The restart effect in social dilemmas shows humans are self-interested not altruistic.

Do economic games show evidence of altruistic or self-interested motivations in humans? A huge body of empirical work has found contrasting results. While many participants routinely make costly decisions that benefit strangers, consistent with the hypothesis that humans exhibit a biologically novel form of altruism (or "prosociality"), many participants also typically learn to pay fewer costs with experience, consistent with self-interested individuals adapting to an unfamiliar environment. Key to resolving this debate is explaining the famous "restart effect," a puzzling enigma whereby failing cooperation in public goods games can be briefly rescued by a surprise restart. Here we replicate this canonical result, often taken as evidence of uniquely human altruism, and show that it 1) disappears when cooperation is invisible, meaning individuals can no longer affect the behavior of their groupmates, consistent with strategically motivated, self-interested, cooperation; and 2) still occurs even when individuals are knowingly grouped with computer players programmed to replicate human decisions, consistent with confusion. These results show that the restart effect can be explained by a mixture of self-interest and irrational beliefs about the game's payoffs, and not altruism. Consequently, our results suggest that public goods games have often been measuring self-interested but confused behaviors and reject the idea that conventional theories of evolution cannot explain the results of economic games.

Self-interested learning is more important than fair-minded conditional cooperation in public-goods games.

Why does human cooperation often unravel in economic experiments despite a promising start? Previous studies have interpreted the decline as the reaction of disappointed altruists retaliating in response to non-altruists (Conditional Cooperators hypothesis). This interpretation has been considered evidence of a uniquely human form of cooperation, motivated by an altruistic concern for equality ('fairness') and requiring special evolutionary explanations. However, experiments have typically shown individuals not only information about the decisions of their groupmates (social information) but also information about their own payoffs. Showing both confounds explanations based on conditional cooperation with explanations based on confused individuals learning how to better play the game (Confused Learners hypothesis). Here we experimentally decouple these two forms of information, and thus these two hypotheses, in a repeated public-goods game. Analysing 616 Swiss university participants, we find that payoff information leads to a greater decline, supporting the Confused Learners hypothesis. In contrast, social information has a small or negligible effect, contradicting the Conditional Cooperators hypothesis. We also find widespread evidence of both confusion and selfish motives, suggesting that human cooperation is maybe not so unique after all.

The Black Box as a Control for Payoff-Based Learning in Economic Games.

The black box method was developed as an "asocial control" to allow for payoff-based learning while eliminating social responses in repeated public goods games. Players are told they must decide how many virtual coins they want to input into a virtual black box that will provide uncertain returns. However, in truth, they are playing with each other in a repeated social game. By "black boxing" the game's social aspects and payoff structure, the method creates a population of self-interested but ignorant or confused individuals that must learn the game's payoffs. This low-information environment, stripped of social concerns, provides an alternative, empirically derived null hypothesis for testing social behaviours, as opposed to the theoretical predictions of rational self-interested agents (Homo economicus). However, a potential problem is that participants can unwittingly affect the learning of other participants. Here, we test a solution to this problem in a range of public goods games by making participants interact, unknowingly, with simulated players ("computerised black box"). We find no significant differences in rates of learning between the original and the computerised black box, therefore either method can be used to investigate learning in games. These results, along with the fact that simulated agents can be programmed to behave in different ways, mean that the computerised black box has great potential for complementing studies of how individuals and groups learn under different environments in social dilemmas.

A preference to learn from successful rather than common behaviours in human social dilemmas.

Human cooperation is often claimed to be special and requiring explanations based on gene-culture coevolution favouring a desire to copy common social behaviours. If this is true, then individuals should be motivated to both observe and copy common social behaviours. Previous economic experiments, using the public goods game, have suggested individuals' desire to sacrifice for the common good and to copy common social behaviours. However, previous experiments have often not shown examples of success. Here we test, on 489 participants, whether individuals are more motivated to learn about, and more likely to copy, either common or successful behaviours. Using the same social dilemma and standard instructions, we find that individuals were primarily motivated to learn from successful rather than common behaviours. Consequently, social learning disfavoured costly cooperation, even when individuals could observe a stable, pro-social level of cooperation. Our results call into question explanations for human cooperation based on cultural evolution and/or a desire to conform with common social behaviours. Instead, our results indicate that participants were motivated by personal gain, but initially confused, despite receiving standard instructions. When individuals could learn from success, they learned to cooperate less, suggesting that human cooperation is maybe not so special after all.

