Feyerabend, P. K. 1981. How to defend society against Latour, B. 1987. Science in action: How tofollow scientists
science. In I. Hacking (ed.), Scientific revolutions,
and engineers through society. Open University Press,
pp. 156-167. Oxford Univ. Press, Oxford.
Milton Keynes, England.
Francon.J. 1938 (1939 translation). The mind oj the Leplin.J. (ed.) 1984. Scientific realism. Univ. of Calif.
bees. Methuen, London.
Press, Berkeley.
Franks, F. 1981. Polywater. MIT Press, Cambridge. Liske, E. and W.J. Davis. 1987. Courtship and matFrisch, V. von. 1947. The dances of the honey bee.
ing behaviour of the Chinese praying mantis,
Bull. Anim. Behav. 5:1-32.
Tenodera aridifolia sinensis. Anim. Behav. 35:15241537.
Frisch, K. von. 1950. Bees, their vision, chemical senses,
and language. Cornell Univ. Press, Ithaca, New Loehle, C. 1987. Hypothesis testing in ecology: PsyYork.
chological aspects and the importance of theory
Frisch, K. von. 1954. The dancing bees: An account of
maturation. Ecology 62:397-409.
the life and senses of the honey bee. Methuen, Lon- Maeterlinck, M. 1901. The life of the bee. (Translated
don.
by Alfred Sutro.) Dodd, Mead and Co., New York.
Getz, W. M. 1988. Dialects and the evolution of the Mahoney, M. J. 1976. Scientist as subject: The psychohoneybee dance language. Trends Evol. Ecol. 3:
logical imperative. Ballinger (Lippincott), Cam32-33.
bridge.
Gilbert, G. N. and M. Mulkay. 1984. Opening Pan- Marais, E. 1969. The soul of the ape. (Introduction by
dora's box. Cambridge Univ. Press, New York.
Robert Ardrey.) Atheneum, New York.
Gould, J. L. 1976. The dance-language controversy. Masterson, M. 1970. The nature of a paradigm. In
Q. Rev. Biol. 51:211-244.
I. Lakatos and A. Musgrave (eds.), Criticism and
Gould.J. L., M. Henerey, and M. C. MacLeod. 1970.
the growth of knowledge, pp. 59-88. Proceedings of
Communication of direction by the honey bee.
the International Colloquium in the Philosophy
Science 169:544-554.
of Science, London, 1965. Vol. 4.
Griffith, B. C. and N. C. Mullins. 1972. Coherent Mclntosh, R. P. 1987. Pluralism in ecology. Annu.
social groups in scientific change. Science 177:
Rev. Ecol. Syst. 18:321-341.
959-964.
Michalos, A. C. 1971. The Popper-Carnap controversy.
Haight, H. V. 1926. Around the world in twentyMartinus Nijhoff, The Hague, Netherlands.
four hours. Sci. Mon. 22:469-472.
Miller, D. (ed.) 1985. Popper selections. Princeton
Hartwick, R. F. 1976. Beach orientation in talitrid
University Press, Princeton, NJ.
amphipods: Capacities and strategies. Behav. Ecol. Nye, M. J. 1980. N-rays: An episode in the history
Sociobiol. 1:447-458.
and psychology of science. Hist. Stud, in Phys.
Holldobler, B. 1971. Homing in the harvester ant
Sci. 11:125-156.
Pogonomyrmex badius. Science 171:1149-1151. Peters, D. P. and J. Ceci. 1982. Peer-review practices
Jackson, C. I. and J. W. Prados. 1983. Honor in
of psychological journals: The fate of published
science. Amer. Sci. 71:462-464.
articles, submitted again. Behav. Brain Sci. 5:187255.
Johnson, D. L. 1967. Honeybees: Do they use the
direction information contained in their dance Platt, John. 1964. Strong inference. Science 146:
maneuver? Science 155:847-849.
347-353.
Keller, E. F. 1983. A feeling for the organism: The life Polanyi, M. 1958. Personal knowledge: Towards a postand work of Barbara McClintock. Freeman, New
critical philosophy. Univ. of Chicago Press, ChiYork.
cago.
Kneller, G. F. 1978. Science as a human endeavor. Popper, K. 1957. Philosophy of science: A personal
Columbia Univ. Press, New York.
report. In C. A. Mace (ed.), British philosophy in
Kuhn, T. S. 1970a (1962). The structure of scientific
the mid-century, pp. 155—191. Macmillan, New
revolutions, 2nd ed., enlarged. Foundations of the
York.
unity of science, Vol. ii, No. 2. Univ. of Chicago Popper, K. 1963. Conjectures and refutations. Harper,
Press, Chicago.
New York.
Kuhn, T. S. 19706. Logic of discovery or psychology Popper, K. 1985 (1977). The mind-body problem.
of research? In I. Lakatosand A. Musgrave (eds.),
In D. Miller (ed.), Popper selections, pp. 265-275.
