The Effect of Income and Wealth on Behavioral Strategies, Personality Traits, and Preferences.

Individuals living in either harsh or favorable environments display well-documented psychological and behavioral differences. For example, people in favorable environments tend to be more future-oriented, trust strangers more, and have more explorative preferences. To account for such differences, psychologists have turned to evolutionary biology and behavioral ecology, in particular, the literature on life-history theory and pace-of-life syndrome. However, critics have found that the theoretical foundations of these approaches are fragile and that differences in life expectancy cannot explain vast psychological and behavioral differences. In this article, we build on the theory of optimal resource allocation to propose an alternative framework. We hypothesize that the quantity of resources available, such as income, has downstream consequences on psychological traits, leading to the emergence of behavioral syndromes. We show that more resources lead to more long-term orientation, more tolerance of variance, and more investment in low marginal-benefit needs. At the behavioral level, this translates, among others, into more large-scale cooperation, more investment in health, and more exploration. These individual-level differences in behavior, in turn, account for cultural phenomena such as puritanism, authoritarianism, and innovation.

Peace is a form of cooperation, and so are the cultural technologies which make peace possible.

While necessary parts of the puzzle, cultural technologies are insufficient to explain peace. They are a form of second-order cooperation - a cooperative interaction designed to incentivize first-order cooperation. We propose an explanation for peacemaking cultural technologies, and therefore peace, based on the reputational incentives for second-order cooperation.

Reciprocal contracts - not competitive acquisition - explain the moral psychology of ownership.

We applaud Boyer's attempt to ground the psychology of ownership partly in a cooperative logic. In this commentary, we propose to go further and ground the psychology of ownership solely in a cooperative logic. The predictions of bargaining theory, we argue, completely contradict the actual features of ownership intuitions. Ownership is only about the calculation of mutually beneficial, reciprocal contracts.

The puritanical moral contract: Purity, cooperation, and the architecture of the moral mind.

Commentators raise fundamental questions about the notion of purity (sect. R1), the architecture of moral cognition (sect. R2), the functional relationship between morality and cooperation (sect. R3), the role of folk-theories of self-control in moral judgment (sect. R4), and the cultural variation of morality (sect. R5). In our response, we address all these issues by clarifying our theory of puritanism, responding to counter-arguments, and incorporating welcome corrections and extensions.

Cultural evolution from the producers' standpoint.

Standard approaches to cultural evolution focus on the recipients or consumers. This does not take into account the fitness costs incurred in producing the behaviours or artefacts that become cultural, i.e. widespread in a social group. We argue that cultural evolution models should focus on these fitness costs and benefits of cultural production, particularly in the domain of 'symbolic' culture. In this approach, cultural products can be considered as a part of the extended phenotype of producers, which can affect the fitness of recipients in a positive way (through cooperation) but also in a detrimental way (through manipulation and exploitation). Taking the producers' perspective may help explain the specific features of many kinds of cultural products.

Human-environment feedback and the consistency of proenvironmental behavior.

Addressing global environmental crises such as anthropogenic climate change requires the consistent adoption of proenvironmental behavior by a large part of a population. Here, we develop a mathematical model of a simple behavior-environment feedback loop to ask how the individual assessment of the environmental state combines with social interactions to influence the consistent adoption of proenvironmental behavior, and how this feeds back to the perceived environmental state. In this stochastic individual-based model, individuals can switch between two behaviors, 'active' (or actively proenvironmental) and 'baseline', differing in their perceived cost (higher for the active behavior) and environmental impact (lower for the active behavior). We show that the deterministic dynamics and the stochastic fluctuations of the system can be approximated by ordinary differential equations and a Ornstein-Uhlenbeck type process. By definition, the proenvironmental behavior is adopted consistently when, at population stationary state, its frequency is high and random fluctuations in frequency are small. We find that the combination of social and environmental feedbacks can promote the spread of costly proenvironmental behavior when neither, operating in isolation, would. To be adopted consistently, strong social pressure for proenvironmental action is necessary but not sufficient-social interactions must occur on a faster timescale compared to individual assessment, and the difference in environmental impact must be small. This simple model suggests a scenario to achieve large reductions in environmental impact, which involves incrementally more active and potentially more costly behavior being consistently adopted under increasing social pressure for proenvironmentalism.

