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Evolutionary Game Theory, Natural Selection, and Darwinian Dynamics: Thomas L. Vincent, Joel S. Brown

Book Description:

All of life is a game and evolution by natural selection is no exception. The evolutionary game theory developed in this book provides the tools necessary for understanding many of nature's mysteries, including co-evolution, speciation, extinction and the major biological questions regarding fit of form and function, diversity, procession, and the distribution and abundance of life. Mathematics for the evolutionary game are developed based on Darwin's postulates leading to the concept of a fitness generating function (G-function). G-function is a tool that simplifies notation and plays an important role developing Darwinian dynamics that drive natural selection. Natural selection may result in special outcomes such as the evolutionarily stable strategy (ESS). An ESS maximum principle is formulated and its graphical representation as an adaptive landscape illuminates concepts such as adaptation, Fisher's Fundamental Theorem of Natural Selection, and the nature of life's evolutionary game.

Review:

First, full disclosure: I am a colleague and friend of the authors, Thomas L. Vincent and Joel S. Brown, and I reviewed the entire book during its writing.

Game theory is a fairly recent development in mathematics, having been introduced in the 1940's. Evolutionary Game Theory is more recent yet - Maynard Smith and Price put it on the map with their publication in Nature in 1973 on the Logic of Animal Conflict. Maynard Smith then more fully elaborated the application of matrix games to evolution with his 1982 volume, Evolution and the Theory of Games. Vincent and Brown trace their contribution to the pioneering developments of Maynard Smith, but in this volume, they go much further. As I reviewed the eleven chapters as they were first written, I felt the privilege of observing, first hand, the construction of a great edifice. In this edifice, the dynamics of ecology is dovetailed with the dynamics of heritable strategies. The tool that accomplishes this is the fitness generating function, known as the G-function. Particularly brilliant is the invention of the virtual strategy, a scalar or vector "place holder" in the G-function. The great virtue of the virtual strategy is that it represents any focal individual taking on any strategy within the entire strategy set of the species. The fitness generating function then determines the fitness for that virtual strategy within the biotic and abiotic environment defined by the set of arguments (e.g., resident strategies, their population sizes, abundance of resources, etc.) defining the G-function. With G-function in hand, Evolutionary Game Theorists now have a mathematical Darwinism - a formal mathematical expression of Darwin's three postulates: a) like begets like; b) organisms struggle for existence; c) heritable traits help determine the outcome of the struggle. With the G-function, we can predict both the dynamics of heritable strategies and the adaptive outcome of natural selection.

Vincent and Brown begin, in Chapter 1, with an historical and philosophical overview of Evolutionary Game Theory and its relationship to the more traditional approach of Evolutionary Genetics. They then proceed to lay the mathematical foundations (Chapters 2 - 7), constructing the theory of Evolutionary Games and the G-function. These chapters each contain useful examples, teaching the student of evolutionary games how to apply the G-function. Noteworthy is that most all of the examples in these chapters represent continuous, as opposed to matrix games. In matrix games, which constitute the bulk of early development of Evolutionary Game Theory, and with which most readers are probably most familiar, strategies are discrete rather than continuous. However, the continuous games elaborated by Vincent and Brown (and now, many others) are of far more useful application in Evolutionary Ecology. Key contributions here are the precise mathematical definition of Maynard Smith's seminal Evolutionarily Stable Strategy (ESS) in Chapter 6, and the formulation of the ESS Maximum Principle in Chapter 7. This principle establishes the well-recognized properties of the ESS of invasion resistance and convergent stability, but also the fit of form and function - the ESS strategy is an adaptation - it maximizes individual fitness given the circumstances.

Chapter 8, which treats species concepts, speciation, and extinction, is particularly enlightening. Here the G-function shines! Under traditional approaches, a huge chasm, conceptual and methodological, separates microevolution and macroevolution. Vincent and Brown, armed with the G-function, unify the two: Microevolution is repeatable and reversible evolutionary dynamics within a G-function. Macroevolution is the production of novel G-functions. They demonstrate the versatility of the G-function approach to Evolutionary Game Theory in their discussion of three contexts for extinction (which is as integral to evolution as is speciation). Vincent and Brown introduce many key concepts in Chapter 8. Perhaps most important is their strategy species concept, which relies on their definition of the species archetype. They provide a particularly cogent definition of a species that is ecologically keystone (its presence promotes the persistence, in ecological time, of other species in the community), but they also point out that a species can by evolutionarily keystone - when its presence increases the numbers of species at an ESS. Using these developments, Vincent and Brown investigate mechanisms of speciation, including sympatric speciation, allopatric speciation, adaptive radiations, coevolution, Wright's shifting balance theory, and incumbent replacement. They conclude with a tour de force: a concise and brilliant discussion of the Procession of Life. As they aptly demonstrate, with the G-function approach to the Game of Life, theories such as Punctuated Equilibrium, oft cited as a contradiction of Darwinian Evolution, instead result naturally from Darwin's three postulates!

Chapter 9 is perhaps the least exciting chapter, but it serves the utilitarian purpose of melding the matrix approach to Evolutionary Game Theory with the G-function approach. This is, indeed, required reading for those who think matrix games are the only game in town.

Chapters 10 and 11 are well worth the wait and development. In these chapters, Vincent and Brown apply the G-function to an impressive diversity of problems arising in the beautiful metaphor of Hutchinson, the Ecological Theater and Evolutionary Play. Though the diversity of topics covered in these two chapters is impressive, as Vincent and Brown state, it represents only a subset of the problems that can be investigated with G-functions. Chapter 10 addresses "basic" issues of Evolutionary Ecology - a who's who of fundamental subjects. These include: Habitat selection and the ideal free distribution; Consumer-resource games, with examples on plant competition and root-shoot ratio; Carcinogenesis (a must read for all interested in Darwinian Medicine); Flowering time for annual plants; Root competition; and Foraging games.

Chapter 11 turns to the G-function as a fundamental tool for Applied Evolutionary Ecology. Here Vincent and Brown examine: Evolutionary responses to harvesting; Resource management and conservation; and Chemotherapy-driven evolution. They contrast management based on ecological enlightenment with that based on evolutionary enlightenment (prescriptions based on each emphasis are not always identical!). They point out the resemblance of control of a cancer with chemotherapy with control of a population through hunting. The analysis is striking, with the main message that if all cancer cells are not destroyed by a chemotherapy session, the survivors will evolve as the first step of what they call chemotherapy-driven evolution. If ever Evolutionary Ecologists were looking for a raison d'etre, here they have it!

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