Weber 1989

EVOLUTION IN THERMODYNAMIC PERSPECTIVE:
AN ECOLOGICAL APPROACH
Bruce H. Weber
Department of Chemistry and Biochemistry
California State University, Fullerton
Fullerton, CA 92634
David J. Depew
Department of Philosophy
California State University, Fullerton
Fullerton, CA 92634
C. Dyke
Department of Philosophy
Temple University
Philadelphia, PA 19122
Stanley N. Salthe
Department of Biology
Brooklyn College
City University of New York
Brooklyn, NY 11210
Eric D. Schneider
Chesapeake Biological Laboratory
University of Maryland
Solomons, MD 20688
Robert E. Ulanowicz
Chesapeake Biological Laboratory
University of Maryland
Solomons, MD 20688
Jeffrey S. Wicken
Behrend College
Pennsylvania State University
Erie, PA 16509
Biology and Philosophy 4: 373-405, 1989.

© 1989 KluwerAcademic Publishers. Printedin the Netherlands.
374 BRUCE H. WEBER ET AL.
Evolution in Thermodynamic Perspective: An

Ecological Approach
ABSTRACT: Recognition that biological systems are stabilized far from equilibrium by
self-organizing, informed, autocatalytic cycles and structures that dissipate unusable energy
and matter has led to recent attempts to reformulate evolutionary theory. We hold that
such insights are consistent with the broad development of the Darwinian Tradition and
with the concept of natural selection. Biological systems are selected that are not only more
efficient than competitors but also enhance the integrity of the web of energetic relations in
which they are embedded. But the expansion of the informational phase space, upon which
selection acts, is also guaranteed by the properties of open informational-energetic systems.
This provides a directionality and irreversibility to evolutionary processes that are not
reflected in current theory.
For this thermodynamically-based program to progress, we believe that biological
information should not be treated in isolation from energy flows, and that the ecological
perspective must be given descriptive and explanatory primacy. Levels of the ecological
hierarchy are relational parts of ecological systems in which there are stable, informed
patterns of energy flow and entropic dissipation. Isomorphies between developmental
patterns and ecological succession are revealing because they suggest that much of the
encoded metabolic information in biological systems is internalized ecological information.
The geneological hierarchy, to the extent that its information content reflects internalized
ecological information, can therefore be redescribed as an ecological hierarchy.
This thermodynamic approach to evolution frees evolutionary theory from dependence
on a crypto-Newtonian language more appropriate to closed equilibrial systems than
to biological systems. It grounds biology non-reductively in physical law, and drives a
conceptual wedge between functions of artifacts and functions of natural systems. This
countenances legitimate use of teleology grounded in natural, teleomatic laws.
KEY WORDS: Nonequilibrium thermodynamics, information, informed patterns of energy

flow, ecological hierarchy, genealogical hierarchy, succession, development, Darwinian
tradition, reductionism, teleology, natural selection
INTRODUCTION
It is universally conceded that biological systems conform to the con-

straints of the Second Law of Thermodynamics. But most workers have
not thought that this, or any other physical law, plays a positive role in
explicitly biological reasoning. Recently, however, the notion has gained
currency in some quarters that thermodynamical laws do indeed play an
informative role in undergirding biological, and especially evolutionary,
explanations. This opinion has gained ground in proportion as thermo-
dynamics itself has been expanded to deal with open systems stabilized far
from thermodynamic equilibrium. For biological systems at all levels of
scale ranging from cellular pathways to ecological systems possess to a
high degree the generic properties exhibited by all far-from-equilibrium
systems.
EVOLUTION IN THERMODYNAMIC PERSPECTIVE 375
In this connection, three highly general propositions have assumed

particular importance:
P1: Biological systems are stabilized far from equilibrium by way of self-organizing,
autocatalytic structures that serve as pathways for the dissipation of unusable
energy and material. Because biological systems are "dissipative structures" in this
sense, entropy production and organization are positively correlated (Prigogine,
1980).
P2: Biological systems that dissipate energy more effectively than others will proliferate
differentially over competitors, not only because they will utilize resources more
effectively than their competitors, but also because they will enhance the integrity
and persistence of the web of energetic relations of which they are themselves
parts. That natural selection may be conceived as serving this function implies a
restatement, rather than a refutation, of the Darwinian tradition (Lotka, 1922,
1925; Wicken, 1987).
P3: Informational complexity, in the form of genetic variation, is subject to the Second
Law, which forbids errorless replication for real dissipative systems with finite
energy sources. Over time, there is not only an exploration of genetic phase space
but an expansion of that phase space. The increase of genetic phase space so
guaranteed is an important explanatory factor in phylogeny (Brooks and Wiley,
1986, 1988).
The fact that P1 does not entail P2 and that P2 does not entail P3 suggests
that exploration of the relation between evolution and nonequilibrium
thermodynamics has not yet resulted in an articulated paradigm and that
much foundational work remains to be done. It is the opinion of the
authors of this paper that these inchoate ideas cannot be developed and
assessed adequately, or their implications for current evolutionary theory
set forth, unless the ecological, and hence energetic, framework in which all
such processes are set is given descriptive and explanatory primacy. We
mean this in at least the following three senses:
P4: Just as ecological systems are relational parts of the entire terrestrial biosphere, so
all the other levels of what is sometimes called the ecological hierarchy (Eldredge
and Salthe, 1984, Eldredge, 1985, Salthe, 1985) are relationalparts of ecological
systems. These include ecosystems, populations, organisms; and, within the eco-
nomy of the organism itself, cells and biological molecules. All of these are
relatively stable, informed patterns of energetic flow and entropic dissipation,
between which selection occurs at various levels.
P5: Biological systems exhibit diachronic developmental patterns non-trivially analo-
gous to those of ecological systems. The pattern of ecological succession is found
as well in the pattern of individual organismic growth.
The point here is not the discredited one that ecological systems are
"super-organisms," exhibiting patterns paradigmatically found in develop-
mental biology, but, on the contrary, that developmental patterns are
informed patterns of energy flow that have been in-corp-orated (literally)
by way of information-bearing macromolecules into metabolic pathways.
P6: Much, but not all, encoded information, which enhances autocatalytic processes by
increased replicative fidelity, is stored metabolic information and hence [by P5] is
internalized ecological information. The genealogical hierarchy that has been dis-
tinguished from the ecological hierarchy (Eldredge, Eldredge and Salthe, Salthe) is
itself (differently considered) an ecological hierarchy to the extent that its informa-
tion reflects internalized ecological information. Thus the fact that the branching
patterns of phylogeny depend on the Second Law does not by itself imply that this
process can be understood either causally or descriptively apart from analyzing
energetic pathways.
We put these points forward both as a contribution to theoretical
biology and with a view to suggesting that this perspective has important
implications for standing issues in the philosophy of evolutionary theory
and for the philosophy of biology more generally.
The fact that natural selection may be conceived as differential survival
of informed pathways of energy flow does not undermine the core (in
Lakatos' [1970] sense) of the Darwinian tradition. Darwinism holds
fundamentally that evolutionary change will occur whenever historially
conserved variation is accompanied by differential retention (Lewontin,
1974). Selection among differential kinetic pathways in accord with
thermodynamical imperatives fully meets these conditions. Thus these
ideas are not put forward to "falsify" Darwinism or to render it obsolete in
the light of a more general theory. On the contrary, our effort is to update
and enrich the Darwinian tradition by reorienting evolutionary science
around ecology, thermodynamics and information science in a way that
allows us more deeply to understand the underlying dynamics of natural
selection, while keeping population genetics as an integral part of the
expanded whole. We are thus conservatively attuned to the Darwinian
tradition, although we do not retain an expectation of the hegemony of
selective processes, but expect that in this richer context it is likely that
other evolutionary principles will be recognized as equally important.
What this reorientation does call for is recognition of the fact that the
Darwinian tradition, as it has hitherto been articulated, has been encoded
within a framework of high-level assumptions about systems dynamics that
can be called approximately Newtonian or equilibrial. Taken up by Darwin
through Lyell and Malthus, and reflecting the general cultural milieu in
which Darwin worked, equilibrial assumptions have been spectacularly
successful in organizing several centuries of productive research. Neo-
Darwinism is a research tradition that reflects this success. Its fundamental
principle, the Hardy-Weinburg Equilibrium Formula, announces its dy-
namical commitments on its face. It is true that neo-Darwinism reflects
a shift from the determinism of Newtonianism, properly so called, to
statistical descriptions and "population thinking," on the model, much
emulated by Fisher, of statistical mechanics. It nonetheless remains true
that the connection between equilibrium thinking and Darwinism is a
contingent one; and that further development of the Darwinian tradition
may now require loosening the grip of such paradigmatic structures:
P7: Because biological systems are stabilized far from equilibrium they are imper-
spicuously described, and only weakly explained, in a crypto-Newtonian language
appropriate to isolated or closed equilibrium systems.
One measure of progress in the development of any science is its ability

