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Theoretical Integration, Cooperation, and
Theories as Tracking Devices
James Griesemer
Departments of Philosophy and Center for Population Biology
University of California, Davis, CA, USA
[email protected]
The theoretical problem of integrating evolution, heredity, development, and cognition has a long pedigree with a complicated history. Many of these fields were considered the subjects
of one science in the 19th century (Maienschein 1987). Leading
theoretical biologists of the age wrote large, expansive treatises
that biologists now can read only with equal measures of wonder and incredulity: wonder at their wide and deep learning and
incredulity at their peculiar visions of biological integration.
Think of Herbert Spencer (1900, pt 2: 367): “Evolution is an
integration of matter and concomitant dissipation of motion;
during which the matter passes from a relatively indefinite,
incoherent homogeneity to a relatively definite, coherent heterogeneity and during which the retained motion undergoes a
parallel transformation.” Or consider Ernst Haeckel (Gastraea
theory), Hans Driesch (entelechies), August Weismann (biophors, ids, idants), and even Charles Darwin (use inheritance,
pangenesis). None of these ideas were especially crazy in their
time, but they drove integration projects that are quite different
from ours.
Historically, each of these special subjects has been a
contender for the role of primary theory from which an understanding of all biological phenomena would flow. To take but
a few examples, Dobzhansky (1973: 125) gave us “Nothing in
biology makes sense except in the light of evolution” on behalf of Darwin’s theory. De Vries gave us his Mutationstheorie
and Johannsen his pure line theory as alternative explanations
of the root cause of hereditary change. Both were intended
originally to replace natural selection as theoretically primary.
Driesch gave us entelechies of individual development as a fundamental philosophical principle for the biological sciences.
James Mark Baldwin, Pierre Teilhard de Chardin, Francisco
Varela, Rupert Riedl, and Donald Campbell, in very different
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ways, put cognition in the center of theoretical concern for
general biological science, evolution in particular.
Thinking about integration from the point of view of any
of these specialties has traditionally depended on a notion of
unification by explanatory theory reduction: explanations flow
from a fundamental primary theory to phenomena that are
organized at the surfaces, so to speak, of those phenomena
considered central for the primary theory—for example, trait
heredity and organism development in the light of evolution
in Darwin’s day, or population change and the development of
hybrids in the light of Mendelian heredity circa 1905, recapitulation and phylogeny in the light of ontogeny in Haeckel’s,
de Beer’s, and Gould’s days (Haeckel 1866; de Beer 1930;
Gould 1977), learning as coequal to inheritance and selection
as a force in evolution.
In the 20th century, phenomena have tended to be organized hierarchically, in terms of the composition of matter
from the smallest to the largest units, so that there is a general
expectation and drift of reductionism toward lower levels of
compositional organization—to the behavior of smaller and
smaller bits of matter—as the primary concern of fundamental
theory. In biology, this reductionist drift led theorists from organisms to organs and tissues, to cells, and on to molecules as
the foci of theoretical interest. Genes, the “molecules of life,”
have been the focus of biological reductionism for nearly all
of that century.
A different model of integration centers on cooperation and communication among theoretical and phenomenal
equals, rather than on imperialism and competition for primacy
and fundamentality, which reduces or replaces one theory by
another or trivializes one explanandum as epiphenomenal to
another. An explanatory domain can become integrated when
its bumps, twists, and turns are smoothly traversable, but we
need not achieve integration by leveling the domain and making it conceptually homogeneous, just as nation-states can be
unified by the smooth flow of goods, services, people, and
ideas across their borders rather than by the obliteration of
local and regional differences making flow irrelevant.
September 27, 2005; accepted October 1, 2005
c 2006 Konrad Lorenz Institute for Evolution and Cognition Research
Biological Theory 1(1) 2006, 4–7. James Griesemer
Increasingly in science, as in other endeavors, “Cooperative work now routinely spans organizational, geographical
and other boundaries” (Gerson in press). Likewise, explanatory reductionism can be practiced as a multidisciplinary,
cooperative heuristic that serves as a means of triangulation to
truer theories (Wimsatt 1987, 2002). A cooperative approach
to theory making facilitates explanatory traversals in all
directions between models, levels, and perspectives, not just
from higher levels to lower ones. There is no reason we cannot
and should not turn reductionism on its head and use genetic
theories as tools to build developmental ones as well as
seek reduction of developmental phenomena to gene activity
(Griesemer 2000, 2002, 2005a). What is left of primacy or
foundations is simply the idea that we start from what we
know and move out from there, guided by our theoretical
commitments. And when we meet others guided by different
commitments, we try to reconcile our differences.
