The Selfish Gene

by

Richard Dawkins

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The Selfish Gene: Chapter 12 Summary & Analysis

Summary
Analysis
 “Nice guys finish last” is a common saying. But Dawkins thinks there’s also a sense in which nice guys finish first. He thinks about birds who are “grudgers” (those that pick parasites off other birds, but remember the ones that don’t return the favor and ignore them the next time around). That strategy actually beats out the “cheat” strategy (accepting help with parasites but not reciprocating). Dawkins thinks the “grudger” is the kind of “nice guy” who “finishes first.” This is the individual who engages in “reciprocal altruism.”   Dawkins agrees with Robert Axelrod and Hamilton that many wild animals are “engaged in ceaseless games of the Prisoner’s Dilemma, played out in evolutionary time,” which explains why nice guys finish first.
Dawkins revisits reciprocal altruism, because he wants to convince the reader more fully that there definitely isn’t altruism in nature. He uses the Prisoner’s Dilemma (which calculates optimal outcomes between individuals who interact without knowing what the other person will do next) to prove his case. The Prisoner’s Dilemma shows that the individual’s genes benefit more when the individual is nice at first. Dawkins thinks the phrase “nice guys finish first” captures the sense in which altruistic behavior is underscored by selfish genes.
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Quotes
The Prisoner’s Dilemma is a strategy game in which two people each have two cards. One card says “cooperate” and the other says “defect.” Both players have the same cards. Each chooses one card and puts it face-down on the table, and the cards are revealed at the same time. If both people play “cooperate,” they each win $300. If both people play “defect,” they each pay a $10 fine. If one person plays “cooperate” and the other plays “defect,” the person who plays “cooperate” has to pay $100, and the person who plays “defect” wins $500.   
In the Prisoner’s Dilemma game, two friends are interacting, but neither knows what the other one will do. They both win a lot of money if they “cooperate.” If both “defect,” they each lose a bit, but not very much. However, if one of them cooperates when the other defects, the person who cooperated loses everything, and the defector wins a lot (but only a bit more than if they both cooperated). The game is set up to test a person’s likelihood of cooperating (or being altruistic).
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Even though mutual cooperation is the best strategy (both people win money), most people will play “defect,” because the risk of being the only one to cooperate is too high. This is why the game is called a “dilemma.” The most logical thing to do is defect—it won’t yield the best potential outcome, but it’s the least risky strategy.  
The “dilemma” arises because cooperating is the best strategy for both players combined (both win). But when an individual cooperates, there’s a chance that the individual will lose everything if the other person defects. The risk of a big loss makes most people defect, even if they’d prefer to cooperate. In other words, the only way to avoid a big loss is to be selfish. 
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There’s another version of the game called the “Iterated (Repeated) Prisoner’s Dilemma.” In this version, the same players play the game over and over again, meaning they know what cards the other player chose in the previous round. Dawkins thinks the birds who were picking parasites off each other’s backs were playing the iterated Prisoner’s Dilemma, since they picked parasites multiple times and remembered the other birds’ strategies. 
The best strategies to play change a bit when the game is repeated (iterated) and each player knows what happened in the last round. Dawkins thinks the iterated game comes closest to what happens in nature, because many organisms can remember who helped them in the past and who didn’t, so this is the one he’ll use to explain why reciprocal altruism (or mutual cooperation) is ultimately selfish.
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Axelrod ran a competition where he programmed a computer to play all the possible strategies in the iterated Prisoner’s Dilemma (including one strategy that was simply “random”) 200 times. There are 15 possible strategies. It turns out that the winning strategy was “Tit for Tat,” meaning be nice on the first round, and in the next round play whatever card one’s opponent played in the previous round.
The best performing strategy when the game is repeated many times is “Tit for Tat.” Since it entails acting nice at first, then doing what one’s opponent does, Tit for Tat is somewhat similar to the “grudger” strategy in nature. Tit for Tat shows how being “nice” at first can yield the best selfish payoff.
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Axelrod ran the “Tit for Tat” strategy against a lot of other strategies, including some very similar ones. Two similar strategies are “Naïve Prober” (which is basically “Tit for Tat” with a random extra “Defect” thrown in now and again), and “Remorseful Prober” (which will randomly play “Defect,” but then play “Cooperate” on the next round). Dawkins calls these “nasty” strategies because they involve defecting first. Tit for Tat, however, is a “nice” strategy, because it only defects in retaliation. It turns out, that when ranked, the eight best strategies were “nice” strategies (in which the player never defects unprovoked), and the strategies that trailed behind were “nasty” strategies (in which the player does defect unprovoked, in some way or another).
Axelrod shows that even upon further testing, all the strategies that involve acting “nice” at first give players better outcomes than the strategies that begin with acting “nasty.” Dawkins stresses this to show that acting nice at first in nature (as grudgers do) is actually the best way for an individual to maximize their winnings in the game of life. This “nice” behavior happens in nature because the genes that program their survival machines to act that way are the most successful at keeping their survival machines alive.
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 “Tit for Tat” is also a forgiving strategy: one only play the card one’s opponent played on the previous round. The “grudger” strategy that birds play when picking parasites off each other, in contrast, is unforgiving. The bird remembers that another bird didn’t help them for the whole game and never helps them again, even if the cheating bird changes its mind and helps the grudger. It turns out that the “grudger” strategy ranked next to last in Axelrod’s simulation. Dawkins thinks this means that the best strategies are both “nice” and “forgiving.”
