On the relationship between evolutionary and psychological definitions of altruism and selfishness

I examine the relationship between evolutionary definitions of altruism that are based on fitness effects and psychological definitions that are based on the motives of the actor. I show that evolutionary altruism can be motivated by proximate mechanisms that are psychologically either altruistic or selfish. I also show that evolutionary definitions do rely upon motives as a metaphor in which the outcome of natural selection is compared to the decisions of a psychologically selfish (or altruistic) individual. Ignoring the precise nature of both psychological and evolutionary definitions has obscured many important issues, including the biological roots of psychological altruism.     

Altruism and fairness: Unnatural selection?

Darwin admitted that the evolution of moral phenomena such as altruism and fairness, which are usually in opposition to the maximization of individual reproductive success, was not easily accounted for by natural selection. Later, authors have proposed additional mechanisms, including kin selection, inclusive fitness, and reciprocal altruism. In the present work, we explore the extent to which sexual selection has played a role in the appearance of human moral traits. It has been suggested that because certain moral virtues, including altruism and kindness, are sexually attractive, their evolution could have been shaped by the process of sexual selection. Our review suggests that although it is possible that sexual selection played such a role, it is difficult to determine the extent of its relevance, the specific form of this influence, and its interplay with other evolutionary mechanisms.     

Levels of selection, altruism, and primate behavior.

Altruistic behaviors seem anomalous from a traditional view of Darwinian natural selection, and evolutionary explanations for them have generated much discussion. The debate centers around four major explanations: classic individual-level selection, reciprocity and game theory, kin selection, and trait-group selection. The historical context and defining criteria of each model must be reviewed before its validity can be assessed. Of these proposed mechanisms, group selection historically has been the most controversial. Although the extent to which empirical data support group selection hypotheses is uncertain, there is evidence for group-level selection among avirulent virus strains and foraging ant queens. Researchers studying mammalian behavior, particularly primatologists, have largely dismissed models of group-level selection. Most discussion of altruism among primates has focused on differences in fitness among individuals within a single group, but students of altruistic behaviors exhibited by primates also need to investigate intergroup variation with respect to these behaviors. Various altruistic behaviors are likely to have evolved through different forms of selection, and each example of apparent altruism therefore needs to be evaluated separately.     

The handicap principle and the argument of subversion from within

This paper examines the very disparate positions that various actors have taken towards the argument of subversion from within (a classical argument against the evolution of altruism by group selection) in a set of related debates on group selection, altruism and the handicap principle. Using this set of debates as a case study, this paper argues that different applications of epistemic values were one of the factors behind the disagreements between John Maynard Smith and Amotz Zahavi over a number of important evolutionary issues. The paper also argues that these different applications were connected to important epistemological differences related in part (but not solely) to their disciplinary background. Apart from conflicting evolutionary views concerning the theoretical feasibility of the handicap effect, these antagonists both differed in the confidence they ascribed to mathematical modeling and over the hereditary basis for altruistic behavior.     

The evolution of cooperation and altruism--a general framework and a classification of models

One of the enduring puzzles in biology and the social sciences is the origin and persistence of intraspecific cooperation and altruism in humans and other species. Hundreds of theoretical models have been proposed and there is much confusion about the relationship between these models. To clarify the situation, we developed a synthetic conceptual framework that delineates the conditions necessary for the evolution of altruism and cooperation. We show that at least one of the four following conditions needs to be fulfilled: direct benefits to the focal individual performing a cooperative act; direct or indirect information allowing a better than random guess about whether a given individual will behave cooperatively in repeated reciprocal interactions; preferential interactions between related individuals; and genetic correlation between genes coding for altruism and phenotypic traits that can be identified. When one or more of these conditions are met, altruism or cooperation can evolve if the cost-to-benefit ratio of altruistic and cooperative acts is greater than a threshold value. The cost-to-benefit ratio can be altered by coercion, punishment and policing which therefore act as mechanisms facilitating the evolution of altruism and cooperation. All the models proposed so far are explicitly or implicitly built on these general principles, allowing us to classify them into four general categories.     

Bioaltruism reconsidered

Altruistic behavior is often regarded as sociobiology's most central theoretical problem, but is it? Altruism in biology, bioaltruism, has many meanings, which can be grouped into two categories. The first I will callcommon bioaltruism. It is primarily of ethological relevance. The second,evolutionary bioaltruism, is a special category in evolutionary respects in that it may indeed pose a problem for evolutionary theory. These categories are logically independent. Moreover, both of them are logically different from altruism in its everyday psychological or moral sense. Sociobiological examples of bioaltruistic behavior concern bioaltruism in the first sense only, so the theoretical problem ‘altruism’ is supposed to pose, is indeed nothing but a theoretical problem and the bioaltruism that actually occurs has no evolutionary relevance. Nevertheless, evolutionary theory is relevant to our understanding of the possibility of common bioaltruism, and that possibility — even though bioaltruism is conceptually different from ethical altruism — is relevant for ethicists: it sheds light on what we can ask people to do or not to do.     