Moral disciplining: The cognitive and evolutionary foundations of puritanical morality.

Why do many societies moralize apparently harmless pleasures, such as lust, gluttony, alcohol, drugs, and even music and dance? Why do they erect temperance, asceticism, sobriety, modesty, and piety as cardinal moral virtues? According to existing theories, this puritanical morality cannot be reduced to concerns for harm and fairness: It must emerge from cognitive systems that did not evolve for cooperation (e.g., disgust-based "purity" concerns). Here, we argue that, despite appearances, puritanical morality is no exception to the cooperative function of moral cognition. It emerges in response to a key feature of cooperation, namely that cooperation is (ultimately) a long-term strategy, requiring (proximately) the self-control of appetites for immediate gratification. Puritanical moralizations condemn behaviors which, although inherently harmless, are perceived as indirectly facilitating uncooperative behaviors, by impairing the self-control required to refrain from cheating. Drinking, drugs, immodest clothing, and unruly music and dance are condemned as stimulating short-term impulses, thus facilitating uncooperative behaviors (e.g., violence, adultery, free-riding). Overindulgence in harmless bodily pleasures (e.g., masturbation, gluttony) is perceived as making people slave to their urges, thus altering abilities to resist future antisocial temptations. Daily self-discipline, ascetic temperance, and pious ritual observance are perceived as cultivating the self-control required to honor prosocial obligations. We review psychological, historical, and ethnographic evidence supporting this account. We use this theory to explain the fall of puritanism in western, educated, industrialized, rich, and democratic (WEIRD) societies, and discuss the cultural evolution of puritanical norms. Explaining puritanical norms does not require adding mechanisms unrelated to cooperation in our models of the moral mind.

Policy search with rare significant events: Choosing the right partner to cooperate with.

This paper focuses on a class of reinforcement learning problems where significant events are rare and limited to a single positive reward per episode. A typical example is that of an agent who has to choose a partner to cooperate with, while a large number of partners are simply not interested in cooperating, regardless of what the agent has to offer. We address this problem in a continuous state and action space with two different kinds of search methods: a gradient policy search method and a direct policy search method using an evolution strategy. We show that when significant events are rare, gradient information is also scarce, making it difficult for policy gradient search methods to find an optimal policy, with or without a deep neural architecture. On the other hand, we show that direct policy search methods are invariant to the rarity of significant events, which is yet another confirmation of the unique role evolutionary algorithms has to play as a reinforcement learning method.

Cooperation as a signal of time preferences.

Many evolutionary models explain why we cooperate with non-kin, but few explain why cooperative behaviour and trust vary. Here, we introduce a model of cooperation as a signal of time preferences, which addresses this variability. At equilibrium in our model (i) future-oriented individuals are more motivated to cooperate, (ii) future-oriented populations have access to a wider range of cooperative opportunities, and (iii) spontaneous and inconspicuous cooperation reveal stronger preference for the future, and therefore inspire more trust. Our theory sheds light on the variability of cooperative behaviour and trust. Since affluence tends to align with time preferences, results (i) and (ii) explain why cooperation is often associated with affluence, in surveys and field studies. Time preferences also explain why we trust others based on proxies for impulsivity, and, following result (iii), why uncalculating, subtle and one-shot cooperators are deemed particularly trustworthy. Time preferences provide a powerful and parsimonious explanatory lens, through which we can better understand the variability of trust and cooperation.

The evolution of music: One trait, many ultimate-level explanations.

We propose an approach reconciling the ultimate-level explanations proposed by Savage et al. and Mehr et al. as to why music evolved. We also question the current adaptationist view of culture, which too often fails to disentangle distinct fitness benefits.

Nothing better to do? Environment quality and the evolution of cooperation by partner choice.