to make close contact with other sciences. In the present case contact is
urged between evolutionary biology and physics and informational science.
It is especially important to point out at the outset, therefore, that
P8: While we take a dim view of the 'autonomy-of-biology' position commonly de-
fended by neo-Darwinians (Mayr, 1982; 1985), the emerging paradigm espoused
here does not seek to "reduce" biology to chemistry and physics.
No science has a permanent, natural, defensible frontier, any more than

the Roman Empire did. Rather, just as the most productive estuaries in
the natural world are those most open to environmental exchanges, so too
the most productive science is that which looks for connection with, and
fertilization from, other disciplines. In the present instance, we seek to
expand biology by judiciously incorporating concepts from the physical
and informational sciences. Far from leading to the reduction of biology to
physics this effort effectively points out what is unique about the biological
world - the stabilization and articulation of energetic pathways by
informational macromolecules, and by the conservation and expansion of
functional information through phylogenetic time. It also changes and
expands the boundaries of physics and chemistry. It is, in short, an
interfield theory (Darden and Maull, 1977) rather than a reduction of one
theory to another.
This cross-fertilization is especially effective in two related areas. It
provides the deep causal principles underlying variation, constraint and
selection, and it allows us to incorporate the emergence of life within the
resultant framework. Darwin would have approved. The inability of the
science of his day to account for the existence of living organizations
vexed him. Reluctantly, he was obliged to take life as a 'given' in order to
move evolutionary thought onto a solid, scientific plane. Now, however,
we can talk about the self-organizing principles that lead to life's emer-
gence in a way that is entirely consonant with the broad Darwinian
schema referred to above.
This perspective has implications, finally, for problems about biological
teleology. The well known tendency of adaptationist explanations in
evolutionary biology to degenerate into illegitimate design-teleology or
"just so stories" represents an acknowledged weakness in current evolu-
tionary theorizing (Gould and Lewontin, 1979). This tendency comes
from viewing natural selection as an alternative means to achieving the
same kind of effects as those achieved by design and craft. Like crafts-
manship, natural selection seems on this view to presuppose a picture in
which raw materials are worked up into functional wholes that are them-
selves regarded as assemblages of decomposible and ontologically distinct

parts. Among the disadvantages of this assumption is that it sustains the
conceptual possibility of creationist arguments in a scientific milieu that
has overwhelmingly powerful empirical arguments against them. Our view
is that:
P9: When informed patterns of energy flow are treated as units of selection a con-
ceptual wedge is driven between the functions performed by artifacts and the
functions performed by natural processes, whose self-organization results in path-
ways for energy flow and entropic dissipation. This allows us to distinguish legiti-
mate from illegitimate senses of 'teleology', and to ground the former in natural,
teleomatic laws.
ECOSYSTEMS PHENOMENOLOGY, ASCENDENCY AND DEVELOPMENT
The focus of ecology is relational. Ecologists concern themselves primarily

with the processes that connect species and populations into symbiotic
relationships, food chains and other patterns. The fascination of ecologists
with homeostatic and mutally adaptational processes so evident among
communities of populations has given rise to the view of an ecosystem as a
"superorganism" (Cf. Clements and Shelford, 1939, and recently Lovelock,
1979). However, the scorn justly heaped on such notions by biologists and
philosophers of science, as well as the tendentious social and political
philosophy to which it lent credibility, led to a chastening of ecology. It
was eventually reintegrated into the neo-Darwinian Synthesis as a dis-
cipline in which ecological systems were construed as the outcome of
predator-prey relations governed by the Lotka-Volterra equation and of
other dynamics that could be accounted for in terms of competition
among individual organisms for scarce resources. Group selection hypoth-
eses were discredited. With these developments, however, the emphasis of
early ecologists on the community as a pattern of energy and matter flow
was also lost. Attention shifted, under the influence of neo-Darwinian
ideas, toward "diversity of population levels" as a surrogate for diversity of
flows.
Modern ecosystems phenomenology traces its origins to the students of
G. E. Hutchinson, who emphasized the importance of measuring palpable
transfers of material and energy among the elements of a living com-
munity. Lindeman (1942) sought to portray the influence of the Second
Law as a constraint that ordered the "trophic pyramid" inherent in the
energy flows occurring in a shallow water lake ecosystem. With his death it
fell to Eugene and Howard T. Odum to advocate the measurement of
networks of flows occurring in ecosystems. Early among Eugene Odum's
(1953) observations was the notion that ecosystem homeostasis could be
maintained through the existence of compensatory flow pathways. That is,
if more than one concatenation of transfers linked two arbitrary species in

a community, the unexpected disruption of one or some of those pathways
might be compensated by augmented flow through those linkages that
remained relatively unperturbed.
Eugene Odum took advantage of the flow compensation idea to set
forth an entire suite of observations on the development of ecosystems.
He listed twenty-four attributes that were correlated with mature com-
munites in the later stages of succession (E. Odum, 1969; Schneider,
1988). These are combinable into five categories. Mature communities
tend to possess:
i. increased energy flow
ii. greater variety of species
iii. more narrow trophic specialization by their members
iv. enhanced amount of cycling
v. longer retention of media in the system.
At the heart of this ordering process stands the kind of positive
feedback inherent in autocatalytic cycling studied by Eugene and Howard
Odum (Odum and Odum, 1982; Odum, 1988). In its simplest idealization
autocatalytic feedback may be thought to result from a concatenation of
positive influences, where one item in the chain catalyzes another. In the
ideal cases, the causal links in the loop are the only ones present, and the
system appears as a wholly autonomous structure. Of course, real causal
loops are always embedded within larger networks of causalities. If one
observes only some of the elements of a loop, they will appear to function
in a non-autonomous fashion at the behest of the boundary conditions that
drive them. However, once the scale of observation is enlarged to include
all members of the loop, its overall autonomy then emerges as an attribute
of the expanded system as a whole. Seen in this light, the extensive effect
of a feedback loop is to enhance the aggregate system activity. Thus an
increase in the activity of any element in the loop will propagate to
augment the activities of all loop members, including itself.
Seen functionally, the role of autocatalytic feedback is to exert selection
pressure upon its own members and to compete with neighboring loops
for system resources. Any change in the properties of a loop element that
abets its action upon the next member will be rewarded in the next pass.
Conversely, changes that diminish autocatalytic activity are self-inhibiting,
and the elements in such loops are subject to replacement by newer, more
effective components. One can readily imagine, then, a situation wherein
all the original members of a loop are replaced in turn, while the loop
itself persists beyond the more transitory durations of its components. The
same feedback cycle will then have played an active role in influencing
what the replacement parts would be.
The overall directional effects of positive autocatalytic feedback loops
in networks of flows define the ascendency of the system (Ulanowicz,
1986). Ascendency quantifies the total energy transfer in the system and
the level of the interconnectiveness among the components of that eco-
system. Change in ascendency reflects the effects of positive feedback as
an-agent in the successive restructurings of the system. Ascendency is
measured by dividing the ecosystem into compartments that serve as
nodes of the network. Directed graphs are then drawn indicating flows.
The mathematical techniques required to estimate ascendency are a
combination of Leontieff's input-output analyses with information theory
(see Appendix I for a description).
Of course, no system can grow and develop without bounds. Against
the inherent tendency of systems networks to become ever more articu-
lated stands the vulnerability of such configurations to perturbations.
Redundant pathways offer a certain "strength-in-reserve" by virtue of the
compensatory rerouting they afford. If a perturbation is entirely new to a
community, the system's ability to adapt successfully to it will hinge on
the availability of residual, less effective pathways, or even on outright
stochastic configurations (Von Foerster, 1960; Atlan, 1974; Conrad,
1983). Losses from the system and the multiplicity of inputs also detract
from the rise in ascendency. It is, in fact, never possible to eliminate
completely all exogenous transfers. Some of the dissipative losses are in
fact necessary to maintain structure at lower levels; and some of the
exports to other systems from links in higher-order reward loops. Finally,
a system with but a single input is highly vulnerable to disturbance in that
source. Hence, that external factors will be the most pronounced factor in
a system's fate follows from the very same principle in virtue of which
ascendency rewards the increasingly modular stability that we refer to as
hierarchical. Each of these influences detracts from the rise in ascendency.
These detractors can be quantified in mathematical terms similar in form
to those used to measure the ascendency itself. It was, in fact, to quantify
such "strength-in-reserve" that Rutledge et al. (1976) first correctly
applied information theory to flow networks.
Let us look at a number of phenomenological patterns in ecological
succession in the light of these principles:
P10: Ecosystems favor species that, in funnelling energy into their own production and
reproduction, also increase the total energy flow through the ecosystem.
P1: In succession there is a maximization of energy flow into organizational progaga-
tion and a minimization of metabolic costs.
P12: After an initial increase, a monotonic average decrease in the intensity of energy
flow (i.e. a weight-specific energy flow) through an open system occurs.
P13: A continual, hyperbolic increase in complicatedness (= size + number of com-
ponents + organization) develops.
P14: An increase in internal stability occurs as the rate of development slows down.
P15: There is a corresponding decrease in stability to perturbations.
P10 is a statement of Lotka's power principle (Lotka, 1922, 1925). P11

