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Sex Ratios and the Power of Game Theory

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The sex ratio is the ratio of males to females in a population

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Sex Ratios Are Approximately 50:50 In Most Species

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Including Humans

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Puzzle: Why are sex ratios at birth approximately 50:50 in most species?

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Is it to ensure that everyone has a mate?

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From: Trivers 1976

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Is it to maximize the size of the species?

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No.

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The real answer, thanks to Fisher (1930)

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Simplest case: Female and male babies require approximately the same parental investment No inbreeding

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FACT: Every child must have exactly one mother and one father

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Suppose there are: 100 parents: 75 females, and 25 males 100 offspring Each male expects 100/25 = 4 offspring Each female expects 100/75 = 1.33 offspring Males have more offspring

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Parents of males have more grandkids

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If everyone else gives birth to offspring at a ratio of 1 male to every 3 females (25:75) You’d do better by having more male offspring 25:75 is not a Nash Equilibrium

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Same argument holds for any sex ratio

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Except 50:50 Then, male and female offspring have the same expected number of offspring And you can’t do better by having more male or female offspring

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50:50 is the unique Nash Equilibrium

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Are we sure evolution will lead to 50:50?

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If everyone gives birth to offspring at a ratio of 3 females for every male (25:75) A mutant gene arises that leads to more male offspring The individual with the mutant gene will have more grandkids, and the gene will end up disproportionally represented in two generations Since individuals with this gene have more male offspring, this would increase the sex ratio

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Until… the population hits 50:50 At 50:50, the mutant’s male offspring won’t give more grandkids And the mutant will stop spreading

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At 50:50, no mutant does better than existing population And if it does by chance, the mutant will do worse and we’ll return to 50:50

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Evolution leads to Nash Equilibrium

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Robust Doesn’t matter if: Males die before maturity, since result is driven by the expected number of children Polygyny, for same reason Population size isn’t constant Sex determined by one sex

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Wait a minute. Sex ratios aren’t always 50:50!

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Sex Ratios vs. Parental Investment Source: Trivers 1976

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Sex Ratios Amongst Inbred Wasps Source: Herre 1987

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These exceptions are actually predicted by the theory

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What if parents invest more in one sex?

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Suppose females twice as costly to make as males

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Is 50:50 an equilibrium?

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Male and female offspring yield the same number of grandkids But male offspring cost ½ as much

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If everyone else gives birth to offspring at a ratio of one male to every female (50:50) Can do better by having primarily male offspring, and thus more offspring 50:50 is not a Nash Equilibrium

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In equilibrium, would have more males

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Novel prediction that fits the data!

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What if there is inbreeding?

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Male offspring mate only with their sisters

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Maximize number of grandkids by having mostly females

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Novel prediction that fits the data!

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Our analysis of sex ratios is the gold standard for treating game theory as a science

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Identify A Puzzle: Why are sex ratios at birth approximately 50:50 in most species?

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Use Data to Eliminate Alternative Explanations Ensure equal ratio at maturity? Maximize size of species?

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Use Data to Eliminate Alternative Explanations

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Use game theory to solve the puzzle (In a way that’s robust)

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Check that dynamics lead to Nash (In this case, evolution)

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Use the theory to find novel predictions If females require more parental investment, there will be more males in equilibrium

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Find Empirical Evidence to Support It

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Local mate competition In 1967 Hamilton described his classic LMC model: –When populations are structured such that mating takes place locally and related.

Local mate competition In 1967 Hamilton described his classic LMC model: –When populations are structured such that mating takes place locally and related.

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