AMER. ZOOL., 29:1057-1060 (1989)
Prologue: The Necessity of the Organism1
LOUISE RUSSERT-KRAEMER
Department of Zoology, SCEN-632, University of Arkansas,
Fayetteville, Arkansas 72701
AND
WALTER J. BOCK
Department of Biological Sciences, Columbia University,
New York, New York 10027
This symposium is directed to an emphasis and understanding of the argument that
the organism, and the time-space organization of the organism, must constitute a
central focus in modern biology and in all
biological explanations. We insist on moving beyond the historically persistent dispute between claims of reductionists and
holists. That is not what we are about. We
are firmly committed, from the outset, to
the essential power of reductionistic techniques in all science. To reject reductionism in biological study is to retreat from
analysis—a defeatist approach. But we are
also strongly aware that reductionist procedures will not unilaterally delineate, let
alone solve, most problems in biology. In
the present instance, for example, careful
analysis of various bits and pieces of organisms is just part of a much larger, much
more challenging task. Sophisticated present-day, reductionary techniques will have
failed us if their results cannot in turn be
applied to critical syncretization in the difficult task of understanding whole, living
organisms. We think the time for examining and dealing with these concerns is
here. Accordingly we have organized this
symposium.
A fundamental problem in modern biology, as expressed in the composition
of biological departments in North American universities, in available research support and in the distribution of published
papers, is that two areas of research and
teaching are emphasized. On the one hand,
major stress is placed on molecular and
cellular biology, with interactions between
individual cells being the highest level of
organization considered, and with absolutely no interest in how these molecules
and cells interact to form tissues, organs,
organ systems and whole organisms. On
the other hand are behavioral-ecological
studies in which the individual organisms
are treated as so many black boxes which
interact with each other and with environmental factors, but with no understanding
of the inner workings of the organisms
which underlie these interactions. This
view, that ethology, the study of the biological basis of animal behavior, was slated
to dwindle within a couple of decades to a
narrow conduit between the ever ballooning fields of molecular biology and population genetics, was asserted by E. O. Wilson (1975) in a rather cavalier fashion.
Organismic biology is as yet an undernourished orphan in biology, unwanted
because it is not understood by biologists
interested in levels above and below the
central one of individual organisms. Yet,
the expected vigorous argument from the
organismic biologists has not developed.
We hope this symposium will constitute a
vantage point from which, at least, a discussion may be initiated.
This problem is well illustrated in the
recent volume, A Feeling for the Organism.
The Life and Work of Barbara McClintock, by
Evelyn Keller (1983) which was in many
ways a starting point for this symposium.
The central theme of this historical analysis is that Barbara McClintock's extremely
1
From the Symposium on Is the Organism Necessary?
presented at the annual meeting of the American thorough understanding of the corn
Society of Zoologists, 27-30 December 1987, at New organism was a major factor in her discovOrleans, Louisiana.
ery of transposition of genetic material
1057
1058
L. RUSSERT-KRAEMER AND W. J. BOCK
(jumping genes), and for her eventual winning of the Nobel Prize. The importance
of this "feeling for the organism" is
emphasized by a sad antithetical story. Professor Alberto Mancinelli, the lone botanist in the Department of Biological Sciences at Columbia University, told one of
us (Bock) that following the announcement
of Barbara McClintock's Nobel Prize, he
received a puzzling series of telephone calls
from leading geneticists in the New York
City region. All of them asked him the same
question about the relationship between the
genetic material of the individual peas
(seeds) in a pea pod and that of the parent
plant. Only after several calls with this
peculiar question did Professor Mancinelli
realize the underlying confusion; this
thinking was rather beyond his comprehension as he is a good organismic biologist. These geneticists assumed that simply
because the peas were lined up in a single
row within the pod, they were analogous
to the ascospores lined up in a row within
the ascus (spore case) in Neurospora and
other Ascomycetes (sac fungi). Therefore,
pea seeds within a pod should be the result
of reduction-division from the same adult
diploid cell, and hence be especially suitable for genetic study—as had been discovered some time ago by Gregor Mendel
but for quite different reasons. Development of "a feeling for the organism" can
be critical in the making of significant discoveries. Without it, the biologist may as
well resign him/herself to plodding along,
doing "normal scientific work," and as
Thomas Kuhn (1970) describes it, filling
in the unexciting details.