and P12 together might be thought of as the allocation of a given energy

flow to an increasing number of parts insofar as these systems grow. It
may also, to some degree, be a consequence of P13 when the decrease
mentioned in P12 becomes absolute. P13 is the locus for discourse about
macroscopic information and its increase. P12 and P13 are a result of the
action of the Lotka/Odum maximum power principle, which is fully
realized only in immature stages. P14 could be a consequence of P12 or
P13, either separately or together. P11 through P14 reflect the influence
of Prigogine's principle of minimum entropy production, which only
begins to be realized in these systems during senescence. P14 and P15
together define senescence.
These rules of succession are also applicable to the global ecosphere,
which is itself a thermodynamically open system that processes radiant
energy from the sun, and exports what is not retained as heat to the sink
of space. In a broad sense we note that a decrease in the intensity of
energy flow across the surface of the earth has taken place since Hadean
times (P12), when the planet was much hotter, was unscreened from the
sun, and was subjected to numerous impacts from bodies that would
subsequently have burned up in the atmosphere. Meanwhile, an increase
in stored information has occurred (P13) - first with the separation of
phases in the Hadean period, then with the gradual separation of ores,
then with an increase in the diversity of living forms, and finally, just a
little while ago, with the increase in the complicatedness of human artifact-
mediated production. Whether this process has been, or will be, accom-
panied by an increase in internal stability and a decrease in stability from
external insults (P14-15) remains an open question.
We may also see the same phenomenological rules of succession at
work in the development of social systems, presumably among all social
animals but in a particularly vivid way in human societies. Cities, for
example, are dissipative structures (Dyke, 1988b). They exhibit universal
patterns of economic self-organization in which internal complexity,
including the peculiar kind of information contained in cultural meanings
and other ideological forms, results in progressively more effective ways of
directing energetic flow through the system (Odum, 1988). Cities recur
and persist as the chief nodal links between the kinship systems out of
which they arise and the national and transnational economic systems into
which they are (sometimes) linked. The history of their rise, competition
as attractors and their eventual, inevitable decline closely parallels the
phenomenological rules we have traced.
These phenomenological symmetries between ecosphere dynamics,
ecosystem succession and social evolution are striking. They are, however,
far less striking than the fact that the patterns seen in P1O-P15 are also
observable in, and applicable to, the development of individual organisms
(Zotin, 1972). The first three rules (P10, P11, P12) indicate that the
organism gradually becomes more energetic, while its specific metabolic

energy decreases over most of its life. The fourth (P13) indicates that our
descriptions of an organism's energetic pathways become ever more
cumbersome as it develops. The fifth (P14) indicates that an organism's
internal tendency to change tends to dwindle with age. Finally, the sixth
(P15) rule suggests that, as aging continues, the organism is more and
more easily disrupted by fluctuations in its environment. Viewed as a
whole, these rules suggest that the stored information curve is a kind of
inverse of the energy intensity curve. While the organism is changing most
rapidly, it, like the ecosystem, has the greatest specific energy flow through
it. On the other hand, increasing complexity results in less flexibility,
curtailed metabolic rates and an increased vulnerability to insults. The
organism's ability to heal or regenerate as it works on an ever increasing
number of homeorhetic problems decreases until a limit is reached and
the organism is recycled.
These considerations have suggested to some the old, but recurrent
view that the larger systems within which organisms are contained are
developmental entities, or "superorganisms," in their own right. This is a
misleading way of calling attention to the often-neglected reality of
systems above the organismic level, such as the stable feedback cycles
between the atmosphere and the biosphere remarked on by proponents of
"the Gaia hypothesis," as well as to the fact that organisms are embedded
in these larger systems as relational parts. The fact is, however, that
organisms are the proximate and proper locus of replicating molecules,
and that the greater integrity that we see in organisms, as compared to
higher-level ecological units, is not just a matter of our own epistemic
perspective, but instead reflects the powerful agency and real effects of the
appearance of replicating macromolecules. It is organisms alone, or at
most certain informational-sharing symbiotic systems, that have employed
informational macromolecules to achieve stable, homeostatic and homeor-
hetic metabolic pathways, together with reproductive fidelity. For ability to
do these things is the very definition of an organism. An organism is an
informed autocatalytic system possessing, in virtue of information stored in
macromolecules, an internal organization of kinetic relationshipssuch that
it is able to maintain itself by pulling environmental resources into its own
production and faithful reproduction and by dissipating unusable energy to
appropriate sinks (Wicken, 1987). Thus in interpreting the observed
symmetry between the rules of ecological succession and the patterns of
organismic development it is far more informative to speak of organisms
as highly integrated, homeorhetic ecosystems than to speak of ecosystems
as superorganisms. This view carries with it the suggestion that organisms
originated, and continue in part to evolve, by (literally) incorporating-
storing and deploying in developmental programs - information that, in
the first instance, describes patterns of energy flow in more generic kinds
of autocatalytic cycling systems. It also implies that organisms continue to

be informatively described as autocatalytic systems even after the trans-
formation brought on by replication, with the greater sources of variation
and internal structure which that entails, and that these facts are still
causally relevant to their evolution.
AN ECOLOGICAL APPROACH TO EVOLUTIONARY DYNAMICS
How might these things have occurred - and why? We may push these
phenomenological reflections in a causal direction by noting, in the first
instance, that the patterns noted in P10-P1 5 conform to the fundamental
propositions of open systems hermodyamics. We can reformulate P10
and P11, for example, in a more causal way by asserting that:
P16: Natural selection will favor species that, in funnelling energy into their own
production and reproduction, also increase the total flow of energy through their
ecosystems;
and that
P17: Natural selection includes (in addition to this flow-expanding principle) an effi-
ciency principle, which promotes a maximization of flow into organizational
progagation and a minimization of flow into metabolic costs.
These two principles make opposing contributions to the organization-

