Abstract

Darwinian fitness describes the capacity of an organism to appropriate resources from the environment and to convert these resources into net-offspring production. Studies of competition between related types indicate that fitness is analytically described by entropy, a statistical measure which is positively correlated with population stability, and describes the number of accessible pathways of energy flow between the individuals in the population. Directionality theory is a mathematical model of the evolutionary process based on the concept evolutionary entropy as the measure of fitness. The theory predicts that the changes which occur as a population evolves from one non-equilibrium steady state to another are described by the following directionality principle–fundamental theorem of evolution: (a) an increase in evolutionary entropy when resource composition is diverse, and resource abundance constant; (b) a decrease in evolutionary entropy when resource composition is singular, and resource abundance variable. Evolutionary entropy characterizes the dynamics of energy flow between the individual elements in various classes of biological networks: (a) where the units are individuals parameterized by age, and their age-specific fecundity and mortality; where the units are metabolites, and the transitions are the biochemical reactions that convert substrates to products; (c) where the units are social groups, and the forces are the cooperative and competitive interactions between the individual groups. % This article reviews the analytical basis of the evolutionary entropic principle, and describes applications of directionality theory to the study of evolutionary dynamics in two biological systems; (i) social networks–the evolution of cooperation; (ii) metabolic networks–the evolution of body size. Statistical thermodynamics is a mathematical model of macroscopic behavior in inanimate matter based on entropy, a statistical measure which describes the number of ways the molecules that compose the a material aggregate can be arranged to attain the same total energy. This theory predicts an increase in thermodynamic entropy as the system evolves towards its equilibrium state. We will delineate the relation between directionality theory and statistical thermodynamics, and review the claim that the entropic principle for thermodynamic systems is the limit, as the resource production rate tends to zero, and population size tends to infinity, of the entropic principle for evolutionary systems.

Highlights

  • One of the remarkable features of the organic world is the large diversity of plants and animals that exist and the high degree of adaptation of these organisms to their natural environment

  • The question which emerges from this observation is: Can this directional change in the morphology and physiology of organisms be explained in terms of a dynamic model which takes into account the fact that biopopulations are composed of genetically unique individuals?

  • We will appeal to the depiction of thermodynamic and evolutionary entropy given by Equations (91) and (92) to show that the thermodynamic principle is the limit, as the resource production rate tends to zero and the population size tends to infinity, of the evolutionary entropic principle

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Summary

Introduction

One of the remarkable features of the organic world is the large diversity of plants and animals that exist and the high degree of adaptation of these organisms to their natural environment. This new class of models showed that the outcome of competition between related types is determined by the rate at which the population appropriates the exiguous resources This rate can be analytically described by the statistical parameter, evolutionary entropy. Competitive outcome, when limited resource constraint prevails, is decided by the rate at which an individual acquires energy from the environment and converts this energy into demographic currency This rate is precisely the macroscopic parameter, evolutionary entropy. These three modes of selection and the relations between them will be described in Sections 1.1 to 1.4 These sections provide a conceptual overview of the main tenets of directionality theory: We furnish a historical account of earlier efforts, due to Lotka and Fisher, to develop an analytic model of evolutionary dynamics within the framework of the statistical thermodynamics of Boltzmann.

Evolutionary Selection–Malthusian and Entropic
Malthusian Systems
Evolutionary Entropic Systems
Malthusian and Evolutionary Entropic Systems
Thermodynamic Selection
Thermodynamic and Evolutionary Processes
The Selection Principles
Historical Digression
Bioenergetic Considerations—Lotka and the Thermodynamic Paradigm
Demographic Systems—Fisher and the Malthusian Paradigm
Organization of Paper
Entropy
Thermodynamic Entropy
Evolutionary Entropy
Statistical Measures
Thermodynamic Entropy and Evolutionary Entropy
Macroscopic Variables in Biological Networks
The Dynamics of Biological Networks
The Steady State
The Macroscopic Variables
The Entropic Selection Principle
Incumbent and Variant Populations
Invasion-Extinction Dynamics
Evolutionary Stability
Selection Principle and Resource Constraints
Demographic Parameters and Resource Constraints
Complexity and Stability of Biological Networks
Complexity and Evolutionary Entropy
Robustness and the Fluctuation Decay Rate
Entropy and Robustness
The Evolution of Robustness
The Evolution of Cooperation
Evolutionary Entropy and Sociality
Evolutionary Entropy as a Measure of Social Organization
Relatedness and Social Network
A Genetic Theory of the Evolution of Social Behavior
Inclusive Fitness Theory
Inclusive Fitness Theory and the Entropic Theory of Sociality
Directionality Theory
The Dynamics of the Incumbent
The Composite Population
The Fundamental Theorem of Evolution
Empirical Considerations
10. Directionality Theory
10.1. The Dynamics of Speciation
10.1.1. Mean Fixation Time
10.1.2. Entropy and Speciation Rate
10.2. The Dynamics of Extinction
10.3. Background Extinction
10.4. Mass Extinction
11. Directionality Principles in Macroevolution
11.1. Speciation and Background Extinction
11.2. Speciation and Alternation of Background and Mass Extinction
11.3. The Evolution of Mean Body Size
11.4. The Evolution of Maximum Body Size
12. Evolutionary Dynamics and Thermodynamics
12.1. The Entropic Principles: A Characterization
13. The Entropic Principles
14. The Origin of Metabolic-Replicating Systems
14.1. Inanimate Matter and Living Systems
14.2. The Continuation Rule and the Origin of Life
Findings
15. Conclusions
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