Criticism and the growth of knowledge, pp. 1—23.
Princeton Univ. Press, New Jersey.
Cambridge Univ. Press, London.
Roeder, K. D. 1935. An experimental analysis of the
Lakatos, I. 1968. Changes in the problem of inducsexual behavior of the praying mantis (Mantis relitive logic. In I. Lakatos (ed.), The problem of inducgiosa L.). Biol. Bull. 69:203-220.
tive logic, pp. 315-417. North-Holland Publ. Co., Rosin, R. 1978. The honey bee "language" controAmsterdam.
versy. J. Theor. Biol. 72:589-602.
Lakatos, I. and A. Musgrave. (eds.) 1970. Criticism Rosin, R. 1980a. Paradoxesof the honey-bee "dance
language" hypothesis. J. Theor. Biol. 84:775and the growth of knowledge. Proc. of the Interna800.
tional Colloquium in the Philosophy of Science,
Vol. 4: London, 1965. Cambridge Univ. Press, Rosin, R. 19806. The honey-bee "dance language"
London.
hypothesis and the foundations of biology and
behavior. J. Theor. Biol. 87:457-481.
Langer, S. K. 1967. Mind: An essay on human feeling,
Vol. 1. The Johns Hopkins Press, Baltimore.
Rosin, R. 1984. Further analysis of the honey bee
CONCEPT- VS. ORGANISM-CENTERED BIOLOGY
1197
"dance language" controversy. I. Presumed Wells, P. H. and A. M. Wenner. 1973. Do bees have
proofs for the "dance language" hypothesis, by
a language? Nature 241:171-174.
Soviet scientists. J. Theor. Biol. 107:417-442.
Wenner, A. M. 1967. Honeybees: Do they use the
Rosin, R. 1988. Do honey bees still have a "dance
distance information contained in their dance
language"? Amer. Bee J. 128:267-268.
maneuver? Science 155:847-849.
Shipley, B. and P. A. Keddy. 1987. The individual- Wenner, A. M. 1974. Information transfer in honistic and community-unit concepts as falsifiable
eybees: A population approach. In L. Krames, T.
hypotheses. Vegetatio 69:47-55.
Alloway, and P. Pliner (eds.), Nonverbal communication: Advances in the study of communication and
Silvernale, M. N. 1965. Zoology. Macmillan, New York.
effect, Vol. 1, pp. 133-169. Plenum Press, New
Smithers, C. N. 1977. Seasonal distribution and
York.
breeding status of Danaus plexippus (L.) (Lepidoptera: Nymphalidae) in Australia. J. Aust. Wenner, A. M. and A. M. Harris. 1989. Do CaliforEntomol. Soc. 16:175-184.
nia monarchs undergo long distance directed
migration? (Symposium volume: Second InterTheocharis, T. and M. Psimopoulos. 1987. Where
national Conference on the Monarch Butterfly,
science has gone wrong. Nature 329:595-598.
Los Angeles, 24-27 September 1986.) (In press)
Thompson, 1831. On the metamorphoses of decapodous Crustacea. Zool. Journ. 5:383-384.
Wenner, A. M. and P. H. Wells. 1987. The honey
Townsend, C. H. T. 1926. Around the world in a
bee dance language controversy: The search for
daylight day: A problem in flight. Sci. Mon. 22:
"truth" vs the search for useful information.
309-311.
Amer. Bee J. 127:130-131.
Townsend, C. H. T. 1927. On the Cephenemyia mech- Wenner, A. M. and P. H. Wells. 1990. Anatomy of a
controversy: The question ofa "language" among bees.
anism and the daylight-day circuit of the earth
Columbia Univ. Press, New York. (In press)
by flight. J. N.Y. Entomol. Soc. 35:245-252.
Townsend, C. H. T. 1939. Speed of Cephenemyia. J. Wenner, A. M., P. H. Wells, and D. L.Johnson. 1969.
N.Y. Entomol. Soc. 47:43-46.
Honeybee recruitment to food sources: Olfaction
or language? Science 164:84-86.
Wehner, R. 1976. Polarized light navigation by
West wood, J. O. 1835. On the supposed existence
insects. Sci. Amer. 235:106-115.
of metamorphoses in the Crustacea. Philos. Trans.
Wehner, R. and P. Duelli. 1971. The spatial orienRoy. Soc, London, pt. 2, pp. 311-328, pi. 4.
tation of ants Cataglyphis bicolor, before and after
sunrise. Experientia 27:1364-1366.
Williams, C. B. 1958. Insect migration. Collins, London.
Wehner, R. and R. Menzel. 1969. Homing in the ant
Wilson, E. O. 1972. (Letter Exchange.) Sci. Amer.
Cataglyphis bicolor. Science 164:192-194.
227:6.
Weimer, W. B. 1979. Notes on the methodology of scientific research. Lawrence Erlbaum Assoc, Hillsdale, N.J.