dissipation relationship. P16 implies a teleomatic drive to expand the
energetic economy through evolutionary innovations that multiply kinetic
pathways for dissipative flow. P17 leads to specialization and mutualism:
In unperturbed ecosystems specific dissipation (per unit of biomass)
decreases as the elaboration of mutualistic networks under selection
pressure increases biomass/throughput ratios. These principles are yoked
together by way of the deeper principles of nonequilibrium thermody-
namics. As developed by Prigogine and his school, these state that open
systems will dissipate energy as entropy through self-organizing kinetic
pathways. These emerge from spontaneous, autocatalytic cycling (P1).
Seen in this light Lotka's principles (in the form of P16 and P17) provide
an explanatorybacking for the phenomenology of P10-P 15.
This approach implies that natural selection is best conceived within a
framework of physical law. Individuals, in the normal sense of organismic
units, will not be the paradigmatic units of selection, as they are on the
traditional, morphologically-based conception of natural selection, which
retains echoes of discourse about craftsmanship and design. Rather, like
ecosystems, individual organisms will be construed as informed patterns of
thermodynamic flow. On this view cells, organisms, populations, and
ecosystems will all be units of selection, but it is ecosystems that will
constitute the paradigm case and the matrix within which the others arise.
Accordingly, the language in which natural selection is to be conceived,
and in which adaptative scenarios are to be couched, is a language of
process and pattern rather than of products and entities. In this language
what is selected are not parts, in the morphological-artifactual sense, but
differential patterns of energy flow. Some flow patterns will necessarily be
superior to others in commanding resources, and will be selected on that
basis.
The concomitant reconception introduces a dimension of directionality
into the notion of fitness. Those populations are fittest that best enhance
the autocatalytic behavior of the reward loops in which they participate.
The effect of such competition is an emergent tendency during the later
stages of ecosystem development toward more highly articulated networks
of flow, wherein those pathways that foster more efficient transfers
flourish at the expense of less effective routes.
Three immediate implications of this reconceptualization are worth
noting. First, the competitive success of flow patterns is, at least in part,
explained in terms of self-organizing systems dynamics (Csanyi, 1985),
rather than being ascribed, as it generally has been by the Darwinian
tradition, exclusively to externally imposed forces. It is true that initial and
boundary conditions, such as limiting resources, are important factors. But
causality is not to be conceived in the first instance as a matter of external
pressures being exerted on systems that would otherwise remain un-
changed.
Second, flow patterns are selected because they contribute to the
stability and coherence of the systems of which they are a part. Thus
individual interest is contextualized within community dynamics. The
nitrogen cycle illustrates this well. Plants take up nitrate from the soil and
convert it to amino acids and nucleotides, then to proteins and nucleic
acids. Decomposers hydrolyze these polymers, then deaminate the mon-
omers, releasing ammonia. This serves as an energy source for Nitrosomas
bacteria, which oxidize it to nitrite, which is in turn oxidized by Nitrobacter
to nitrate. Each microbe is obviously in business for itself, exploiting
energy sources for survival and reproduction. But it is the ecosystemic
cycle to which they contribute that makes these niches available, and it is
their activity within this cycle that sustains the continued self-maintainance
of the large flow pattern. In this picture, constraint and selection cannot be
conceived apart from each other, or as opposing forces that are somehow
balanced off. Rather the fact that the competitive success of higher-order
ecosystem flow imposes constraints on lower-order entities is an essential
component of the very possibility of isolating units of selection and
ascribing adaptations to them. To the theme of selection, as it has emerged
thus far within the Darwinian tradition, therefore, we do not propose to
add a new theme of constraint. Instead, the theme of constraint makes the
theme of selection coherent and complete for the first time. However, not
all traits need be selected. Some can arise through neutral molecular
evolution (Kimura, 1983), through endogenous characteristics of genetic
regulatory patterns considered as statistical ensembles (Kauffman, 1985;
see also Demongeot et al., 1985), or through exploration of the possi-
bilities of developmental alternatives such as neoteny.
Third, in this context units of selection, as they appear in the writings of
those who advocate an "expanded synthesis" (Gould, 1982), are neces-
sarily members of an ecological hierarchy. For cells, organisms, popula-
tions, ecosystems and regional biota are all phase-separated systems of
thermodynamic cycling and flow; and cycling and flow will proceed most
efficiently and stably by hierarchical or modular organization (Odum,
1988). Thus, recent calls for an hierarchical expansion of the Darwinian
tradition may now be seen to be grounded in the deep structure of
evolutionary theory. Recent hierarchical thinking is not called for, as it
may have appeared at first, merely because the need has been felt for
conceptual models that usefully accomodate new data. Rather hierarchical
structure turns out to be a basicfact about the biological world.
To these considerations we may add the important consideration that
the vexing problem of the origin of life can be productively and sugges-
tively conceptualized within this framework. From the conventional point
of view one must begin either with proteins or with nucleic acids, and
then wonder how to get them together into a faithful, functioning self-
replicating system, even if we assume that the RNA has catalytic activity.
A bootstrapping problem then occurs. As a result the considerable
conceptual and explanatory resources of 'hypercycle theory' fail to be fully
exploited. However, within the ecological, thermodynamic context that we
have been exploring, hypercycles now can be generalized as autocatalytic
cycles of selection and flow within which nucleic acids and proteins are
seen to co-evolve, along with energy metabolism, under the imperative of
physical laws, possibly in the phase-separated microspheres (Fox, 1984)
or in an amphiphile bilayer vesicle (Morowitz, Heinz and Deamer, 1988)
that could provide a proto-organismic setting for the acquisition of genetic
information under thermodynamic flows. Accordingly, the chicken-egg
problem, which Darwin bracketed long ago by taking living organization
as primitive in his scheme, is brought within sight of a solution.
The source of the coevolutionary nexus in the thermodynamic space
within which life emerges can readily be shown without getting into
equations. Thermodynamic flows require specific mechanisms in order
to achieve irreversibility. Those in turn require cycles. For example, a
photoreceptor X, excited to X*, must have some dissipative route back to
the ground state to command irreversible energy flow. The emergence of
informed dissipative routes is synchronous with the emergence of living
organization. Proteins and nucleic acids evolved informationally in this
dissipative setting. Consider the following dissipative route (Figure 1):

Protoreceptor X delivers excitation energy to amino acid phosphorylation,
indicated by A-P. The excited monomers then polymerize to poly-
peptides PP. If these feed back to catalyze either step leading to their
production, as indicated by the arrows in Figure 1, then one has an
autocatalytic system capable of pulling energy resources into the pro-
pagation of that polypeptide. Such capacities have been demonstrated
experimentally (e.g. Fox, 1984). Abiotic polypeptides, operating in phase-
separated microspheres, also absorb polynucleotides, and engage them in
mutually-stabilizing proto-ribosomal complexes. Since lysine-rich poly-
nucleotide complexes have catalytic activities for both amino acid and
nucleotide polymerization, one can expand the scheme as indicated in
Figure 2. Here, a nucleic acid cycle has been coupled synergistically with
the protein cycle. In this diagram T represents nucleic acid template, N
nucleotide monomer and N-P phosphorylated nucleotide. PP catalyzes
the formation both of itself and template T. T provides the template for its
own replication and informs the production of the catalytic polypeptide.
The system as a whole becomes a nucleus for acquiring information
through competitions for energy fluxes. The chicken-egg problem is solved
in principle, if not in historical particularity, by giving nucleic acid and
protein equal ontological status within an information-acquiring dissipative
package, thus projecting the ecological/genealogical interaction dynamics
back to the origin of life.
The solution to this problem is, moreover, of a piece with an adequate
theory of subsequent evolutionary dynamics. We have defined a living
thing as an informed autocatalytic system that sustains itself by environ-
+ Pi
X*A
Fig. 1. A hypothetical autocatalytic cycle involving phosphorylation, polymerization, and

hydrolysis where polymer PP feeds back positively to either of its precursors.
7A A P
PP A-P
E S
N-P
NP NI
Fig. 2. A minimum 'realistic' autocatalytic cycle in which template T and polypeptide PP

work synergistically to pull resources into themselves and degrade energy.
mental energy exchanges and that varies under the drive to configurational
randomness. Statistical thermodynamics provides two important principles
of randomization that connect life's emergence with its subsequent evolu-
tionary diversification. One is the degradation of free energy to heat. This
principle leads, in the first instance, to the 'chemical imperialism' of living
matter, by which organisms are able to funnel resources into the perpetua-
tion and propagation of their own organizational types. The other is the
drive to configurational disorder, which ensures that new configurations of
matter will be delivered into the testing ground of the ecological arena.
Mutation and sexual recombination both derive from this entropic drive
(Wicken, 1987). Accordingly:
P18: The principles of variation, constraint and selection that began their operation in
life's emergence, and were themselves responsible for that emergence, are also at
work in biological evolution.
As successful patterns of dissipation emerge, so too does a tightening of

the nucleic acid-protein relationship. With that tightening comes the
constraint on variation characteristic of the biological world. The inor-
ganic world thus flows smoothly into the organic world, and genetic
information accumulates under thermodynamic selection for stable pat-
terns of entropy production. The internalization by natural selection of
informational possibilities stabilizes the energetic (i.e. metabolic) flow
patterns, and thus becomes metabolic information. This stabilization
defines the boundaries of the organism. But this metabolic information is
in fact selected because it allows organisms to function as relational parts

of ecological wholes: It has in part been selected because it stabilizes the
network of ecological relationships. Thus, to the extent that it reflects this
process, metabolic information is internalized, or, as we have stated
previously, incorporated ecological information (P5). As metabolic in-
formation accumulates, dissipative flows are guided into new creative
channels, which reduce specific entropy production in ever more complex
networks through the generation of information space in the form of
variational possibilities. The operation of such complex whole systems
physically requires the production of yet more variation, or informational
possibilities, in a new selective round. This in turn makes possible new
ecological pathways and niches. (The idea that the environment shapes
the informational content of the genome through feedback interactions
between genotype and phenotype has also been explored by Riedel
[1977]. This concept does not conflict with the traditional Darwinian
insistence that variation is random in relation to selection. The feedback
loop runs over the long run. However, if phenomena violating the normal
Darwinian rule are confirmed, the view expressed here will be able better
lo conceptually absorb this than traditional Darwinism.)
ENTROPY, INFORMATION, AND GENEALOGY
The relation between the Second Law and the kinetic phase space
represented by metabolic processes is subject to much controversy. No
neo-Darwinian has denied that organisms must conform to the dissipative
requirements of the Second Law. But when units of selection are thought
of as analogous to machines few interesting connections between natural
selection and the Second Law appear. By envisioning natural selection as
occurring among informed patterns of energy flow, however, this paper
makes that connection more perspicuous and tighter. From this perspec-
tive there will be deep reasons for expecting that evolution, and phylo-
genetic branching, will occur, since the Second Law will demand pathways
down which entropy can be dissipated. Despite this advantage over
neo-Darwinism, however, one might still hold on this view of the matter
that the Second Law merely provides the possibility space within which
selection at the level of metabolic processes, and evolutionary diversifi-
cation, are to be construed, and that little if any causal or explanatory
force is to be given to the Second Law itself. Alternatively one might hold
that there is a non-trivial causal reciprocity between the expansion of
information space demanded by the Second Law and the kinetic-
metabolic processes that embody and manifest that expansion. If kinetic
pathways are means for the dissipation of entropy, we can, on the one
hand, say that the fitnesses of these pathways are the proximate cause of
diversity, given the background requirements of the Second Law. But it is

