Abstract

The general attributes of ecosystems are examined and a naturally occurring reference ecosystem is established, comparable with the isolated system of classical thermodynamics. Such an autonomous system with a stable, periodic input of energy is shown to assume certain structural characteristics that have an identifiable thermodynamic basis. Individual species tend to assume a state of least dissipation; this is most clearly evident in the dominant species (the species with the best integration of energy acquisition and conservation). It is concluded that ecosystem structure results from the antagonistic interaction of two nearly equal forces. These forces have their origin in the Principle of Most Action (least dissipation or least entropy production) and the universal Principle of Least Action. Most action is contingent on the equipartitioning of the energy available, through uniform interaction of similar individuals. The trend to Least action is contingent on increased dissipation attained through increasing diversity and increasing complexity. These principles exhibit a basic asymmetry. Given the operation of these opposing principles over evolutionary time, it is argued that ecosystems originated in the vicinity of thermodynamic equilibrium through the resonant amplification of reversible fluctuations. On account of the basic asymmetry the system was able to evolve away from thermodynamic equilibrium provided that it remained within the vicinity of ergodynamic equilibrium (equilibrium maintained by internal work, where the opposing forces are equal and opposite).At the highest level of generalization there appear to be three principles operating: i) maximum association of free-energy and materials; ii) energy conservation (deceleration of the energy flow) through symmetric interaction and increased homogeneity; and iii) the principle of least action which induces acceleration of the energy flow through asymmetrical interaction. The opposition and asymmetry of the two forces give rise to natural selection and evolution.