The effects of partner choice have been documented in a large number of biological systems such as sexual markets, interspecific mutualisms, or human cooperation. There are, however, a number of situations in which one would expect this mechanism to play a role, but where no such effect has ever been demonstrated. This is the case in particular in many intraspecific interactions, such as collective hunts, in non-human animals. Here we use individual-based simulations to solve this apparent paradox. We show that the conditions for partner choice to operate are in fact restrictive. They entail that individuals can compare social opportunities and choose the best. The challenge is that social opportunities are often rare because they necessitate the co-occurrence of (i) at least one available partner, and (ii) a resource to exploit together with this partner. This has three consequences. First, partner choice cannot lead to the evolution of cooperation when resources are scarce, which explains that this mechanism could never be observed in many cases of intraspecific cooperation in animals. Second, partner choice can operate when partners constitute in themselves a resource, which is the case in sexual interactions and interspecific mutualisms. Third, partner choice can lead to the evolution of cooperation when individuals live in a rich environment, and/or when they are highly efficient at extracting resources from their environment.

The emergence of cooperation by evolutionary generalization.

In principle, any cooperative behaviour can be evolutionarily stable as long as it is incentivized by a reward from the beneficiary, a mechanism that has been called reciprocal cooperation. However, what makes this mechanism so powerful also has an evolutionary downside. Reciprocal cooperation faces a chicken-and-egg problem of the same kind as communication: it requires two functions to evolve at the same time-cooperation and response to cooperation. As a result, it can only emerge if one side first evolves for another reason, and is then recycled into a reciprocal function. Developing an evolutionary model in which we make use of machine learning techniques, we show that this occurs if the fact to cooperate and reward others' cooperation become general abilities that extend beyond the set of contexts for which they have initially been selected. Drawing on an evolutionary analogy with the concept of generalization, we identify the conditions necessary for this to happen. This allows us to understand the peculiar distribution of reciprocal cooperation in the wild, virtually absent in most species-or limited to situations where individuals have partially overlapping interests, but pervasive in the human species.

Indirect genetic effects allow escape from the inefficient equilibrium in a coordination game.

Social interactions involving coordination between individuals are subject to an "evolutionary trap." Once a suboptimal strategy has evolved, mutants playing an alternative strategy are counterselected because they fail to coordinate with the majority. This creates a detrimental situation from which evolution cannot escape, preventing the evolution of efficient collective behaviors. Here, we study this problem using evolutionary robotics simulations. We first confirm the existence of an evolutionary trap in a simple setting. We then, however, reveal that evolution can solve this problem in a more realistic setting where individuals need to coordinate with one another. In this setting, simulated robots evolve an ability to adapt plastically their behavior to one another, as this improves the efficiency of their interaction. This ability has an unintended evolutionary consequence: a genetic mutation affecting one individual's behavior also indirectly alters their partner's behavior because the two individuals influence one another. As a consequence of this indirect genetic effect, pairs of partners can virtually change strategy together with a single mutation, and the evolutionary barrier between alternative strategies disappears. This finding reveals a general principle that could play a role in nature to smoothen the transition to efficient collective behaviors in all games with multiple equilibriums.

Cultural evolution by capital accumulation.

In this article, we model cultural knowledge as a capital in which individuals invest at a cost. To this end, following other models of cultural evolution, we explicitly consider the investments made by individuals in culture as life history decisions. Our aim is to understand what then determines the dynamics of cultural accumulation. We show that culture can accumulate provided it improves the efficiency of people's lives in such a way as to increase their productivity or, said differently, provided the knowledge created by previous generations improves the ability of subsequent generations to invest in new knowledge. Our central message is that this positive feedback allowing cultural accumulation can occur for many different reasons. It can occur if cultural knowledge increases people's productivity, including in domains that have no connection with knowledge, because it frees up time that people can then spend learning and/or innovating. We also show that it can occur if cultural knowledge, and thus the higher level of resources that results from increased productivity, leads individuals to modify their life history decisions through phenotypic plasticity. Finally, we show that it can occur if technical knowledge reduces the effective cost of its own acquisition via division of labour. These results suggest that culture should not be defined only as a set of knowledge and skills but, more generally, as all the capital that has been produced by previous generations and that continues to affect current generations.

Why cooperation is not running away.