also informative to say that in selecting pathways imperatives of the
Second Law explain selection and diversification as a whole. From this
largest perspective, the Second Law has causal implications. But at the
detailed level the causal accent must be on kinetics.
Might one say that the quantity that increases is not entropy, at least in
the energetic sense, but rather information space, and that the Second Law
is causally responsible for this quite apart from whatever function is being
performed at the energetic level? This view has been vigorously advocated
by Brooks and Wiley (1986, 1988), who argue as follows. The concept of
entropy can be generalized beyond its original energetic and later statis-
tical senses to an informational definition. This makes the growth of
information in genetic systems the expected result of the operation of the
Second Law without requiring direct reference to energy flows. Shannon's
concepts of transmitter, channel and receiver are extended to biological
systems and Gatlin's notions of divergence from maximal informational
"entropy" is hierarchically expanded. Unlike Shannon entropies, however,
which can be defined formally, the informational entropies being meas-
ured here are said to be physically grounded in biological systems (Brooks
and Wiley, 1986, 1988; Brooks, Cumming and LeBlond, 1988; Collier,
1986). In Gatlin's work, total information cannot increase over phylo-
genetic time. In Brooks and Wiley's theory it can. Accordingly, they follow
Layzer (1977) in holding that information is the lag between the unlimited
expansion of the possibility phase space, Hmax, and constraints, Hobs. This
lag, which is seen as the paradigm of entropic processes generally con-
sidered, recurs at every level of biological processing. The hierarchical
information theory so developed represents a generalized expression of
the Boltzmann entropy measure considered as arrays under constraint.
Although Boltzmann entropy measures were originally invoked for
energy transformations, Brooks and Wiley view the most general expres-
sion of the Second Law as having only a contingent relationship to energy-
based processes. They argue that energy transformation in entities in the
ecological hierarchy is a necessary condition for the expansion of bio-
logical information, but that the information passed on is not in itself a
measure of energetic processes. On this view, the ever-increasing noise
due to errors in replication results from species acting as individuated
"information-dissipative" structures, driven by the Second Law. This
process is thought to be sufficient to account for diversity. Thus it is
thought to be reasonable, indeed necessary, in order to get closure
conditions for constructing phylogenies, to ignore input from environ-
mental boundary conditions. Any deviation from the expected informa-
tional distributions is then attributed to the action of natural selection,
energetically considered, which affects the rate of cladogenesis and prunes
the variation so produced. New -channels become forever separated, and
"noise" on the overloaded channels is reduced. These concepts are made

operational by way of the notion of breakdowns in reproductive "cohesion
entropy". This is said to explain, as well as to measure, the informational-
entropic bifurcations that now appear as cladogenesis.
This attempt to generalize statistical entropy to informational entropy is
based initially on the fact that both formulations use a similar probability
theory (Khinchin, 1957). However, we must go back one step and
consider the relationship of thermodynamic to statistical entropy. The
reason that the Boltzmann microstate/macrostate ensembles are entropi-
cally relevant is precisely because of their relevance to the energetics of
the systems studied and hence to the amount of work that can be gotten
out of such systems. The microstates are energy levels, and the distribution
over these levels is constrained by the requirement of the conservation
of energy. Not every physical microstate/macrostate distinction that
embodies a principle of constraint will yield entropy calculations in the
thermodynamic sense; only those that are appropriately connected to
energetics will. Entropy is not a measure of every sort of disorder, but
only of energetically relevant disorder. Disorder in formal, non-physical
systems is often lumped into a Shannon "entropy," but what relationship
this would have to a true entropy is not clear. This is not meant to deny,
however, that a formal analysis of disorder is useful in formal systems -
or in physical systems, such as Brooks and Wiley envision, for which the
connection to energetics is only a necessary precondition for the existence
of the informational arrays. The question rather becomes one of the
adequacy and robustness of this formal analysis for the particularprob-
lems of interest.
The perspective that we have been advocating entails that information
remains tied to energy transformation and that encoded metabolic infor-
mation is a reflection of environmental boundary conditions. Information
measures relationships and pathways among such flows. For certain
analytical purposes, when tracing phylogenetic patterns, we might concede
that it is reasonable to disregard the fact that genealogical information is,
to a large extent, encoded ecological and metabolic information. Since
cladistic analysis is now generally accepted as the correct way to construct
phylogenetic trees, the use of equations from information theory is
justified if it improves phylogenetic methods and their applications to
biogeography and historical ecology. This does not, however, imply that
the relationship between information theory and thermodynamics pos-
tulated by Brooks and Wiley (which might be characterized in its putative
general form as "infodynamics") has been proven, although it supports the
intuition that information and thermodynamics are somehow connected.
What the relationship of information and thermodynamics actually is
has been a vexing question. Since the 1940s, information engineers and
some general systems theorists have intuited that there is a definite
relationship between thermodynamics and what Shannon called infor-

mational "entropy". This intuition was informally extended initially by
Crick to macromolecular function and more formally to biological systems
in general by Gatlin (1972) and Riedel (1978). More recently, Brooks and
Wiley (1986, 1988), Collier (1986), Wicken (1987), and the contributors
to Weber, Depew and Smith (1988) have explored the implications of
using an enriched thermodynamics of the nonequilibrium sort, fused in
some fashion with information theory, to reformulate evolutionary theory.
Nonetheless, the nature of the relationship of thermodynamic and statisti-
cal concepts of entropy to informational or Shannon entropy remains
problematic (see Peacocke, 1983; Denbigh and Denbigh, 1985; Depew
and Weber, 1988). It is still unclear whether information represents a
generalization of earlier concepts of entropy (Brooks and Wiley, 1986,
1988; Collier, 1986); or is a distinct concept connected to entropy, but
not in a linear fashion (Ferracin et al., 1978; Wicken, 1987); or whether
information and entropy can only be linked in a metaphorical manner
(Stuart, 1985); or whether, finally, present information theory is inad-
equate for the task of describing information and energy transformations
in living systems (Olmsted, 1988).
It might be said that one motive for generalizing entropy to an
informational definition that is not so closely tied to energy flow is that
thermodynamic entropy is strictly defined only at equilibrium, and that the
widely applied minimum entropy-production rule for steady-state systems
is only rigorously applicable to isothermal systems near equilibrium and
subject to weak energy gradients. Entropy therefore seems to be measur-
able, and therefore meaningful, only for such a restricted class of systems.
It must be acknowledged that those of us who want conservatively to keep
entropy tied to energy flows in far from equilibrium systems should keep
this point in mind. Those wishing to maintain a connection of entropy to
energetics must be willing to explore the likelihood that since living
systems are in fact operating far from equilibrium they are, and must be,
dissipating entropy, even if no simple measure of it is available. Those
with realist preconceptions about scientific theories will have much less
trouble with this than those who are wedded to forms of operationalism,
where failure to find an adequate measure of some quantity is tantamount
to failing to define the concept itself. Concrete measures of information,
such as those used to measure ascendency in ecological systems, are,
however, closely enough linked to energy flows that they are useful
instruments in approximating energy processing information in far from
equilibrium systems, such as ecosystems or metabolic processes.
Biological information processing is itself a physical process. Our ability
to manipulate information by the physical processes of a computer
(Landauer, 1988), which has already led to such powerful suggestions and
results, may lead us astray in this connection, for the computer is designed
to transmit and process information in a particularway, availing itself of