Decoupling cooperation and punishment in humans shows that punishment is not an altruistic trait.

Economic experiments have suggested that cooperative humans will altruistically match local levels of cooperation (conditional cooperation) and pay to punish non-cooperators (altruistic punishment). Evolutionary models have suggested that if altruists punish non-altruists this could favour the evolution of costly helping behaviours (cooperation) among strangers. An often-key requirement is that helping behaviours and punishing behaviours form one single conjoined trait (strong reciprocity). Previous economics experiments have provided support for the hypothesis that punishment and cooperation form one conjoined, altruistically motivated, trait. However, such a conjoined trait may be evolutionarily unstable, and previous experiments have confounded a fear of being punished with being surrounded by cooperators, two factors that could favour cooperation. Here, we experimentally decouple the fear of punishment from a cooperative environment and allow cooperation and punishment behaviour to freely separate (420 participants). We show, that if a minority of individuals is made immune to punishment, they (i) learn to stop cooperating on average despite being surrounded by high levels of cooperation, contradicting the idea of conditional cooperation and (ii) often continue to punish, 'hypocritically', showing that cooperation and punishment do not form one, altruistically motivated, linked trait.

Payoff-based learning best explains the rate of decline in cooperation across 237 public-goods games.

What motivates human behaviour in social dilemmas? The results of public goods games are commonly interpreted as showing that humans are altruistically motivated to benefit others. However, there is a competing 'confused learners' hypothesis: that individuals start the game either uncertain or mistaken (confused) and then learn from experience how to improve their payoff (payoff-based learning). Here we (1) show that these competing hypotheses can be differentiated by how they predict contributions should decline over time; and (2) use metadata from 237 published public goods games to test between these competing hypotheses. We found, as predicted by the confused learners hypothesis, that contributions declined faster when individuals had more influence over their own payoffs. This predicted relationship arises because more influence leads to a greater correlation between contributions and payoffs, facilitating learning. Our results suggest that humans, in general, are not altruistically motivated to benefit others but instead learn to help themselves.

Evidence for strategic cooperation in humans.

Humans may cooperate strategically, cooperating at higher levels than expected from their short-term interests, to try and stimulate others to cooperate. To test this hypothesis, we experimentally manipulated the extent an individual's behaviour is known to others, and hence whether or not strategic cooperation is possible. In contrast with many previous studies, we avoided confounding factors by preventing individuals from learning during the game about either pay-offs or about how other individuals behave. We found clear evidence for strategic cooperators-just telling some individuals that their groupmates would be informed about their behaviour led to them tripling their initial level of cooperation, from 17 to 50%. We also found that many individuals play as if they do not understand the game, and their presence obscures the detection of strategic cooperation. Identifying such players allowed us to detect and study strategic motives for cooperation in novel, more powerful, ways.

Social learning and the demise of costly cooperation in humans.

Humans have a sophisticated ability to learn from others, termed social learning, which has allowed us to spread over the planet, construct complex societies, and travel to the moon. It has been hypothesized that social learning has played a pivotal role in making human societies cooperative, by favouring cooperation even when it is not favoured by genetical selection. However, this hypothesis lacks direct experimental testing, and the opposite prediction has also been made, that social learning disfavours cooperation. We experimentally tested how different aspects of social learning affect the level of cooperation in public-goods games. We found that: (i) social information never increased cooperation and usually led to decreased cooperation; (ii) cooperation was lowest when individuals could observe how successful individuals behaved; and (iii) cooperation declined because individuals preferred to copy successful individuals, who cooperated less, rather than copy common behaviours. Overall, these results suggest that individuals use social information to try and improve their own success, and that this can lead to lower levels of cooperation.

Conditional cooperation and confusion in public-goods experiments.

