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.

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.

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.

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.

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.

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 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.

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.

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.

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.

To Cooperate or Not to Cooperate: Why Behavioural Mechanisms Matter.

Mutualistic cooperation often requires multiple individuals to behave in a coordinated fashion. Hence, while the evolutionary stability of mutualistic cooperation poses no particular theoretical difficulty, its evolutionary emergence faces a chicken and egg problem: an individual cannot benefit from cooperating unless other individuals already do so. Here, we use evolutionary robotic simulations to study the consequences of this problem for the evolution of cooperation. In contrast with standard game-theoretic results, we find that the transition from solitary to cooperative strategies is very unlikely, whether interacting individuals are genetically related (cooperation evolves in 20% of all simulations) or unrelated (only 3% of all simulations). We also observe that successful cooperation between individuals requires the evolution of a specific and rather complex behaviour. This behavioural complexity creates a large fitness valley between solitary and cooperative strategies, making the evolutionary transition difficult. These results reveal the need for research on biological mechanisms which may facilitate this transition.

Inclusive fitness theory and eusociality.

Arising from M. A. Nowak, C. E. Tarnita & E. O. Wilson 466, 1057-1062 (2010); Nowak et al. reply. Nowak et al. argue that inclusive fitness theory has been of little value in explaining the natural world, and that it has led to negligible progress in explaining the evolution of eusociality. However, we believe that their arguments are based upon a misunderstanding of evolutionary theory and a misrepresentation of the empirical literature. We will focus our comments on three general issues.

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.

Contingency in the evolutionary emergence of reciprocal cooperation.

Reciprocity is characterized by individuals actively making it beneficial for others to cooperate by responding to them. This makes it a particularly powerful generator of mutual interest, because the benefits accrued by an individual can be redistributed to another. However, reciprocity is a composite biological function, entailing at least two subfunctions: (i) a behavioral ability to provide fitness benefits to others and (ii) a cognitive ability to evaluate the benefits received from others. For reciprocity to evolve, these two subfunctions must appear together, which raises an evolutionary problem of bootstrapping. In this article, I develop mathematical models to study the necessary conditions for the gradual emergence of reciprocity in spite of this bootstrapping problem. I show that the evolution of reciprocity is based on three conditions. First, there must be some variability in behavior. Second, cooperation must pre-evolve for reasons independent of reciprocity. Third, and most significantly, selection favors conditional cooperation only if the cooperation expressed by others is already conditional, that is, if some reciprocity is already present in the first place. In the discussion, I show that these three conditions help explain the specific features of the instances in which reciprocity does occur in the wild. For instance, it accounts for the role of spatial symmetry (as in ungulate allogrooming), the importance of synergistic benefits (as in nuptial gifts), the facilitating role of collective actions (as in many instances of human cooperation), and the potential role of kinship (as in primate grooming).

Partner choice creates fairness in humans.

Many studies demonstrate that partner choice has played an important role in the evolution of human cooperation, but little work has tested its impact on the evolution of human fairness. In experiments involving divisions of money, people become either over-generous or over-selfish when they are in competition to be chosen as cooperative partners. Hence, it is difficult to see how partner choice could result in the evolution of fair, equal divisions. Here, we show that this puzzle can be solved if we consider the outside options on which partner choice operates. We conduct a behavioural experiment, run agent-based simulations and analyse a game-theoretic model to understand how outside options affect partner choice and fairness. All support the conclusion that partner choice leads to fairness only when individuals have equal outside options. We discuss how this condition has been met in our evolutionary history, and the implications of these findings for our understanding of other aspects of fairness less specific than preferences for equal divisions of resources.

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.

A mutualistic approach to morality: the evolution of fairness by partner choice.

What makes humans moral beings? This question can be understood either as a proximate "how" question or as an ultimate "why" question. The "how" question is about the mental and social mechanisms that produce moral judgments and interactions, and has been investigated by psychologists and social scientists. The "why" question is about the fitness consequences that explain why humans have morality, and has been discussed by evolutionary biologists in the context of the evolution of cooperation. Our goal here is to contribute to a fruitful articulation of such proximate and ultimate explanations of human morality. We develop an approach to morality as an adaptation to an environment in which individuals were in competition to be chosen and recruited in mutually advantageous cooperative interactions. In this environment, the best strategy is to treat others with impartiality and to share the costs and benefits of cooperation equally. Those who offer less than others will be left out of cooperation; conversely, those who offer more will be exploited by their partners. In line with this mutualistic approach, the study of a range of economic games involving property rights, collective actions, mutual help and punishment shows that participants' distributions aim at sharing the costs and benefits of interactions in an impartial way. In particular, the distribution of resources is influenced by effort and talent, and the perception of each participant's rights on the resources to be distributed.

Partner choice, fairness, and the extension of morality.

Our discussion of the commentaries begins, at the evolutionary level, with issues raised by our account of the evolution of morality in terms of partner-choice mutualism. We then turn to the cognitive level and the characterization and workings of fairness. In a final section, we discuss the degree to which our fairness-based approach to morality extends to norms that are commonly considered moral even though they are distinct from fairness.