Genetics is not more fundamental than development, it
is just that its theoretical apparatus was rationalized first (see
Gerson in press on the process of rationalization). That is,
its conceptual parts were made to fit and work smoothly and
economically together, in no small measure by the experimental and mathematical insights of Mendel and the cytologists
who studied chromosome behavior in meiosis. Hence genetics tends to be, but need not be, one’s first stop in crafting a
strategy for integration of biological theory. Why should we
think that the first formal, articulated theory we come to in a
domain will turn out to be more fundamental than those arrived
at later, even if it is true that, by getting there first, a theory
tends to become entrenched in our ways of looking for (and
at) subsequent theory (see Wimsatt 1986 on entrenchment)? If
anything, early theories get articulated first because they are
easier to work with, or by accident, or through a lucky break,
or because of a social or political or economic consequence
that motivated early workers to start there. What does any of
that have to do with being “fundamental” or “foundational”
in the logical sense reductionists expect? Mendel’s theory was
easily articulated in an algebraic form because his materials
and experiments were arranged to do just that. It is not just that
Mendel’s mind was prepared to make his famous discovery; his
discovery was prepared to make his theory. The developmentally invariant property Mendel called a factor or “Merkmal”
is just the sort of property that can be easily expressed mathematically (see Griesemer in press).
Conceptual boundaries, e.g., between genetical, developmental, and evolutionary problems, can be traversed by theoretical, experimental, or observational means. If mathematical
results can flow seamlessly between representations of networks of interacting genes and the statistics of populations undergoing natural selection in changing environments, or niche
constructing activities of species can be represented in a model
alongside the contributions of Mendelizing alleles, then the
Biological Theory 1(1) 2006
theoretical domain of evolution can be traversed in integrated
and unified ways across the disciplinary boundaries separating heredity, development, and evolution. If manipulation of
the development of hybrid offspring in breeding experiments
can be traced to consequences for quantitative trait loci or nucleotide sequences located in particular chromosomal regions
and nucleotide changes can be tracked through molecular interactions to phenotypes, then experimental domains can be
traversed across the heredity-development border in an integrated and unified way. If microscopists can track luminescent molecules across stages of embryonic development from
fertilization through differentiation of new germ cells, then
observation can traverse the border between development and
heredity.
If such traversals are the occasion for integration by cooperation, e.g., when scientists approach a phenomenon from
different paths, what can we expect the theoretical products
to look like? What, for example, would a theory of EvoDevo
look like, as opposed to the motley collection of applications
of existing theory we have now: of phylogenetic inference
to and from developmental characters (Gould 1977), of population genetics models to systems of developmental genes
(Nijhout and Paulsen 1997), of molecular developmental models of cis-acting regulatory regions to evolution and phylogeny
(Davidson 2001)? A merger of classical theories does not seem
like the right answer for a new science of EvoDevo, nor for any
of the other integration projects one can imagine for evolution,
heredity, development, and cognition.
Why not? One reason is that these theories were formulated under the assumption that the processes they describe,
while sharing fundamental units, are separable. This is a generalization of the doctrine of Weismannism (Griesemer and
Wimsatt 1989). Weismannism pictures the relation of heredity
and development as separate processes flowing from a common cause: the gene, or, in Weismann’s day, the germ plasm
(see Weismann 1893). Evolution, under Weismannism and
later Mendelism, becomes alteration of the flow and distribution of germ plasm or genes within and among generations
because that process was separated by a theoretical abstraction
from development. By abstraction from inheritance, development becomes the elaboration of germ plasm (gene expression) into the soma (body), whose interactions with external
environments subject germ plasm to forces of evolution. (On
abstraction as a means of theory generalization, see Griesemer
2005b.) With the perspective of Weismannism in hand, theoretical integration looks like a problem concerning the duality
of the gene’s autocatalytic and heterocatalytic roles and their
consequences.
