The chess-board is the world;
the pieces are the phenomena of the universe;
the rules of the game are what we call the laws of Nature.
The player on the other side is hidden from us.
We know that his play is always fair, just, and patient.
But also we know, to our cost, that he never overlooks a mistake,
or makes the smallest allowance for ignorance.

T. H. HUXLEY 1825–1895,
Lay Sermons: A Liberal Education


The phenomenon of cooperative interactions among animals has puzzled biologists since Darwin. Nevertheless, theoretical concepts to study cooperation appeared only a century later and originated in economics and political sciences rather than biology. John von Neumann and Oskar Morgenstern developed a mathematical framework termed Game Theory to describe interactions between individuals. This theory emerged in the wake of World War II and was mainly intended to provide a basis to prevent a nuclear holocaust. John Nash, working at the post-WWII US military think tank, the RAND Corporation, augmented the theory by developing and introducing the concept of equilibria, the so-called Nash equilibria:

An equilibrium is reached as soon as no party can increase its profit by unilaterally deciding differently.

Another generation later, John Maynard-Smith and George R. Price ingeniously related the economic concept of payoff functions with evolutionary fitness as the only relevant currency in evolution. Furthermore, Maynard-Smith refined the concept of Nash equilibria in an evolutionary context and introduced the notion of evolutionarily stable strategies (ESS). All ESS represent a subset of the Nash equilibria because an ESS applies only at the population level and adds stability requirements.

A strategy is called evolutionary stable if a population of individuals homogenously playing this strategy is able to outperform and eliminate a small amount of any mutant strategy introduced into the population.

These achievements mark the advent of an entirely new, approach to behavioral ecology where theoretical models and predictions inspired and continue to inspire numerous experiments and field studies.

Figure 1: The forefathers of game theory and the theory of the evolution of cooperation.
John von Neumann Oskar Morgenstern John Nash John Maynard-Smith
John von Neumann Oskar Morgenstern John Nash John Maynard-Smith


Spatial Prisoner's Dilemma

Prisoner's Dilemma

The Prisoner's Dilemma is probably the most famous mathematical methaphor for modelling the evolution of cooperation. This section gives a brief introduction into the Prisoner's Dilemma and provides examples of the game dynamics in well-mixed populations with random interactions as well as structured populations with limited local interactions.

Spatial Snowdrift game

Snowdrift game

A closely related game, which is also addressing the problem of cooperation but under slightly relaxed conditions, is called the Snowdrift game. This section briefly introduces the Snowdrift game and then, similarly to the Prisoner's dilemma, exemplifies the game dynamics in well-mixed populations as well as structured populations.

Spatial Rock-Scissors-Paper game

Rock-Scissors-Paper game

The fascination of the Rock-Scissors-Paper game is not restricted to children but is equally thrilling for evolutionary biologists. The cyclic dominance of the three strategies can lead to very interesting dynamics both in well-mixed as well as structured populations.

Selected publications on evolutionary game theory:


The development of these pages would not have been possible without the encouragement and the insightful advice of Karl Sigmund. Financial support for the first version of these pages of the Swiss National Science Foundation is gratefully acknowledged.