Group Selection as Behavioral Adaptation to Systematic Risk

Despite many compelling applications in economics, sociobiology, and evolutionary psychology, group selection is still one of the most hotly contested ideas in evolutionary biology. Here we propose a simple evolutionary model of behavior and show that what appears to be group selection may, in fact, simply be the consequence of natural selection occurring in stochastic environments with reproductive risks that are correlated across individuals. Those individuals with highly correlated risks will appear to form “groups”, even if their actions are, in fact, totally autonomous, mindless, and, prior to selection, uniformly randomly distributed in the population. This framework implies that a separate theory of group selection is not strictly necessary to explain observed phenomena such as altruism and cooperation. At the same time, it shows that the notion of group selection does captures a unique aspect of evolution—selection with correlated reproductive risk–that may be sufficiently widespread to warrant a separate term for the phenomenon.     

The evolution of altruism: game theory in multilevel selection and inclusive fitness

Although the prisoner's dilemma (PD) has been used extensively to study reciprocal altruism, here we show that the n-player prisoner's dilemma (NPD) is also central to two other prominent theories of the evolution of altruism: inclusive fitness and multilevel selection. An NPD model captures the essential factors for the evolution of altruism directly in its parameters and integrates important aspects of these two theories such as Hamilton's rule, Simpson's paradox, and the Price covariance equation. The model also suggests a simple interpretation of the Price selection decomposition and an alternative decomposition that is symmetrical and complementary to it. In some situations this alternative shows the temporal changes in within- and between-group selection more clearly than the Price equation. In addition, we provide a new perspective on strong vs. weak altruism by identifying their different underlying game structures (based on absolute fitness) and showing how their evolutionary dynamics are nevertheless similar under selection (based on relative fitness). In contrast to conventional wisdom, the model shows that both strong and weak altruism can evolve in periodically formed random groups of non-conditional strategies if groups are multigenerational. An integrative approach based on the NPD helps unify different perspectives on the evolution of altruism.     

Altruism Can Proliferate through Population Viscosity despite High Random Gene Flow

The ways in which natural selection can allow the proliferation of cooperative behavior have long been seen as a central problem in evolutionary biology. Most of the literature has focused on interactions between pairs of individuals and on linear public goods games. This emphasis has led to the conclusion that even modest levels of migration would pose a serious problem to the spread of altruism through population viscosity in group structured populations. Here we challenge this conclusion, by analyzing evolution in a framework which allows for complex group interactions and random migration among groups. We conclude that contingent forms of strong altruism that benefits equally all group members, regardless of kinship and without greenbeard effects, can spread when rare under realistic group sizes and levels of migration, due to the assortment of genes resulting only from population viscosity. Our analysis combines group-centric and gene-centric perspectives, allows for arbitrary strength of selection, and leads to extensions of Hamilton’s rule for the spread of altruistic alleles, applicable under broad conditions.     

Hamilton vs. Kant: pitting adaptations for altruism against adaptations for moral judgment

Prominent evolutionary theories of morality maintain that the adaptations that underlie moral judgment and behavior function, at least in part, to deliver benefits (or prevent harm) to others. These explanations are based on the theories of kin selection and reciprocal altruism, and they predict that moral systems are designed to maximize Hamiltonian inclusive fitness. In sharp contrast, however, moral judgment often appears Kantian and rule-based. To reconcile this apparent discrepancy, some theorists have claimed that Kantian moral rules result from mechanisms that implement simple heuristics for maximizing welfare. To test this idea, we conducted a set of studies in which subjects (N=1290) decided whether they would kill one person to save five others, varying the relationship of the subject with the others involved (strangers, friends, brothers). Are participants more likely to observe the Kantian rule against killing in decisions about brothers and friends, rather than strangers? We found the reverse. Subjects reported greater willingness to kill a brother or friend than a stranger (in order to save five others of the same type). These results suggest that the rule-based structure of moral cognition is not explained by kin selection, reciprocity, or other altruism theories.     

Effects of information and group structure on evolution of altruism: analysis of two-score model by covariance and contextual analyses

An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.     