We do not wish to assert that organismic
thinking, even when well done, will always
lead to correct solutions; it will not! Good
organismic research depends on valid data
from studies on lower levels of biological
organization. To cite a specific example,
the honey-guides (Indicatoridae) are a small
family of mainly African birds, related to
the barbets (Capitonidae) arid the woodpeckers (Picidae). They are able to feed
and subsist for a time on bee's wax, and
one species even leads mammals, including
humans, to bee hives and feeds on the remnants, mainly the wax, left by the mam-
malian predators. Herbert Friedmann
(1955), having demonstrated that honeyguides were able to survive for about a
month on a diet of pure bee's wax before
suffering from lack of essential minerals
and vitamins, undertook an analysis of their
wax-digestion abilities. Based on extrapolations from mammalian physiology, he
surmised that birds lacked digestive
enzymes for processing the wax they
ingested, and assumed that wax-digestion
in honey-guides was accomplished with the
assistance of symbiotic micro-organisms.
Collaborating with several microbiologists,
Friedmann discovered several microorganisms in the digestive tract of honeyguides which possessed the ability to digest
wax, and concluded that these symbiotic
micro-organisms had a central role in the
wax-digesting abilities of honey-guides
(Friedmann and Kern, 1956; Friedmann et
al., 1957; Friedmann, 1967).
Their conclusions appeared to be most
reasonable and well supported by factual
data, and they have been widely accepted;
unfortunately, however, they are not correct. Subsequently, Place and his associates
(Place and Roby, 1986; Roby et al, 1986;
Diamond and Place, 1988), using an excellent blend of reductionistic and organismic
approaches, demonstrated clearly that
members of several quite different groups
of birds, including honey-guides, possess
the digestive machinery needed to digest
waxes without the assistance of microorganisms. This example illustrates the
absolute requirement of basing integrative, holistic work on correct reductionistic
analyses. The microbiological and the
digestive physiological studies done by
Friedmann and his associates were faulty,
thereby negating their integrative analysis,
a defect overcome by Place and his coworkers.
What are the dimensions of the problem
of organismic thinking—the "feeling for
the organism"?
Are biologists simply "preparationists"?
Alexandrowicz (1967, p. 320) noted that
many neurophysiologists work with isolated sensory tissue of arthropod effector
tissues, such as the muscle receptor organs
(MRO) of the Crustacea. He noted (p. 320)
THE ORGANISM IS NECESSARY
that "It is true that since the existence of
the MRO in Crustacea has been established, a lot of experimental work has been
done with them and they have been subjected to many trials. They have been stimulated with electric current, cooled and
warmed up, bespattered or injected with
various drugs, impaled, stretched and
overstretched. The nerve cells, put so to
speak on the rack, responded in various
ways; they even gave their inquisitors some
valuable information about the mechanism
of their action, but they have not revealed
their most important secret, namely, with
what other nerve elements they are in relation and to exactly that purpose they send
their signals to them. In point of fact, this
question has seldom been asked." Such
questions will not even occur to biologists
unless they go beyond the stage of simply
being only preparationists and start considering the relationship of the particular
feature to the rest of the organism. That
is to start asking the question: How does
this feature fit into the entire construction
of the organism?
Are biologists nothing more than "closet
generalists"? In 1974 at the ASZ Symposium in Tucson, "New Directions in Comparative Physiology and Biochemistry,"
honoring C. Ladd Prosser, Donald Kennedy rose to speak and prefaced his presentation with the observation that he had
listened to his respected colleagues for two
whole days—as they all assured their audience of the particular glories of the particular organisms with which they worked.
Kennedy went on to note that he suspected, nevertheless, that he was in a room
full of "closet generalists." He described a
closet generalist as one who would desert
his own animal at once in favor of the perfect animal if it were found. The perfect
animal for neurological research, Dr. Kennedy described, was one with no more than
a few dozen neural soma in its major ganglion, no neural fibers less than several
hundred micrometers in diameter, and with
completely stereotypic behavior. This
approach may be suitable for the discovery
of functional properties of living organisms, but it is completely inadequate for
inquiries into the evolutionary biology of
1059
living organisms which requires a comparative approach of real organisms with all
of their complexity. Such comparativeevolutionary approaches are essential if one
hopes to obtain a full explanation of the
diversity of features found in living organisms (Bock, 1988).
When we attempt to dissect the foundations of the conflict regarding the
"necessity" of the organism, as the participants in this symposium have done ably,
some surprising findings emerge. The
reader will find that the papers comprising
the proceedings of this symposium on "Is
the Organism Necessary?" constitute a
vantage point from which a very respectable and challenging dialogue can be
mounted. Is Wilson correct? What have we
got to say for ourselves? The core idea that
we would like all members of the audience
to take away from this symposium is that
biological organization on the level of the
organism is central to all biological analyses and that "a feeling for the organism"
is not just a nicety, rather it is the central
ingredient which will make the difference
between success or failure in much biological research.
