Ageing is mostly explained as a by-product of evolution, since selection weakens after reproduction. Biologists still debate whether it is a side effect or an optimised trade-off.
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Evolutionary mismatch is the idea that a trait shaped by one environment can become harmful when the environment changes faster than evolution can respond.
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The Prisoner's Dilemma shows that one-shot interactions favor defection. Cooperation can evolve when interactions repeat, as with reciprocity strategies like tit-for-tat.
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The handicap principle holds that signals are honest because they are costly. It became widely accepted, but a 2019 review argues the idea is erroneous and that honesty can evolve without costs.
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Richard Dawkins's 1982 idea holds that a gene's phenotype includes all its effects on the world, inside or outside the organism's body, such as dams, manipulated hosts and mimicry.
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Both theories explain how altruism evolves and can often be translated into each other mathematically. The dispute is over whether that equivalence means they describe the same causes.
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