A growing number of experimental and theoretical studies show the importance of partner choice as a mechanism to promote the evolution of cooperation, especially in humans. In this paper, we focus on the question of the precise quantitative level of cooperation that should evolve under this mechanism. When individuals compete to be chosen by others, their level of investment in cooperation evolves towards higher values, a process called competitive altruism, or runaway cooperation. Using a classic adaptive dynamics model, we first show that when the cost of changing partner is low, this runaway process can lead to a profitless escalation of cooperation. In the extreme, when partner choice is entirely frictionless, cooperation even increases up to a level where its cost entirely cancels out its benefit. That is, at evolutionary equilibrium, individuals gain the same payoff than if they had not cooperated at all. Second, importing models from matching theory in economics we, however, show that when individuals can plastically modulate their choosiness in function of their own cooperation level, partner choice stops being a runaway competition to outbid others and becomes a competition to form the most optimal partnerships. In this case, when the cost of changing partner tends towards zero, partner choice leads to the evolution of the socially optimum level of cooperation. This last result could explain the observation that human cooperation seems to be often constrained by considerations of social efficiency.

Fairness, more than any other cognitive mechanism, is what explains the content of folk-economic beliefs.

We applaud Boyer & Petersen's (B&P's) article on economic folk beliefs. We believe that it is crucial for the future of democracy to identify the cognitive systems through which people form their beliefs about the working of the economy. In this commentary, we put forward the idea that, although many systems are involved, fairness is probably the main driver of folk-economic beliefs.

Correction: On the evolutionary origins of equity.

[This corrects the article DOI: 10.1371/journal.pone.0173636.].

Grandmothering and cognitive resources are required for the emergence of menopause and extensive post-reproductive lifespan.

Menopause, the permanent cessation of ovulation, occurs in humans well before the end of the expected lifespan, leading to an extensive post-reproductive period which remains a puzzle for evolutionary biologists. All human populations display this particularity; thus, it is difficult to empirically evaluate the conditions for its emergence. In this study, we used artificial neural networks to model the emergence and evolution of allocation decisions related to reproduction in simulated populations. When allocation decisions were allowed to freely evolve, both menopause and extensive post-reproductive life-span emerged under some ecological conditions. This result allowed us to test various hypotheses about the required conditions for the emergence of menopause and extensive post-reproductive life-span. Our findings did not support the Maternal Hypothesis (menopause has evolved to avoid the risk of dying in childbirth, which is higher in older women). In contrast, results supported a shared prediction from the Grandmother Hypothesis and the Embodied Capital Model. Indeed, we found that extensive post-reproductive lifespan allows resource reallocation to increase fertility of the children and survival of the grandchildren. Furthermore, neural capital development and the skill intensiveness of the foraging niche, rather than strength, played a major role in shaping the age profile of somatic and cognitive senescence in our simulated populations. This result supports the Embodied Capital Model rather than the Grand-Mother Hypothesis. Finally, in simulated populations where menopause had already evolved, we found that reduced post-reproductive lifespan lead to reduced children's fertility and grandchildren's survival. The results are discussed in the context of the evolutionary emergence of menopause and extensive post-reproductive life-span.

On the evolutionary origins of equity.

Equity, defined as reward according to contribution, is considered a central aspect of human fairness in both philosophical debates and scientific research. Despite large amounts of research on the evolutionary origins of fairness, the evolutionary rationale behind equity is still unknown. Here, we investigate how equity can be understood in the context of the cooperative environment in which humans evolved. We model a population of individuals who cooperate to produce and divide a resource, and choose their cooperative partners based on how they are willing to divide the resource. Agent-based simulations, an analytical model, and extended simulations using neural networks provide converging evidence that equity is the best evolutionary strategy in such an environment: individuals maximize their fitness by dividing benefits in proportion to their own and their partners' relative contribution. The need to be chosen as a cooperative partner thus creates a selection pressure strong enough to explain the evolution of preferences for equity. We discuss the limitations of our model, the discrepancies between its predictions and empirical data, and how interindividual and intercultural variability fit within this framework.

Both collection risk and waiting costs give rise to the behavioral constellation of deprivation.

Pepper & Nettle explain the behavioral constellation of deprivation (BCD) in terms of differences in collection risk (i.e., the probability of collecting a reward after some delay) between high- and low-socioeconomic-status (SES) populations. We argue that a proper explanation should also include the costs of waiting per se, which are paid even when the benefits are guaranteed.