Biological signals as handicaps
Grafen built an evolutionarily stable strategy model of Zahavi's handicap principle. He reported that a stable honest signalling equilibrium exists when the conditions of the principle hold, that advertising must lower the signaller's fitness, and that the marginal cost of advertising must be lower for the better signaller. If advertising were free or equally costly for good and poor signallers it could not work as a handicap. The paper also discusses economic signalling models and the limits of the model.
Grafen built an evolutionarily stable strategy model of Zahavi's handicap principle. He reported that a stable honest signalling equilibrium exists when the conditions of the principle hold, that advertising must lower the signaller's fitness, and that the marginal cost of advertising must be lower for the better signaller. If advertising were free or equally costly for good and poor signallers it could not work as a handicap. The paper also discusses economic signalling models and the limits of the model.
Written from the abstract and body of the paper (open copy), 2026-11-01.
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Grafen, A. (1990). “Biological signals as handicaps.” Journal of Theoretical Biology, 144(4), 517-546. doi:10.1016/S0022-5193(05)80088-8 https://doi.org/10.1016/S0022-5193(05)80088-8