Economic experiments are often used to study if humans altruistically value the welfare of others. A canonical result from public-good games is that humans vary in how they value the welfare of others, dividing into fair-minded conditional cooperators, who match the cooperation of others, and selfish noncooperators. However, an alternative explanation for the data are that individuals vary in their understanding of how to maximize income, with misunderstanding leading to the appearance of cooperation. We show that (i) individuals divide into the same behavioral types when playing with computers, whom they cannot be concerned with the welfare of; (ii) behavior across games with computers and humans is correlated and can be explained by variation in understanding of how to maximize income; (iii) misunderstanding correlates with higher levels of cooperation; and (iv) standard control questions do not guarantee understanding. These results cast doubt on certain experimental methods and demonstrate that a common assumption in behavioral economics experiments, that choices reveal motivations, will not necessarily hold.

Payoff-based learning explains the decline in cooperation in public goods games.

Economic games such as the public goods game are increasingly being used to measure social behaviours in humans and non-human primates. The results of such games have been used to argue that people are pro-social, and that humans are uniquely altruistic, willingly sacrificing their own welfare in order to benefit others. However, an alternative explanation for the empirical observations is that individuals are mistaken, but learn, during the game, how to improve their personal payoff. We test between these competing hypotheses, by comparing the explanatory power of different behavioural rules, in public goods games, where individuals are given different amounts of information. We find: (i) that individual behaviour is best explained by a learning rule that is trying to maximize personal income; (ii) that conditional cooperation disappears when the consequences of cooperation are made clearer; and (iii) that social preferences, if they exist, are more anti-social than pro-social.

The evolution of altruism in humans.

Humans are an intensely social species, frequently performing costly behaviors that benefit others. Efforts to solve the evolutionary puzzle of altruism have a lengthy history, and recent years have seen many important advances across a range of disciplines. Here we bring together this interdisciplinary body of research and review the main theories that have been proposed to explain human prosociality, with an emphasis on kinship, reciprocity, indirect reciprocity, punishment, and morality. We highlight recent methodological advances that are stimulating research and point to some areas that either remain controversial or merit more attention.

Cooperation, quorum sensing, and evolution of virulence in Staphylococcus aureus.

The virulence and fitness in vivo of the major human pathogen Staphylococcus aureus are associated with a cell-to-cell signaling mechanism known as quorum sensing (QS). QS coordinates the production of virulence factors via the production and sensing of autoinducing peptide (AIP) signal molecules by the agr locus. Here we show, in a wax moth larva virulence model, that (i) QS in S. aureus is a cooperative social trait that provides a benefit to the local population of cells, (ii) agr mutants, which do not produce or respond to QS signal, are able to exploit the benefits provided by the QS of others ("cheat"), allowing them to increase in frequency when in mixed populations with cooperators, (iii) these social interactions between cells determine virulence, with the host mortality rate being negatively correlated to the percentage of agr mutants ("cheats") in a population, and (iv) a higher within-host relatedness (lower strain diversity) selects for QS and hence higher virulence. Our results provide an explanation for why agr mutants show reduced virulence in animal models but can be isolated from infections of humans. More generally, by providing the first evidence that QS is a cooperative social behavior in a Gram-positive bacterium, our results suggest convergent, and potentially widespread, evolution for signaling to coordinate cooperation in bacteria.

Prosocial preferences do not explain human cooperation in public-goods games.

It has become an accepted paradigm that humans have "prosocial preferences" that lead to higher levels of cooperation than those that would maximize their personal financial gain. However, the existence of prosocial preferences has been inferred post hoc from the results of economic games, rather than with direct experimental tests. Here, we test how behavior in a public-goods game is influenced by knowledge of the consequences of actions for other players. We found that (i) individuals cooperate at similar levels, even when they are not informed that their behavior benefits others; (ii) an increased awareness of how cooperation benefits others leads to a reduction, rather than an increase, in the level of cooperation; and (iii) cooperation can be either lower or higher than expected, depending on experimental design. Overall, these results contradict the suggested role of the prosocial preferences hypothesis and show how the complexity of human behavior can lead to misleading conclusions from controlled laboratory experiments.

Kin selection, quorum sensing and virulence in pathogenic bacteria.

