John Maynard Smith

(1920 - )

Compiled by Carrie Cook (December 1999)

J.M. Smith Biography
Time Line

John Maynard Smith was born in 1920 in London. He received a strong education at Eton and Cambridge, from which he graduated with a degree in aeronautical engineering in 1941. He then attended University College in London, where he graduated in 1951 with a degree in zoology. Upon receiving his degree, Maynard Smith was offered a teaching position at his alma mater, where he taught from 1951 until 1965. After leaving University College, he was appointed professor of biology at the University of Sussex in 1965, and he is presently teaching there today. He was recently awarded The Royal Swedish Academy of Sciences 1999 Crafoord Prize for biology in conjunction with Ernst Mayr and George C. Williams of the United States for their work toward a greater understanding of biological evolution and the adaptation of species to their particular environment (URL, 1).

Maynard Smith’s main area of expertise is in genetics, mathematics, and evolutionary biology. Throughout his career, he has concentrated on the fighting strategies among animals, especially within the same species. He feels it is important to understand why animals behave the way they do in nature and the physical encounters that they face against members of their own species. Maynard Smith has conducted research studies of animal fighting strategies and the signals and behaviors that they exhibit before and during battle. Maynard Smith has written many books and journal articles addressing evolution, biology, and game theory, to name a few. Some of these works include The Theory of Evolution (1958), Mathematical Ideas in Biology (1968), Models in Ecology (1974), Evolution of Sex (1978), and Theory of Games (1982). His most recent publications, which were published in 1997, include Role of Mutator Genes in Adaptive Evolution and Evolution of Genetic Redundancy.

Maynard Smith is best known for three different theories. His research on game theory, evolutionarily stable strategy, and hawk and dove strategies has made a big impact on the world of science. These three theories are discussed below.


Game Theory: Game theory refers to “a strategy that considers competitors’ behaviors” (Maier 1998, 501). Game theory was originally viewed as a mathematical theory or method that may be used to analyze how an actor- player- should behave when he cannot have complete information about what another actor- “opponent”- is going to do” in certain situations (URL, 2). Animals, like humans, will hesitate and wait to see what others do before making a decision. Maynard Smith has observed that it is rare for rivals to kill one another in nature. Rivals use threats to symbolize power but have no intention of engaging in conflict (URL, 2). An example of game theory can be applied to two male animals that are competing for the attention of a female with which they wish to mate. Both males entering the battle do not know what the final outcome will bring. They could be physically or fatally wounded, but that is the price that they pay in order to pass on their genes. The decision must be made by both animals as to whether it is worth such a great risk just to be able to copulate with the female. It is a common issue in the animal world, but the idea of competing and not knowing the outcome is an example of game theory. It is uncommon for members of the same species to kill one another in battle because animals cannot grasp a complete understanding of their opponents’ information or capabilities in combat, so it is difficult and pointless to fight one another. It does not accomplish anything and can only lead to problems in nature. Maynard Smith has applied graded signals, which are communication signals that vary in intensity and duration in terms of the sender’s motivation level, to game theory (Maier, 1998).


An example of graded signals in nature is the intensity of a deer roar in battle. Two male deer may be competing for a female. Instead of attacking one another, they compete to see who can roar the loudest and longest, based on who wants to mate more. This idea is an elaboration of the idea of game theory in nature. Game theory and fighting strategies among animals can be related to the evolutionarily stable strategy theory (Maier 1998).

Evolutionarily Stable Strategy: Evolutionarily stable strategy, or ESS, is a strategy that is so effective that it will not likely be replaced by an alternative strategy” (Maier, 1998). Maynard Smith introduced the idea of evolutionarily stable strategy in terms of game theory in his essay Game Theory and The Evolution of Fighting in 1972. He found this concept of ESS to be vital in the new traits among populations. In ESS, different characteristics are balanced within a particular species. He determined that the strategy behind a particular population is to resist the formation of new traits or characteristics, such as the length of a horn or whether to fight or retreat when faced with an opponent, since it may lessen the possibilities for successful reproduction. Members within the same species are in constant competition for the same resources, such as food and the proper mates (URL, 3). If an aggressor kills a member of its own species, one would believe that it has done well by limiting the competition and protecting the remaining members. In actuality, by being the aggressor and always engaging in conflict, an animal may be risking its own life for the sake of power. There needs to be an equilibrium between two species and within species in order to keep nature stable and working as it should. Evolutionarily stable strategy can be applied to the tit-for-tat strategy because it cannot be replaced by other strategies. In regard to both of these strategies, if two parties decide to cooperate in a competition, the cost could be deadly, whereas if one fails to cooperate, there would be a minimal loss. It is better for animals to decide to cooperate rather than compete. The basic theory behind evolutionarily stable strategy is “fight if the opponent is smaller, but retreat if the opponent is larger” (URL, 4). Maynard Smith’s research with evolutionarily stable strategy was so profound and important that it is commonly used in relation to economics and biology today (URL, 5).

Hawk and Dove Strategies: Fighting strategies are vital necessities within nature. The hawk strategy is one that favors fighting an opponent over backing off (Maier,1998). Dove strategies entail withdrawing from an opponent rather than engaging in an attack. If a particular species acted like a dove and was not an aggressor, there would be an advantage to being a hawk. The hawk would dominate over the population, and have a greater chance of locating food, being a predator, as well as possess the capability to reproduce successfully. The downfall to this idea that Maynard Smith found was that, as the population of hawks increases, there would then be a greater likelihood of competition within a population. The hawks may fight more amongst themselves and may risk the possibility of successfully reproducing or death (Maier 1998). With an increased hawk population, the doves may be at an advantage. There could be a positive outcome to a confrontation, where there is less of a chance of injury or death. The idea of hawk and dove strategies is a conditional strategy that serves as an alternative to fighting that animals may develop. According to Maynard Smith, if hawk and dove strategies are genetically influenced, the population will reach a point where there are stable percentages of hawks and doves. This idea of a stable relationship that develops between animals that fight and those that back off is known as stable polymorphism (Maier, 1998). Maynard Smith has found that when there is an equilibrium between the number of hawks and doves within a population, then that species will be able to survive and produce in a fashion that will be beneficial for a given population.

Conclusion: Maynard Smith has studied evolution and has related animal behavior to game theory and fighting strategies. His extensive research and work on game theory, evolutionarily stable strategy, and hawk and dove strategies has provided a greater understanding of animal behavior in relation to their reaction to predators and competition.

Time Line
1920 Born in London
1941 Graduated as an aeronautical engineer from Eton and Cambridge
1951 Degree in zoology from the University College in London
1951- 65 Taught at University College
1958 Published The Theory of Evolution
1965 Appointed professor of biology at the University of Sussex
1968 Punblished Mathematical Ideas in Biology
1972 Evolutionarily Stable Strategy (ESS) introduced in the essay, Game Theory and the Evolution of Fighting
1974 Published Models in Ecology
1978 Published Evolution of Sex
1982 Published Theory of Games
1997 Published Role of Mutator Genes in Adaptive Evolution
1997 Published Evolution of Genetic Redundancy
1999 Awarded the Crafoord Prize from the Royal Swedish Academy of Sciences
1999 Professor emeritus at University of Sussex, UK

Maier, Richard. (1998) Comparative Animal Behavior: An Evolutionary and Ecological Approach. Boston: Allyn and Bacon.

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URL 4. Evolutionarily stable strategy. Helicon Publishing, 1999.

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