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Department of Organismic and Evolutionary Biology, Harvard
University, Cambridge, MA 02138
Communicated by Dudley R. Herschbach, Harvard University,
Cambridge, MA, February 7, 1997
(received for review October 3, 1996)
Directionality in populations of replicating organisms can be
parametrized in terms of a statistical concept: evolutionary entropy.
This parameter, a measure of the variability in the age of reproducing
individuals in a population, is isometric with the macroscopic variable
body size. Evolutionary trends in entropy due to mutation and natural
selection fall into patterns modulated by ecological and demographic
constraints, which are delineated as follows: (i)
density-dependent conditions (a unidirectional increase in evolutionary
entropy), and (ii) density-independent conditions,
(a) slow exponential growth (an increase in entropy); (b) rapid exponential growth, low degree of iteroparity
(a decrease in entropy); and (c) rapid exponential
growth, high degree of iteroparity (random, nondirectional change in
entropy). Directionality in aggregates of inanimate matter can be
parametrized in terms of the statistical concept, thermodynamic
entropy, a measure of disorder. Directional trends in entropy in
aggregates of matter fall into patterns determined by the nature of the
adiabatic constraints, which are characterized as follows:
(i) irreversible processes (an increase in thermodynamic
entropy) and (ii) reversible processes (a constant value
for entropy). This article analyzes the relation between the concepts
that underlie the directionality principles in evolutionary biology and
physical systems. For models of cellular populations, an analytic
relation is derived between generation time, the average length of the
cell cycle, and temperature. This correspondence between generation
time, an evolutionary parameter, and temperature, a thermodynamic
variable, is exploited to show that the increase in evolutionary
entropy that characterizes population processes under density-dependent
conditions represents a nonequilibrium analogue of the second law of
thermodynamics.
Proc. Natl. Acad. Sci. USA
Vol. 94,
pp. 3491-3498,
April 1997
Review
Directionality principles in thermodynamics and evolution
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