Bacterial growth and virulence often depends upon the cooperative release of extracellular factors excreted in response to quorum sensing (QS). We carried out an in vivo selection experiment in mice to examine how QS evolves in response to variation in relatedness (strain diversity), and the consequences for virulence. We started our experiment with two bacterial strains: a wild-type that both produces and responds to QS signal molecules, and a lasR (signal-blind) mutant that does not release extracellular factors in response to signal. We found that: (i) QS leads to greater growth within hosts; (ii) high relatedness favours the QS wild-type; and (iii) low relatedness favours the lasR mutant. Relatedness matters in our experiment because, at relatively low relatedness, the lasR mutant is able to exploit the extracellular factors produced by the cells that respond to QS, and hence increase in frequency. Furthermore, our results suggest that because a higher relatedness favours cooperative QS, and hence leads to higher growth, this will also lead to a higher virulence, giving a relationship between relatedness and virulence that is in the opposite direction to that usually predicted by virulence theory.

Correlates of cooperation in a one-shot high-stakes televised prisoners' dilemma.

Explaining cooperation between non-relatives is a puzzle for both evolutionary biology and the social sciences. In humans, cooperation is often studied in a laboratory setting using economic games such as the prisoners' dilemma. However, such experiments are sometimes criticized for being played for low stakes and by misrepresentative student samples. Golden balls is a televised game show that uses the prisoners' dilemma, with a diverse range of participants, often playing for very large stakes. We use this non-experimental dataset to investigate the factors that influence cooperation when "playing" for considerably larger stakes than found in economic experiments. The game show has earlier stages that allow for an analysis of lying and voting decisions. We found that contestants were sensitive to the stakes involved, cooperating less when the stakes were larger in both absolute and relative terms. We also found that older contestants were more likely to cooperate, that liars received less cooperative behavior, but only if they told a certain type of lie, and that physical contact was associated with reduced cooperation, whereas laughter and promises were reliable signals or cues of cooperation, but were not necessarily detected.

Resistance to extreme strategies, rather than prosocial preferences, can explain human cooperation in public goods games.

The results of numerous economic games suggest that humans behave more cooperatively than would be expected if they were maximizing selfish interests. It has been argued that this is because individuals gain satisfaction from the success of others, and that such prosocial preferences require a novel evolutionary explanation. However, in previous games, imperfect behavior would automatically lead to an increase in cooperation, making it impossible to decouple any form of mistake or error from prosocial cooperative decisions. Here we empirically test between these alternatives by decoupling imperfect behavior from prosocial preferences in modified versions of the public goods game, in which individuals would maximize their selfish gain by completely (100%) cooperating. We found that, although this led to higher levels of cooperation, it did not lead to full cooperation, and individuals still perceived their group mates as competitors. This is inconsistent with either selfish or prosocial preferences, suggesting that the most parsimonious explanation is imperfect behavior triggered by psychological drives that can prevent both complete defection and complete cooperation. More generally, our results illustrate the caution that must be exercised when interpreting the evolutionary implications of economic experiments, especially the absolute level of cooperation in a particular treatment.

Evolutionary cooperation: male cleaner fish aggression may promote female cooperation.

A new study has shown that mixed-sex pairs of cleaner fish provide a better - more cooperative - service than singletons despite pairs facing an apparent Prisoner's dilemma.

Facultative sex ratio adjustment in natural populations of wasps: cues of local mate competition and the precision of adaptation.

Sex ratio theory offers excellent opportunities to examine the extent to which individuals adaptively adjust their behavior in response to local conditions. Hamilton's theory of local mate competition, which predicts female-biased sex ratios in structured populations, has been extended in numerous directions to predict individual behavior in response to factors such as relative fecundity, time of oviposition, and relatedness between cofoundresses and between mates. These extended models assume that foundresses use different sources of information, and they have generally been untested or have only been tested in the laboratory. We use microsatellite markers to describe the wild oviposition behavior of individual foundresses in natural populations of the parasitoid wasp Nasonia vitripennis, and we use the data collected to test these various models. The offspring sex ratio produced by a foundress on a particular host reflected the number of eggs that were laid on that host relative to the number of eggs that were laid on that host by other foundresses. In contrast, the offspring sex ratio was not directly influenced by other potentially important factors, such as the number of foundresses laying eggs on that patch, relative fecundity at the patch level, or relatedness to either a mate or other foundresses on the patch.