In fact, “Tit for Tat” shows that being more altruistic than grudgers are will yield the best results. Grudgers never help someone who betrayed them once before, but Tit for Tat, is more forgiving: individuals will switch back to being nice if the other player does. Dawkins implies that individuals who are more forgiving (altruistic) in nature would actually be the most successful.
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Axelrod called “Tit for Tat” a “robust” strategy. Dawkins calls it an evolutionarily stable strategy. Interestingly, “Tit for Tat” doesn’t win in an environment where most other strategies are nasty. Dawkins thinks this means that evolutionarily stable strategies work when there are a lot of others who act in similar ways. This is exactly what happens in nature, because winning strategies produce offspring that inherit similar behavior tendencies.   
Dawkins stresses that Axelrod’s research is a good model for explaining behaviors in nature, because it also explores scenarios where the players are mostly “nice” or mostly “nasty.” In nature, too, the base disposition of the population will vary depending on the behavioral tendencies that are inherited in a particular population. 
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Axelrod ran his Prisoner’s Dilemma game multiple times. Dawkins thinks each round of games is analogous to a generation. When multiple rounds are played it turns out that the winning strategy is whichever more or less “stable” one is dominant first.  If “always defect” is played often, and early in the game, it will dominate and become the evolutionarily stable strategy.
The Prisoner’s Dilemma shows that different behavior patterns can become evolutionarily stable strategies. The population will settle into a pattern that’s based on acting selfishly all the time, or acting nice at first. There’s no case, however, in which always being nice (or being purely altruistic) will persist from generation to generation.
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Dawkins wonders which strategy (between “Tit for Tat” and “always defect”) wins out in real life, if both are stable. “Tit for Tat” tends to do well when other people play the same strategy. He decides that since individuals tend to live near their kin, it’s likely that if one person tends to play “nice,” their genetic relatives will too, and “Tit for Tat” prevails. An “always defect” family would stay that way, but fair badly in the long run.
In the natural world, populations either “cooperate” and act nice at first, or “defect” and act selfish all the time. However, in the long run, the selfish populations are more likely to die out. Dawkins is implying that acting nice at first (or being somewhat altruistic) is actually the most selfish thing to do in the long run.
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Games are either “zero sum” games (meaning one side wins and the other loses), or “nonzero sum” games (meaning both sides can win). Football, chess, rugby, and tennis are all zero sum games—usually there’s one winner and one loser. Dawkins thinks that many situations in nature are nonzero sum games. If both sides cooperate, they can both win, which explains how “cooperation and mutual assistance can flourish even in a basically selfish world.”
The Prisoner’s Dilemma, like nature, is a “nonzero sum game,” meaning both sides can win. Dawkins reiterates that when mutual cooperation does happen in nature, it only “flourishes” because both sides win big in this scenario. Again, what looks like altruism is explained in terms of the selfish payoffs to those interacting. 
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Dawkins talks about times during World War I when German and British soldiers were nice to each other between battles, even though they were enemies in the war. One Christmas, for example, they put down their guns, drank wine, and celebrated together on the battleground. Dawkins wonders if niceness can evolve in nature in the ways that Axelrod’s simulation games showed (and how the World War I soldiers behaved). Dawkins concludes that it can, if the game is nonzero sum, and if it’s iterated (repeated).
Dawkins thinks that organisms from different species who interact frequently can play the “cooperate” strategy in the game of nature, just as the British and German soldiers did at Christmas during World War I. The Prisoner’s Dilemma shows that cooperation only happens because it’s self-serving: the genetic benefit of mutual cooperation is higher than that of defecting.
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For example, fig trees and fig wasps cooperate. Fig wasps pollinate fig trees, and fig trees provide nourishment for fig wasp eggs. Both sides could be exploitative (or “defect”)—if the wasps lay too many eggs, or if the trees shed flowers with eggs in them—but they “cooperate” because over time (for generations) the benefit to each is higher than if they exploit each other. 
Fig trees and fig wasps exemplify an iterated Prisoner’s Dilemma game in which both players “cooperate.” Fig wasp genes and fig tree genes both benefit more when their survival machines cooperate. Mutual cooperation is an evolutionarily stable strategy because it keeps both sets of genes in the gene pool over multiple generations.
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 Wilkinson’s research on vampire bats indicates that the “cooperate” strategy comes up elsewhere in nature, as well. Wilkinson discovered that vampire bats who are able to feed often donate their food (which is blood) to other bats. He examined 110 donations, and found that 33 were to non-relatives. He then discovered that bats can recognize each other. Dawkins thinks these 33 examples were the “Tit for Tat” policy in action. The bats were sharing their meals in case they needed help next time. Dawkins thinks that he has shown that even in a world driven by selfish genes, “nice guys can finish first.”
Wilkinson’s research shows that a vampire bat who seems altruistic by sharing her food with non-relatives is actually only cooperating so that other bats are more likely return the favor in the future, which might come in handy if she doesn’t score a big meal herself one night. The vampire bat’s supposed altruism, once again, comes hand in hand with a hefty selfish benefit to her genes. Dawkins thinks he has covered enough examples to convince the reader that reciprocal altruism only happens in nature when the individual rewards are higher than acting selfish, meaning these actions only seem altruistic on the surface. 
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