evolutionary game theory. game theory von neumann & morgenstern (1953) studying economic...
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Evolutionary Game Theory
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Game Theory
• Von Neumann & Morgenstern (1953)Studying economic behavior
• Maynard Smith & Price (1973)Why are animal conflicts examples of ‘limited wars’?
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Assumptions
• Infinite population size
• Random mating
• Asexual reproduction
• Frequency dependent fitness
• Genotype can be mapped directly onto phenotype - haplotypes
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Fundamental Concept
• The Evolutionary Stable Strategy (ESS)
“A strategy such that if all members of the population adopt it, then no mutant can invade the population under the influence of selection”
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The Haploid Hawk Dove Game
• Consider two haplod virus genotypes that breed true
• The Hawk genotype encodes a virulent virus strain.
• The Dove genotype encodes an avirulent virus strain
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Fitness payoffs
• The reproductive value of an infected host to a virus is V
• When two virulent viruses (H) coinfect a host there is a cost associated with morbidity C
• When a virulent virus (H) coinfects with a avirulent virus (D), H derives all the benefits V.
• When two avirulent viruses (D) infect a host they obtain approximately half of the resource each V/2
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Payoff matrix
H D
H 1/2(V-C) V
D 0 V/2
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Building the model
• p = frequency of H viruses
• W(H) & W(D), denote mean fitness
• E(H,D) fitness payoff to H infecting a body already infected with D, similar meaning for E(H,H), E(D,H) and E(D,D)
• W0 is the fitness of the virus prior to infection of the host
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Virus fitnesses
• Upon infection of a single host:
W(H) = W0+ pE(H,H) + (1-p)E(H,D)
W(D) = W0 + pE(D,H) + (1-p)E(D,D)
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Determining the ESS conditions
• Consider any two genotypes I & J:W(I) = W0 + pE(I,J) + (1-p)E(I,I)W(J) = W0 + pE(J,J) + (1-p)E(J,I)
• Assume that I is an ESS and J is a rare mutant with frequency p
• If I is an ESS then W(I) > W(J), assuming that p <<1, then,E(I,I) > E(J,I) or (Invasion condition)E(I,I) = E(J,I) and E(I,J) > E(J,J) (Stability)
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ESS solutions to the H & D game
• E(I,I) > E(J,I) or (Invasion condition)E(I,I) = E(J,I) and E(I,J) > E(J,J) (Stability)
• E(D,D) > E(H,D) Never!
• E(H,H) > E(D,H) only of 1/2(V-C) > 0
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Mixed ESS solutions
• What if V<C?• Does this mean that there is no ESS solution to the
game?• An alternative ESS solution can exist if the biology
permits.• This requires either a genotype capable of switching
between H and D or some mix of H & D coexisting in the population.
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Mixed ESS solution
• Consider strategy I as genotype H with probability P and genotype D with probability (1-P).
• For a mixed ESS to exist then:E(A,I) = E(B,I) = E(C,I)…= E(I,I)All pure strategies in support of I must have the same payoff.
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Finding the mixed ESS
• If I is a mixed ESS then E(H,I)= E(D,I):
• E(H,I) = PE(H,H) + (1-P)E(H,D)
• E(D,I) = PE(D,H) + (1-P)E(D,D)
• P(1/2)(V-C) + (1-P)V = P.0 + (1-P)V/2
• Solve for P
• P = V/C
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Testing I with the ESS conditions
• E(I,I) > E(J,I) or (Invasion condition)E(I,I) = E(J,I) and E(I,J) > E(J,J) (Stability)
• We need to see if I meets the stability condition:E(H,I) = E(D,I) = E(I,I) (True)
• Therefore we require that:E(I,D) > E(D,D) & E(I,H) > E(H,H)
• Calculate the above and show that I is an ESS
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Evolution of virulence genes
• When V > C then virulent virus always favoured
• When V < C then some proportion of the population given by V/C will be virulent
• Increasing the cost favours avirulent forms• Reducing the cost favours the virulent
forms
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Game Theory Summary
• Fitness of a gene can depend on frequencies of all other genes in a population -- fitness is frequency dependent
• Game theory provides a tool for determining the equilibrium distribution of genotypes in the population when fitness is frequency dependent
• Key Reference: John Maynard Smith.Evolution and the Theory of Games. CUP. 1982.
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Game theory: anisogamy
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Game Theory: the sex ratio
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Game theory: area of application
• Frequency-dependent selection
• Ignorant about genetic mechanisms
• Parthenogenetic inheritance
• Act as an aid to intuition before building more complex models
• When we do know about genetics it is best to add selection to our population genetics models