one - but only one - model of physical information transactions. The
macromolecular interactions involved in DNA replication, transcription,
and translation provide, however, a point of intersection between ther-
modynamic and informational approaches in which it is clear that the
particular physical grounding is not the same as that of a computer.
Further, other aspects of metabolism, such as informational processing
(integration of non-binary signals rather than receiving a message) by
allosteric enzymes and cell receptors, along with the more complex
operations of the endocrine and neuronal systems (especially the axonless
neurons), need to be explored as unique kinds of biological informational
systems. Similarly, developmental programs in DNA appear insufficient
fully to provide all the information needed for ontogenesis; metabolic
gradients and environmental factors are also important (Stent, 1985).
More speculatively, even DNA in contemporary organisms may not be
immune to informational transformations due to the action of metabolism,
and through metabolism to signals from the extracellular environment,
as presented in the concept of "profane genes" (Campbell, 1982, 1985;
Campbell and Perkins, 1988). All these phenomena fall within the
purview of metabolic information and can be investigated profitably from
an ecological perspective. An approach limiting itself to genetic infor-
mation considered in isolation, on the other hand, will not pick them up.
It must also be borne in mind that every informational component in
biological systems acts as both a channel and a receiver. Thus the
phylogenetic message cannot be disembedded from its metabolic matrix,
both molecular and ecological, except heuristically for analytical purposes
(P6). In our view, then, biological information has to be regarded as
"interpretive" information: Biological information is information about
function, rather than about mere signal differentiation. Introducing the
concept of functionality into the concept of biological information does
not imply that there must be a subjectivity in the "receiver" to "get" the
message. Nor does it imply any intentional teleology. It merely reflects the
fact that functionality, and the kind of meaning that we ascribe to
functions, is a real, objective property of all biological systems. Function
appears in a context - of metabolic pathways within the cell, of cells in
organisms, of organisms in populations, of populations within com-
munities, and so forth. From this perspective, the Second Law is causally
relevant, requiring selection among alternative kinetic pathways for energy
dissipation and telling us that we are entitled to expect that variation and
cladogenesis will occur, but telling us also that investigation of why it
occurs in particularpatterns in particularcases requires attention to more
particular ecological and metabolic facts.
Ultimately attempts to apply information theory to biological systems
require boundary and closure conditions, yet no single stable set of
boundary and closure conditions can be imposed on biological systems

without serious distortion. Thus to use informational entropy constant
care must be exercised to specify what boundary and closure conditions
are assumed, and to recognize corresponding limitations upon inter-
pretation and generalization. Any heuristic closure condition that is
imposed will give quantitative expressions that will be interpretable only
within the experimental environment defined by the closure heuristics
themselves (Dyke, 1988a). That is an inevitable consequence when we
cross the barrier that separates the science of complex systems from the
science of the simple that has, thus far, provided our culture with its
paradigms of successful theorizing. We should expect that new approaches
will be needed as we try to account for organized complexity instead of
organized simplicity (Newton) or disorganized complexity (Boltzmann),
(Serra et al., 1986; Serra and Zanarini, 1986).
REDUCTIONISM, TELEOLOGY AND THE NEWTONIAN PARADIGM
The ecological-energetic perspective presented in this paper informs the

philosophical claims made in P7-P9. We hold in the first instance that the
far-from-equilibrium character of biological systems allows the Darwinian
tradition to be more robustly reformulated by relaxing Newtonian and,
more generally, equilibrial background assumptions. Once such a shift in
background view has occurred, vexing problems of reduction and tele-
ology may be seen in a new light.
We assume here that successful explanatory theories rest, in part, on
highly general, often tacit pictures about the way things are. When such
pictures have proven successful in one field they are often transferred to
others, and sometimes come to pervade an entire culture, on its cognitive
side, forming its peculiar sort of common sense. Following Laudan (1977),
we will call such pictures background assumptions. In this light, Dar-
winism has from its inception been presented and defended in terms of
background assumptions associated with Newtonian systems-dynamics.
These background assumptions were brought into the study of evolving
biological systems by Darwin's encounters with uniformitarian geology
and English economics. Darwin's theory can be described, from this
perspective, as an attempt to extend Newtonian background assumptions
from Lyell's Newtonian or "uniformitarian" geology to biology with a little
help from the Malthusian version of Adam Smith's economics, which had
already brought the same background assumptions to bear on economic
systems.
The entities that enter into a Newtonian system, whatever they may be,
are assumed to exhibit inertial motion or rest. Deviation from this state
occurs only under the influence of impressed forces. At each instant the
system will tend to restore a dynamic equilbrium between inertial motion

and impressed force. What plays the role of inertia in Smith's case is the
assumed tendency of each economic agent to maximize command over
scarce resources. The laws of supply and demand lead to a constantly
threatened, and constantly renewed, dynamic equilibrium by pulling each
agent back toward others. By envisioning the environment as a closed
Malthusian system characterised by intense competition, Darwin suggested
what force could account for the modification of lineages of living things
over time. The intrinsic rate of population increase - a property of
organisms no less deeply embedded in their natures than is inertial mass in
Newtonian entities - is countered by the force of scarcity, an analogue of
gravity that every body exerts on every other. To the extent that each
competing entity is unconstrained by prior relationships to others, herit-
able variations will be indefinitely protracted into adaptive changes in
lineages that increasingly mirror the demands of a stern environment. In
this way Darwin more or less successfully refuted the standing presump-
tion that no natural law could account for the existence of functional
organic traits.
Newtonian systems have the following conceptual presuppositions:
They are closed, deterministic, reversible and decomposible. They are
closed because extrasystemic inputs would dissipate the calculable effect of
forces in determining state changes within the system. Malthus's argument
about populations illustrates the explanatory role of closure conditions
by showing how scarcity, the closure condition of classical economics,
can cause economic equilibrium to occur at a point below that where
exchanges are correlated with the biological survival of all the entities in
the system. Newtonian systems are in addition deterministic: Every state of
the system is fully determined by its prior states. Moreover, Newtonian
systems are reversible. For the laws governing state changes can be
calculated in both direction. Thus every prior or subsequent state of the
system is in principle recoverable. There is no inherent arrow of time in
a Newtonian system. Finally, reversibility implies decomposibility. A
Newtonian system must be atomistic, for if every state change is in
principle recoverable every structure in the system must be an aggregate
of least units that can be assembled or disassembled by whatever forces
are currently operating.
The neo-Darwinian Synthesis retains many of these conceptual pro-
clivities. Sober, for example, has recently provided an analysis of the key
concepts of the Synthesis in which a Newtonian concept of force is fused
with the notion of equilibrium in the classical thermodynamic sense.
According to Sober (1984), the Hardy-Weinberg Equilibrium Formula
provides the inertial baseline against which are measured the effects of
impinging forces on a system that would otherwise remain in the same
state. The system will not change its state until it is deflected onto a new
path by the imposition of external forces, such as selection. In this recon-

struction equilibrium, and deviation from it, are redefined in statistical
terms. For just as in physics classical mechanics gave way to statistical
mechanics, so in evolutionary biology neo-Darwinism meant, in part, a
shift away from deterministic closure at the phenotypic surface to a
two-level statistical reformulation in which gene frequency distributions
underlie and cause phenotypic macrostates. (This shift goes back to R. A.
Fisher, whose work is based on an explicit analogy between increases in
fitness in large interbreeding populations and increases in entropy: Just as
entropy is a function of the number of available microstates, so increases
in organismic fitness are a function of the genetic variance over a sum of
loci [Fisher, 19301.)
This account is and is not Newtonian in the classical sense. It is
Newtonian insofar as it is an equilibrium theory, in which state changes
are entirely a function of exogenous forces. But it is not Newtonian insofar
as it is not based on a picture in which a dynamic equilibrium is achieved
by the resolution of competing forces. Rather, 'equilibrium' is defined by
zero variance (which for diploid species means the Hardy-Weinberg
binomial distribution of genotypes), just as equilibrium is reached in a
physical system when the probabilities of its components being in partic-
ular microstates are stable. Accordingly, the analogue of inertia is not the
Malthusian population parameter, but rather a statistical distribution of
gene frequencies, deviation from which indicates the presence of a
disturbing force. By defining a biological macrospace in genetical micro-
space terms, this statistical reformulation allows for microscopic rever-
sibility and macroscropic irreversibility, thus giving a better explanation of
our intuitions about the irreversibility of biological phenomena than had
the original Darwinism. And, as Robert Rosen has argued (Rosen, 1985),
it is in one respect more Newtonian than Newton, for it defines in way
Darwin could not the real units or atoms demanded for decomposible
systems, namely populations of independently assorting and segregating
genes.
Now it is quite possible to imagine systems that are in every way the
opposite of Newtonian systems, whether deterministically or statistically
considered. The systems that Prigogine calls dissipative structures embody
this possibility. Such systems are open, non-deterministic, irreversible and
non-decomposible. They are open to extrasystemic inputs of matter and
energy, retaining their identities as distinct systems just because of this
openness. Dissipative structures maintain a trajectory far from equilibrium
as long as appropriate boundary conditions are in place. Indeed, by their
own agency they do much to assure that these conditions obtain by
drawing resources into themselves and dissipating entropic debt beyond
their immediate borders. Dissipative structures are also characterized by
tendencies toward spontaneous self-organization and non-deterministic
bifurcation. This occurs because increases in internal entropy production,