Michael Ruse, as a philosopher of science, addresses the seminal epistemological question "Do organisms exist?", considering the distinction between animate
and inanimate objects. He argues that living organisms display teleological behavior, although using a somewhat different
concept of teleology than usually accepted
(see Mayr, 1974). Keith Benson examines
the history of ideas on the concept of the
organism and discusses the relative newness of this concept within biology. Robert
Barnes analyzes our understanding of the
diversity of animals, pointing out changes
in knowledge of. the number of species in
major groups, the recent discovery of a
number of new major taxa, and stresses the
alarming paucity of information about most
animal species on this planet. He shows that
most generalizations about the properties
of major taxa are based on a very small
sample of species. Betsy Dyer analyzes the
universal symbioses of organisms and concludes that the boundaries of individual
organisms become difficult or impossible
1060
L. RUSSERT-KRAEMER AND W. J. BOCK
to delimit. Rather the units which we generally denote as an individual organism may
actually be symbioses of many individuals
of several different types of organisms. Alan
Kohn analyzes levels of organization existing in biology, discussing the two different
sets—functional and taxonomical—and
their role in understanding the theme of
this symposium, namely organization at the
level of individual organisms.
Howard A. and Sylvia G. Lenhoff provides a useful bridge between the historical
analysis of Keith Benson and the main body
of the symposium with a discussion of the
early organismic approach in the 18th century by Abraham Trembley in his landmark studies of Hydra compared with their
present day investigations of this animal.
The core of the symposium is comprised
of the following five papers in which the
several speakers present examples from a
diversity of animal groups and functional
systems to demonstrate that an understanding of biological organization at the
organismal level was critical to the solution
a number of biological problems. Walter
Bock uses several examples from the vertebrates to illustrate the connectivity
approach in structural-functional analyses. Robert Reid discusses ontogenetic
development of adult "gutlessness" of a
clam to illustrate loss of a major organ system under special nutritive conditions.
Louise Russert-Kraemer examines interconnections between the nervous and
reproductive systems, showing the high
degree of plasticity of these systems and
their conjunctions. Bernd Heinrich uses
thermoregulation to show how organismreferenced experiments serve to ferret out
special mechanisms of heat-regulation.
(Unfortunately, sickness prevented Professor Heinrich from attending and presenting his paper in New Orleans.) Ann Kammer discusses neurophysiology of flight and
its relationship to thermoregulation and
metamorphosis in a locust, showing that
careful attentiveness to the organism
guided her successful experimental design.
Adrian Wenner examines the role of
organismic thinking in the important dual
questions of the orientation of the zoologist toward scientific methods and of the
reception of particular empirical results
depending on the firmness with which theories are held. (Richard Dawkins [Oxford
University] presented the final paper at the
symposium entitled "Necessary for What?"
which supported the arguments put forth
by the other participants. Unfortunately
he was not able to provide us with a manuscript for the Proceedings of the Symposium.)
REFERENCES
Alexandrowicz, J. S. 1967. Receptor organs in thoracic and abdominal muscles of Crustacea. Biol.
Rev. 42:288-326.
Bock, W. J. 1988. The nature of explanations in
morphology. Amer. Zool. 28:205-215.
Diamond, A. W. and A. R. Place. 1988. Wax digestion in Black-throated honey-guides, Indicator
indicator. Ibis 130:558-561.
Friedmann, H. 1955. The honey-guides. Bull. U.S.
Nat. Mus. 208:1-292.
Friedmann, H. 1967. Avian symbiosis. In Symbiosis,
Vol. 2, pp. 291-316. Academic Press, New York.
Friedmann, H. and J. Kern. 1956. The problem of
cerophagy or wax-eating in the honey-guides.
Quart. Rev. Biology 31:19-30.
Friedmann, H., J. Kern, andj. H. Rust. 1957. The
domestic chick: A substitute for the honey-guide
as a symbiont with cerolytic microorganisms.
Amer. Nat. 91:321-326.
Keller, Evelyn Fox. 1983. A feeling for the organism.
The life and work of Barbara McClintock. W. H.
Freeman & Co., New York and San Francisco.
Kuhn, Thomas. 1970. The structure of scientific revo-
lutions. 2nd ed. Univ. of Chicago Press, Chicago.
Mayr, E. 1974. Teleological and teleonomic: A new
analysis. Boston Studies Phil. Sci. 14:91-117
(reprinted, 1976, Evolution and the diversity of life,
Harvard Univ. Press, Cambridge, Mass.).
Place, A. R. and D. D. Roby. 1986. Assimilation and
deposition of dietary fatty alcohols in Leach's
Storm Petrel, Oceanodroma leucorhoa. J. Exp. Zool.
240:149-161.
Roby, D. D., A. R. Place, and R. E. Ricklefs. 1986.
Assimilation and deposition of wax esters in
plankivorousseabirds.Journ. Exp. Zool. 238:2941.
Wilson, E. O. 1975. Sociobiology. The new synthesis.
Harvard Univ. Press, Cambridge, Mass.