The group selection controversy

Many thought Darwinian natural selection could not explain altruism. This error led Wynne‐Edwards to explain sustainable exploitation in animals by selection against overexploiting groups. Williams riposted that selection among groups rarely overrides within‐group selection. Hamilton showed that altruism can evolve through kin selection. How strongly does group selection influence evolution? Following Price, Hamilton showed how levels of selection interact: group selection prevails if Hamilton’s rule applies. Several showed that group selection drove some major evolutionary transitions. Following Hamilton’s lead, Queller extended Hamilton’s rule, replacing genealogical relatedness by the regression on an actor’s genotypic altruism of interacting neighbours’ phenotypic altruism. Price’s theorem shows the generality of Hamilton’s rule. All instances of group selection can be viewed as increasing inclusive fitness of autosomal genomes. Nonetheless, to grasp fully how cooperation and altruism evolve, most biologists need more concrete concepts like kin selection, group selection and selection among individuals for their common good.     

Selective pressures for accurate altruism targeting: evidence from digital evolution for difficult-to-test aspects of inclusive fitness theory

Inclusive fitness theory predicts that natural selection will favour altruist genes that are more accurate in targeting altruism only to copies of themselves. In this paper, we provide evidence from digital evolution in support of this prediction by competing multiple altruist-targeting mechanisms that vary in their accuracy in determining whether a potential target for altruism carries a copy of the altruist gene. We compete altruism-targeting mechanisms based on (i) kinship (kin targeting), (ii) genetic similarity at a level greater than that expected of kin (similarity targeting), and (iii) perfect knowledge of the presence of an altruist gene (green beard targeting). Natural selection always favoured the most accurate targeting mechanism available. Our investigations also revealed that evolution did not increase the altruism level when all green beard altruists used the same phenotypic marker. The green beard altruism levels stably increased only when mutations that changed the altruism level also changed the marker (e.g. beard colour), such that beard colour reliably indicated the altruism level. For kin- and similarity-targeting mechanisms, we found that evolution was able to stably adjust altruism levels. Our results confirm that natural selection favours altruist genes that are increasingly accurate in targeting altruism to only their copies. Our work also emphasizes that the concept of targeting accuracy must include both the presence of an altruist gene and the level of altruism it produces.     

Altruism may arise from individual selection

The fact that humans cooperate with non-kin in large groups, or with people they will never meet again, is a long-standing evolutionary puzzle. Altruism, the capacity to perform costly acts that confer benefits on others, is at the core of cooperative behavior. Behavioral experiments show that humans have a predisposition to cooperate with others and to punish non-cooperators at personal cost (so-called strong reciprocity) which, according to standard evolutionary game theory arguments, cannot arise from selection acting on individuals. This has led to the suggestion of group and cultural selection as the only mechanisms that can explain the evolutionary origin of human altruism. We introduce an agent-based model inspired on the Ultimatum Game, that allows us to go beyond the limitations of standard evolutionary game theory and show that individual selection can indeed give rise to strong reciprocity. Our results are consistent with the existence of neural correlates of fairness and in good agreement with observations on humans and monkeys.     

Can natural selection favour altruism between species?

Darwin suggested that the discovery of altruism between species would annihilate his theory of natural selection. However, it has not been formally shown whether between‐species altruism can evolve by natural selection, or why this could never happen. Here, we develop a spatial population genetic model of two interacting species, showing that indiscriminate between species helping can be favoured by natural selection. We then ask if this helping behaviour constitutes altruism between species, using a linear‐regression analysis to separate the total action of natural selection into its direct and indirect (kin selected) components. We show that our model can be interpreted in two ways, as either altruism within species, or altruism between species. This ambiguity arises depending on whether or not we treat genes in the other species as predictors of an individual's fitness, which is equivalent to treating these individuals as agents (actors or recipients). Our formal analysis, which focuses upon evolutionary dynamics rather than agents and their agendas, cannot resolve which is the better approach. Nonetheless, because a within‐species altruism interpretation is always possible, our analysis supports Darwin's suggestion that natural selection does not favour traits that provide benefits exclusively to individuals of other species.     

The robustness of the weak selection approximation for the evolution of altruism against strong selection

The weak selection approximation of population genetics has made possible the analysis of social evolution under a considerable variety of biological scenarios. Despite its extensive usage, the accuracy of weak selection in predicting the emergence of altruism under limited dispersal when selection intensity increases remains unclear. Here, we derive the condition for the spread of an altruistic mutant in the infinite island model of dispersal under a Moran reproductive process and arbitrary strength of selection. The simplicity of the model allows us to compare weak and strong selection regimes analytically. Our results demonstrate that the weak selection approximation is robust to moderate increases in selection intensity and therefore provides a good approximation to understand the invasion of altruism in spatially structured population. In particular, we find that the weak selection approximation is excellent even if selection is very strong, when either migration is much stronger than selection or when patches are large. Importantly, we emphasize that the weak selection approximation provides the ideal condition for the invasion of altruism, and increasing selection intensity will impede the emergence of altruism. We discuss that this should also hold for more complicated life cycles and for culturally transmitted altruism. Using the weak selection approximation is therefore unlikely to miss out on any demographic scenario that lead to the evolution of altruism under limited dispersal.     