a cost of maintaining internal structure, are more than compensated for by
entropy increases due to energy exchanges between the system and the
external environment. Thus internal organization of the system and the
entropy production of the system-plus-environment grow together. Ac-
cordingly, state changes, or phase transitions, in dissipative structures are
not entirely the result of predictable and measurable impressed forces
operating against an inertial default drive. Lacking inertial tendencies that
counterfactually would reassert themselves when impressed forces are
removed, such systems embody a much stronger arrow of time than we
have seen in either of the two kinds of equilibrial systems we have recently
been considering. Dissipative systems are irreversible, historical entities in
a strong sense, and particularly so when the internalization of information
in replicating macromolecules provides strong forces for conservation.
Finally, because they are composed of complex modules that arise histori-
cally and irreversibly across the micro-macro divide, such systems are only
partially decomposible. They incorporate their historical trajectories into
their very nature.
By shifting the background assumptions of the Darwinian tradition
from those of Newtonian or equilibrial systems to those of complex
non-equilibrium systems, richer connections between physics and evolu-
tionary biology can be forged, especially by way of the ecological interface
between these fields, which gives us deep physical laws on the one hand
and biological systems on the other that are clearly energy processing, and
therefore entropy producing, entities stabilized far from equilibrium. If,
however, one wishes to call such an accomplishment a reduction of
evolutionary biology to physics, chemistry and information theory, one
will have first to revise the received view of what a theoretical reduction is.
As normally understood, the concepts of reduction and reductionism
arise within (roughly) positivist conceptions about the philosophy of
science. This fact brings with it a bias in favor of studying Newtonian
systems as paradigms of mature science. Historically, it was Newtonian
physics that first provided a model toward which, some millenial day,
other sciences might hope to be "reduced". Formally considered, more-
over, covering laws plus initial conditions deductively imply explanations
and predictions only on assumptions about system closure that formally
mirror the material closure conditions of Newtonian systems. The identity
of prediction and explanation rests on Newtonian assumptions about
reversibility, just as the deductive nature of prediction and explanation
reflects Newtonian determinism. Most important in the present context,
however, is the assumption embodied in the received philosophy of
science that progress in subsuming a class of phenomena under laws of a
higher order of generality will normally be correlated with success in
resolving the behavior of wholes into that of smaller and separate parts -
a phenomenon we see vividly in the case of the statistical theory of gases.

The explanation of phenomenal laws of gases in terms of the probability
distributions of particles is a triumph of reduction in this sense. Reduction,
if this is indeed a paradigm of it, implies that productive research
strategies will usually or presumptively culminate in both theoretical and
ontological reduction.
This program can be vividly witnessed in Dawkins' (1976) genic
reduction, in which it is colorfully claimed that organisms are merely
vehicles for the replication of genes, and that organismic traits are selected
in accord with this measure of fitness. In the case, however, of the theory
envisioned in this paper, the illumination of evolutionary biology by the
laws of thermodynamics actually undergirds a modest holism with respect
to entities - none of which will behave according to the specifications for
Newtonian systems, any more than the theories that explain that behavior
will mirror their formal meta-theoretical counterparts in the philosophy of
science. Subsumption of biological phenomena under physical and infor-
mational laws leads to explanations in which the spontaneous emergence
and behavior of macro-systems are shown to be functions of self-
organizing complexity in entities that are relationally constituted as
wholes.
Rather, therefore, than refurbish the concept of reduction to accomo-
date this unexpected divergence between "theory reduction" and "entity
reduction" - a divergence that was bound to occur as soon as science
began to cross the "complexity barrier" (Rosen, 1985; Dyke, 1988a) - it
appears to us to be preferable to speak of the interanimation between
previously isolated fields rather than the reduction of one theory to
another (Darden and Maull, 1977).
The methodological implications of this situation should be mentioned
here in brief. They are studied in detail in Dyke (1988a). Laws provide
not deductive premises, but constraints on the space within which explana-
tory scenarios are possible, as well as expectations of systems behavior
based on very general considerations. Initial and local boundary conditions
are then elevated to central importance in a search for "robust theorems"
(Levins, 1966) that arise at the intersections of different models as they
are brought to bear on particular problems. Laws can be cited as causal
only when we ask such general questions as why a given system is a system
of some type rather than another.
This perspective has implications for other highly charged issues in the
philosophy of biology. One such issue is that of biological teleology.
Traditionally, talk about biological teleology has referred, sometimes
ambiguously, to two different things: (1) the functional adaptedness of
organisms to environments and of organs to organisms; and (2) the overall
direction, and hence alleged directedness, of the evolutionary process
toward greater complexity (and/or other directional properties). It can
hardly be denied that the Darwinian tradition has been hostile to (2). It is
not nearly so hostile, however, to (1). In recent decades, in fact, philoso-
phers, notably Wright (1973), have analyzed teleological statements
roughly as claims to the effect that
X is there because it does Y.
Such claims answer "what for" questions and yield "in order to" answers.
Their general form is common to both design arguments and to natural
selection arguments. If the former make reference to the intentions of an
artificer as the reason an entity has some feature, the latter make reference
to the accumulated effects of natural selection as the reason that some trait
"is there". There is no reason, therefore, to deny that the latter sorts of
arguments are teleological in this sense. Brandon has given the best
analysis of this form. According to him (Brandon, 1981), natural selection
explanations cite "the effects of past instances of A (or precursers of A)
and show how these effects increased the adaptedness of A's possessors
(or the possessors of A's precursors) and so led to the evolution of A"
(Brandon: 1981: 103).
The problem is that genuine adaptionist accounts can easily be replaced
by disguised design arguments. Gould and Lewontin have been highly
effective, for example, in pointing out how easily Wilsonian sociobiology
slides down this slippery slope into disguised design by retailing '"just-so
stories" (Gould and Lewontin, 1979). This slide from selectionist rea-
soning to intentionalist "crypto-teleology" occurs in proportion as the
intentional form is treated as paradigmatic, and the natural-selectionist
form is regarded as an analogue of it - as Darwin himself clearly treated
it. It is for this reason that, the clever work of analytic philosophers aside,
teleology is often regarded by biologists as equivalent to the intentional
form, and is accordingly either denounced or confessed to in accord with
one's proclivities. In the latter case it generates creationism.
One way to break this link, and to drive the wedge between natural
selection and design forms of teleological argumentation that Wright's and
Brandon's work invites, is to anchor natural selection in natural laws and
processes, and so to distance the sort of functional arguments that refer to
natural selection from arguments referring to intentional design. However,
because many evolutionary theorists have sought to protect the autonomy
of biology from physics (Mayr, 1982, 1985, 1988; but see Rosenberg,
1985), and hence are loathe to countenance physical laws as proper
premisses or principles of their science, it is precisely this sort of assur-
ance that has been lacking from discussions of teleology in evolutionary
theory. It is this failure, we believe, that allows creationist arguments, as
well as inadequately formulated sociobiological arguments, to retain
undeserved places as conceptual possibilities in cultural space.
In discussions of this subject it is customary to distinguish among teleo-
matic (or end-resulting) processes, teleonomic (or end-directed) processes