Weismannism, however, faces a severe problem as a fundamental doctrine for biology despite the successes of this
rationalization strategy in the 20th century. First, as Buss
(1983, 1987) pointed out, the distribution of the Weismannian
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Theoretical Integration, Cooperation, and Theories as Tracking Devices
“preformistic” mode of development is spotty and rather rare
among living phyla, so it ill-serves the general theoretical
project. Second, the autocatalytic and heterocatalytic properties of the gene are not intrinsic to nucleic acid polymers but are
“in” the systems in which they are embedded and these systems
engage the “separate” processes of inheritance, development,
and evolution simultaneously and integratedly. Replication is
a cellular process. Genetic encoding and decoding are cellular
processes. They are not properties of genes.
Consider, for example, where the genetic code lies. The
triplet code is best described as residing in the aminoacyltRNA synthetase enzymes that can simultaneously recognize
particular amino acid species and particular tRNA species.
The set of relations between these two recognition functions
constitutes the genetic code and is embodied in the collection
of enzymes that cells have at any particular living moment.
Not only does this entail that a genetic code can exist without
any genes (a test tube of enzymes will do), but it also implies
that any failure to transmit one or more of the 20 classes of
these enzymes from parent cell to offspring cell constitutes a
change in a genetic code. To an evolved, complex cell, such
failures are always lethal “mutations.” Mutation is in quotation marks because, of course, the transmission of enzymes
from parent to offspring is not usually implied by the term.
Indeed, since enzymes are part of the soma and their function
is translation, not replication, their role in the autocatalytic
function of genes is obscured. And yet, if not for their direct
hereditary transmission, these enzymes could not play any role
in gene heterocatalysis. It does no good to pretend, by invoking Weismannism, that enzyme properties can be explained
by gene expression one generation back, reducing the problem to one of heterocatalysis in a previous generation, because
something must explain the inheritance of the enzymes in the
previous generation in order to account for the heterocatalytic
function of genes there. This molecular chicken-egg problem
is, in fact, the problem of Weismannism restated in molecular
terms. It cannot be invoked to explain the causal roles of genes
without begging the question of the origin of the functional
relation already established between the enzymes and the
genes.
The temptation is to address this kind of theoretical problem by assuming a weak “contextualism,” seeking to integrate
evolution and development by sorting out the “right” contexts
in which to invoke developmental principles and those in which
to invoke genetic/hereditary ones. That would be another form
of theoretical segregation like the nature/nurture dichotomy,
one which Susan Oyama (2000) was right to criticize. Integrating theory cannot simply be a matter of identifying the “right”
context for different sorts of principles; moreover, because the
contexts themselves have histories—they are no more fixed
than genes are. More importantly, biological contexts, like biological processes, are entwined. The context of gene expres6
sion is also the context of gene heredity in the sense that the
forces influencing the distribution and probability of transmission occur in the context of gene expression. And the context
of gene heredity is also the context of gene expression, since
the configurations governing gene expression depend on “maternal” inheritance and the conditions transmitted from parent
to offspring.
An adequately integrated biological theory will be one
which respects the entwinement of biological processes, even
while it abstracts them for reasons of mathematical tractability, experimental control, and observational tracking. In fact,
it might be best to drop old assumptions about what theories
are like for the sake of making progress toward integration.
One common thread through my remarks has been that, with
whatever mix of theoretical, experimental, and observational
techniques biologists proceed, their activity always involves
tracking: following genes, cells, organisms, mathematical
quantities, light spots on film, phenotypes in genetic hybrids,
and so on. Perhaps it is time to get away from a physicscentred view of theories and theory making and adopt one
more appropriate to biology. As far as EvoDevo goes, we might
begin with the recognition that development is both cause and
consequence of genetic continuity, so the project of tracking
development through inheritance and inheritance through development will require new ways of thinking about theories
and theoretical integration.
An exciting prospect for this new journal, Biological Theory: Integrating Development, Evolution, and Cognition, is
that each of its subdomains developed using tracking methods suited to its particular phenomena and model systems.
As contributions to the journal traverse the boundaries, new
opportunities and new ways of theory making will arise, provided investigators remain open to the project of theoretical
integration by cooperation and collaboration.
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