Cultural transmission and the evolution of human behaviour: a general approach based on the Price equation

Transmitted culture can be viewed as an inheritance system somewhat independent of genes that is subject to processes of descent with modification in its own right. Although many authors have conceptualized cultural change as a Darwinian process, there is no generally agreed formal framework for defining key concepts such as natural selection, fitness, relatedness and altruism for the cultural case. Here, we present and explore such a framework using the Price equation. Assuming an isolated, independently measurable culturally transmitted trait, we show that cultural natural selection maximizes cultural fitness, a distinct quantity from genetic fitness, and also that cultural relatedness and cultural altruism are not reducible to or necessarily related to their genetic counterparts. We show that antagonistic coevolution will occur between genes and culture whenever cultural fitness is not perfectly aligned with genetic fitness, as genetic selection will shape psychological mechanisms to avoid susceptibility to cultural traits that bear a genetic fitness cost. We discuss the difficulties with conceptualizing cultural change using the framework of evolutionary theory, the degree to which cultural evolution is autonomous from genetic evolution, and the extent to which cultural change should be seen as a Darwinian process. We argue that the nonselection components of evolutionary change are much more important for culture than for genes, and that this and other important differences from the genetic case mean that different approaches and emphases are needed for cultural than genetic processes.     

Altruists Proliferate Even at a Selective Disadvantage within Their Own Niche

The evolutionary origin of altruism is a long-standing puzzle. Numerous explanations have been proposed, most prominently based on inclusive fitness or group selection. One possibility that has not yet been considered is that new niches will be created disproportionately often when altruism appears, perhaps by chance, causing altruists to be over-represented in such new niches. This effect is a novel variant of group selection in which altruistic groups benefit by discovering unoccupied niches instead of by competing for the limited resources within a single niche. Both an analytical population genetics model and computational simulations support that altruism systematically arises due to this side effect of increased carrying capacity even when it is strongly selected against within any given niche. In fact, even when selection is very strongly negative and altruism does not develop in most populations, it can still be expected to be observed in a consistent fraction of species. The ecological structure provided by niches thereby may be sufficient for altruists to proliferate even if they are always at a disadvantage within each niche considered individually.     

Origins of altruism diversity I: the diverse ecological roles of altruistic strategies and their evolutionary responses to local competition

Nature abounds with a rich variety of altruistic strategies, including public resource enhancement, resource provisioning, communal foraging, alarm calling, and nest defense. Yet, despite their vastly different ecological roles, current theory typically treats diverse altruistic traits as being favored under the same general conditions. Here, we introduce greater ecological realism into social evolution theory and find evidence of at least four distinct modes of altruism. Contrary to existing theory, we find that altruistic traits contributing to "resource-enhancement" (e.g., siderophore production, provisioning, agriculture) and "resource-efficiency" (e.g., pack hunting, communication) are most strongly favored when there is strong local competition. These resource-based modes of helping are "K-strategies" that increase a social group's growth yield, and should characterize species with scarce resources and/or high local crowding caused by low mortality, high fecundity, and/or mortality occurring late in the process of resource-acquisition. The opposite conditions, namely weak local competition (abundant resource, low crowding), favor survival (e.g., nest defense) and fecundity (e.g., nurse workers) altruism, which are "r-strategies" that increase a social group's growth rate. We find that survival altruism is uniquely favored by a novel evolutionary force that we call "sunk cost selection." Sunk cost selection favors helping that prevents resources from being wasted on individuals destined to die before reproduction. Our results contribute to explaining the observed natural diversity of altruistic strategies, reveal the necessary connection between the evolution and the ecology of sociality, and correct the widespread but inaccurate view that local competition uniformly impedes the evolution of altruism.     

Sober &; Wilson’s evolutionary arguments for psychological altruism: a reassessment

In their book Unto Others, Sober and Wilson argue that various evolutionary considerations (based on the logic of natural selection) lend support to the truth of psychological altruism. However, recently, Stephen Stich has raised a number of challenges to their reasoning: in particular, he claims that three out of the four evolutionary arguments they give are internally unconvincing, and that the one that is initially plausible fails to take into account recent findings from cognitive science and thus leaves open a number of egoistic responses. These challenges make it necessary to reassess the plausibility of Sober & Wilson’s evolutionary account—which is what I aim to do in this paper. In particular, I try to show that, as a matter of fact, Sober & Wilson’s case remains compelling, as some of Stich’s concerns rest on a confusion, and those that do not are not sufficiently strong to establish all the conclusions he is after. The upshot is that no reason has been given to abandon the view that evolutionary theory has advanced the debate surrounding psychological altruism.