and teleological (or goal seeking) processes (O'Grady, 1984; O'Grady and
Brooks, 1988; Mayr, 1988). Our suggestion is that selectionist arguments,
whose subjects are patterns of informed energy flow (rather than traits
or parts that are too closely modelled on artifacts), are grounded in
teleomatic laws of nature, especially the Second Law (see also Wicken,
1984). Teleonomic feedback within and between ecological systems, as
well as between energy flows and information expansion, arises within this
framework. Thus teleonomic processes, occurring within the possibility
space of teleomatic requirements for dissipation, provide cycles of cause
and effect that fully meet the criteria envisioned by Wright and Brandon
for biological teleology. For the objects of selection that result from this
reiterated process are there, as Wright and Brandon require, just because
they provide dissipative pathways in an informed system of energy flow.
If natural selectionist arguments were put into this form, and referred to
these processes, there would be less occasion for confusing them with
disguised design arguments, and hence for refusing to talk about teleology,
at least in the sense of the term defended by Wright and Brandon. An
advantage of this analysis is that it firmly grounds functional explanations
in natural law without undermining genuine functionality. Thus we can
have a legitimate place in science for functional explanations comple-
mentary to mechanistic explanations. As science shifts its focus of atten-
tion from simple to complex systems, the role of functional explanations
will, in fact, increase relative to mechanistic explanations, since mech-
anistic explanations are rooted in simple systems. Explanations based on
the concept of patterns of informed energy flow in open systems far from
equilibrium are of the teleonomic, and teleological, form because conse-
quences (entropy production) are invoked as reasons for the existence of a
real unit of selection.
Note, however, that the premisses that generate these arguments also
seem to countenance talk about the overall direction of evolution -
teleology in sense (2). If kinetic pathways are naturally selected means
whereby a necessary and inevitable entropic dept is dissipated, are not
these pathways there for the sake of facilitating dissipation; and must we
not say, therefore, that the function of the kinetic pathways is to dissipate
entropy? But since the laws underlying this process are rooted in very
deep, irreversible physical processes, acceptance of this point leads us to
expect that there will be an overall pattern to evolution as dissipative
pathways complexify in real ecological space. It seems, then, that we
cannot talk about local functionality and adaptedness without countenanc-
ing larger patterns of evolutionary complexification and directionality.
From this perspective, when teleology in sense (1) is conceived in the way
just suggested, it seems to countenance certain observations that have
traditionally led to teleology in sense (2). To the extent that the Darwinian
tradition has, in its resolute rejection of cosmic design, often run rough-
shod over what is physically and biologically sound in those observations,
we should welcome an invitation to renew the Darwinian tradition in a
direction that promises to explain macroevolutionary phenomena in terms
of purely natural processes without inviting regression to cosmic design.
In sum: The shift to nonequilibrium thermodynamic background as-
sumptions as a framework for the further development of the Darwinian
tradition leads to the expectation of evolution, and so relieves Darwinian
explanations of temptations to disguised teleology of the 'design' sort. No
longer need Darwinians argue that nature has been able to achieve the
same kind of effect that an engineer-god could attain by 'using' different
means. It is telling that Paley's watchmaker does not disappear in Dawkins'
recent account of evolutionary theory (Dawkins, 1986). He merely is said
to be 'blind.' From our perspective, however, there is no watchmaker,
blind or sighted - for there is no watch. Natural organization is not an
artifact, or anything like it, but instead a teleonomic, and in a certain sense
teleological consequence of the teleomatic action of energy flows. 2
NOTES
i This paper is the result of discussions among the authors and others, which began at the
third meeting on the History, Philosophy, and Social Science of Biology held at the
Virginia Polytechnic Institute and State University in Blacksburg, Virginia, June 1987.
2 Some of us have concerns about the ideological and/or theological extrapolations that
the reader might be inclined to draw from the issues discussed in this section.'
APPENDIX I
Estimating Network Ascendency
Robert E. Ulanowicz
The concept of network ascendency combines the attribute of system size, or level of
activity with the relational notion of dynamic organization.
To quantify the level of system activity input-output theory (Leontieff, 1951) employs
the total system throughput, or aggregate of all the transfers occurring among the system
elements. For example, if i and j are two arbitrary members of a collection of n nodes in a
directed graph, then the transfer of some particular medium between j and i will be
denoted as Tji. The activity level of a specific node i can be gauged either by the sum of
inputs to the node,
T = Ti
or by the collection of all flows issuing from i,
Ti = E Tiq.
(Ti = T' at steady state.) The activity level of the whole system or total system throughput,
T, is equal to the aggregate of the activities at all nodes,
T = Ti= T.
Measuring the level of network organization is only slightly more complicated. We

begin by defining an organized network as one about which there exists little ambiguity as
to where medium at any node will next flow. Ambiguity is associated with uncertainty, and
conversely a decrease in uncertainty bespeaks of information. Hence, it is by employing
elements from probability and information theories that network organization can be
estimated.
Let aj denote the event, "A quantum of medium leaves node j," and b the similar
happening, "A quantum of medium enters node i." The probabilities of each of these
events actually transpiring are denoted by p(aj) and p(bi), respectively. Furthermore, the
joint probability that the quantum both leaves j and enters i will be represented by p(aj, bi).
The average apriori uncertainty about the sources and sinks of flows is given by the
Shannon-Wiener index of uncertainty calculated using the joint probabilities, i.e.,
H = -K L p(aj, bi) log p(aj, bi),
where K is an arbitrary scalar constant.

Knowledge about where (on the average) quanta originate and where they end up [the
p(aj) and p(bi), respectively] decreases this ambiguity, H, by an amount known as the
"average mutual information", denoted here by I and calculated as
I= K ~ E p(aj, bi) log [p(aj,bi)/p(aj)p(bi)].
One can show (Aczel and Daroczy, 1975) that for any properly defined set of probability
distributions
H I 0.
Systems that are highly organized, i.e., those wherein knowing the location of a quantum
also imparts a reasonable knowledge of whither it will go possess a value of the mutual
information, I, that is close to its upper limit, H. That is, most of the ambiguity about which
pathway the medium will take has been resolved. Conversely, in systems that are less
structured, knowing the location of a given quantum tells almost nothing about where that
material is most likely to move and I is insignificantly small. Hence, the mutual informa-
tion, I, is an appropriate index for network organization.
It remains to estimate I in terms of measured system flows. Toward this end it is helpful
first to ask how one would estimate the constitutive probabilities. If T represents that total
amount flowing in the system and T'i is the amount of that flow observed to be entering
node i, then p(bi) can be estimated by the ratio Tf/T. Similarly, the estimator for p(aj) will
become Tj/T. In like manner, the joint probability p(aj, bi) that a quantum will both leave j
and enter i can be assigned the value Tij/T. Substituting these estimates into the formula
for I yields
I = K E ET/T)log
log (TT/T/Ti).
(T/T)
i j
There remains the problem of what value to assign to the undefined scalar constant K.
Conventional practice is to choose a base for the logarithm and arbitrarily set K = 1. But
Trius and McIrvine (1971) argue that K should be chosen so as to impart physical dimen-
sions to the quantity it is scaling. As we have already elected to gauge system "size" using
the total system throughput, T, it is both appropriate and consistent to set K = T, thereby
making the mutual information reflect the size of the system we are observing.
The resulting product of two factors, one representing system size and the other ne-
twork organization, is called the network ascendency (Ulanowicz, 1980) and is denoted by
the letter "A". Knowing the values of all the Tpjin the system allows one to assign an actual
number to the ascendency according to the formula
A= Tji log (Tji T/T; Ti).

i j
Ulanowicz (1986) argues how any increase in A reveals the unitary process of growth and
development transpiring in the system, and he shows how one may investigate the limits to
growth using certain components of the difference by which A falls short of its upper limit
as set by the scaled version of H.
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EVOLUTION IN THERMODYNAMIC PERSPECTIVE:

Biology and Philosophy 4: 373-405, 1989.

Evolution in Thermodynamic Perspective: An

KEY WORDS: Nonequilibrium thermodynamics, information, informed patterns of energy

It is universally conceded that biological systems conform to the con-

In this connection, three highly general propositions have assumed

One measure of progress in the development of any science is its ability

No science has a permanent, natural, defensible frontier, any more than

selves regarded as assemblages of decomposible and ontologically distinct

ECOSYSTEMS PHENOMENOLOGY, ASCENDENCY AND DEVELOPMENT

The focus of ecology is relational. Ecologists concern themselves primarily

if more than one concatenation of transfers linked two arbitrary species in

P10 is a statement of Lotka's power principle (Lotka, 1922, 1925). P11

and P12 together might be thought of as the allocation of a given energy

organism gradually becomes more energetic, while its specific metabolic

of autocatalytic cycling systems. It also implies that organisms continue to

AN ECOLOGICAL APPROACH TO EVOLUTIONARY DYNAMICS

These two principles make opposing contributions to the organization-

dissipative setting. Consider the following dissipative route (Figure 1):

Fig. 1. A hypothetical autocatalytic cycle involving phosphorylation, polymerization, and

Fig. 2. A minimum 'realistic' autocatalytic cycle in which template T and polypeptide PP

As successful patterns of dissipation emerge, so too does a tightening of

in fact selected because it allows organisms to function as relational parts

ENTROPY, INFORMATION, AND GENEALOGY

diversity, given the background requirements of the Second Law. But it is

"noise" on the overloaded channels is reduced. These concepts are made

relationship between thermodynamics and what Shannon called infor-

to transmit and process information in a particularway, availing itself of

boundary and closure conditions can be imposed on biological systems

REDUCTIONISM, TELEOLOGY AND THE NEWTONIAN PARADIGM

The ecological-energetic perspective presented in this paper informs the

system will tend to restore a dynamic equilbrium between inertial motion

path by the imposition of external forces, such as selection. In this recon-

bifurcation. This occurs because increases in internal entropy production,

a phenomenon we see vividly in the case of the statistical theory of gases.

matic (or end-resulting) processes, teleonomic (or end-directed) processes

or by the collection of all flows issuing from i,

Measuring the level of network organization is only slightly more complicated. We

H = -K L p(aj, bi) log p(aj, bi),

where K is an arbitrary scalar constant.

I= K ~ E p(aj, bi) log [p(aj,bi)/p(aj)p(bi)].

A= Tji log (Tji T/T; Ti).

Aczel, J. and Z. Daroczy: 1975, On Measures of Information and Their Characterization,

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