The handicap principle is a hypothesis proposed by the biologist Amotz Zahavi to explain how evolution may lead to "honest" or reliable signalling between animals which have an obvious motivation to bluff or deceive each other. It suggests that costly signals must be reliable signals, costing the signaller something that could not be afforded by an individual with less of a particular trait. For example, in sexual selection, the theory suggests that animals of greater biological fitness signal this status through handicapping behaviour, or morphology that effectively lowers this quality. The central idea is that sexually selected traits function like conspicuous consumption, signalling the ability to afford to squander a resource. Receivers then know that the signal indicates quality, because inferior-quality signallers are unable to produce such wastefully extravagant signals.
The handicap principle was proposed in 1975 by Israeli biologist Amotz Zahavi.
The generality of the phenomenon is the matter of some debate and disagreement, and Zahavi's views on the scope and importance of handicaps in biology have not been accepted by the mainstream. Nevertheless, the idea has been very influential, with most researchers in the field believing that the theory explains some aspects of animal communication.
Though the handicap principle was initially controversial,—John Maynard Smith was a notable early critic of Zahavi's ideas—it has gained wider acceptance because it is supported by game theory models, most notably Alan Grafen's signalling game model. This is essentially a rediscovery of Michael Spence's job market signalling model, where the job applicant signals their quality by declaring a costly education. In Grafen's model, the courting male's quality is signalled by investment in an extravagant trait—such as the peacock's tail. In both cases, the signal is reliable if the cost to the signaller of producing it is proportionately lower for higher-quality signallers than it is for lower-quality signallers (Fig. 2).
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Within evolutionary biology, signalling theory is a body of theoretical work examining communication between individuals, both within species and across species. The central question is when organisms with conflicting interests, such as in sexual selection, should be expected to provide honest signals (no presumption being made of conscious intention) rather than cheating. Mathematical models describe how signalling can contribute to an evolutionarily stable strategy.
Evolutionary game theory (EGT) is the application of game theory to evolving populations in biology. It defines a framework of contests, strategies, and analytics into which Darwinian competition can be modelled. It originated in 1973 with John Maynard Smith and George R. Price's formalisation of contests, analysed as strategies, and the mathematical criteria that can be used to predict the results of competing strategies. Evolutionary game theory differs from classical game theory in focusing more on the dynamics of strategy change.
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