(Autumn, K. 1995. Chapter 1, Section 2. Integrative Biology. Ph.D.
Dissertation, University of California at Berkeley)
Copyright (c) 1995 - All rights reserved
Kellar Autumn
Museum of Vertebrate Zoology and Department of Integrative Biology
3101 Valley Life Sciences Building
University of California, Berkeley CA 94720-3160
email: gecko@garnet.berkeley.edu
INTEGRATIVE BIOLOGY
Reductionism and holism form one of the fundamental dichotomies in science
in general and biology in particular (Reippel 1988). Reductionism has been
the predominant paradigm in biology during the 20th century (Benson 1989)
-as witnessed by the current trend to use molecular explanations for all
aspects of life (Savageau 1991). It could be argued that science is fundamentally
reductionist in its methodology, and that as biology progresses, its explanations
must be on an increasingly small scale. On the contrary, I will argue that
a truly progressive biology is not solely reductionist. Holism has been
historically associated with metaphysics, and the eclipse of vitalism by
(reductionist) materialism during the enlightenment (Benson 1989) has given
holism a negative connotation in science. However when viewed in a modern
context, holism need not be linked in any way to metaphysics (Ruse 1989).
Holism is at the philosophical core of studies of complex, integrated systems
such as living organisms (Wake et al. 1983; Wake and Larson 1987; Russert-Kraemer
and Bock 1989; Pahl-Wostl 1993). The holistic approach to organismal biology
has as its central tenet the assumption that a living organism is greater
than the sum of its parts (Ruse 1989). This approach is integrative
because it focuses on the interaction or integration of parts in a functioning
organism. Integrative biology may use typically reductionist methods, but
in a philosophically holistic framework. The difference between reductionist
and integrative biology is not simply a philosophical distinction; the integrative
approach to biology differs dramatically from reductionist biology in both
its goals and its methods. While the goal of reductionist biology is to
understand life in terms of simple deterministic principles analogous to
Newtonian physics or chemistry (Savageau 1991), the goal of integrative
biology is to understand the structure, function, and history of organisms
and their environments. These are not mutually exclusive goals. Much of
the progress made by reductionist biology can be applied to reach the more
inclusive goal of integrative biology.
Because organisms and their environments form complex integrated systems,
the most robust biology will be an integrative biology (Ruse 1989; Russert-Kraemer
and Bock 1989; Savageau 1991). Organisms and their environments are studied
(of necessity) on many hierarchical levels -for instance the focal unit
of biological organization can be a gene, cell, organism, population, or
species, depending on the focal level of study (Jacob 1977; Bock 1989).
Scientific explanations rely on establishing causal relationships among
units of study (Bock 1989). One the most fundamental methodological differences
between reductionist and integrative biology is in models of causality.
Reductionist biology has a much more restrictive model of causality than
does integrative biology. The reductionist model is one of unidirectional
causation (Fig. 1.1). Reductionism assumes that
effects at higher levels of biological organization can always be reduced
to causes at lower levels (Jacob 1977). For instance, the reductionist approach
assumes that effects at organismal level can always be reduced to causes
at the gene level. Models of causality in integrative biology are much more
complex. The integrative model is one of multidirectional causality that
traces causal threads within and between hierarchical levels, and through
time. For instance, population dynamics and natural selection on individual
organisms can determine which genes are replicated in future generations.
In the integrative model, causality can occur from the highest levels to
the lowest: temperature dependent sex determination is an example of how
the environment can affect gene expression (Ewert et al. 1994; Lang and
Andrews 1994; Viets et al. 1994). Only one of many possible directions of
causation is from gene to organism. In this sense, integrative biology encompasses
the methods of reductionist biology but goes much farther. Reductionist
biology has made tremendous advances in the "parts catalog" (Savageau
1991) of life. However, the reductionist approach has failed to make significant
progress in the study of complex systems (Savageau 1991). This is primarily
because system dynamics often cannot be reduced to a linear causal model.
It is time to move beyond reductionism to expand the understanding of how
parts fit together to form functioning, evolving organisms.