Self-protection promotes altruism

Self-protection tendencies allowed our human ancestors to survive and thrive. In three experiments, we find that individuals who have a salient self-protection motive are more altruistic to others, such as by helping them out or offering them more money in the dictator game paradigm. Self-protecting individuals desire to “bind together” as there is “safety in numbers”, and being altruistic to others should be one (but not the only) way to achieve this goal. Consistent with this reasoning, we find across three behavioral experiments using both non-monetary (Experiment 1) and monetary altruistic contexts (Experiments 2–3) that self-protecting individuals are more altruistic when the altruism is not anonymous (Experiment 1) and when they have the reasonable expectation of future interaction with the recipient (Experiment 2), both of which are situations that should increase affiliation. The effect attenuates when altruism does not help self-protecting individuals, such as when money is donated to impersonal organizations rather than individuals (Experiment 3). We finally discuss the theoretical contributions as well as limitations of our work.     

Multilevel selection and human altruism

Views on the evolution of altruism based upon multilevel selection on structured populations pay little attention to the difference between fortuitous and deliberate processes leading to assortative grouping. Altruism may evolve when assortative grouping is fortuitously produced by forces external to the organism. But when it is deliberately produced by the same proximate mechanism that controls altruistic responses, as in humans, exploitation of altruists by selfish individuals is unlikely and altruism evolves as an individually advantageous trait. Groups formed with altruists of this sort are special, because they are not affected by subversion from within. A synergistic process where altruism is selected both at the individual and at the group level can take place.     

Relatedness and altruism in Polistes wasps

Genetic relatedness is expected to play a crucial role in theevolution of altruistic behaviors such as worker behavior inthe social insects. If individuals sacrifice their own reproduction,then the genes for this sacrifice will be lost unless theseindividuals aid the reproduction of others who share the genes.This leads to the prediction that altruism should be most commonin species with high relatedness among potential beneficiaries.Here we report an attempt to test for such an association. Weestimated both the incidence of altruism and the relatednessto potential beneficiaries in foundresses of seven species ofpaper wasps. The predicted positive correlation was not found,and we conclude that factors other than relatedness are moreimportant in determining interspecific differences in the incidenceof altruism.     

Heterogeneity stabilizes reciprocal altruism interactions

In considering the phenomena of reciprocal altruism few would dispute that there are differences in individual quality-in particular, that for some individuals, at least on occasion, the cost of doing favors will exceed the potential of future benefits. That is, at any given time, a typical population is heterogeneous with respect to the affordability of reciprocal altruism. However, methodological limitations of the traditional analytical framework-Single Type (symmetric) Evolutionary Game Theory-have restricted previous analytical efforts to addressing populations idealized in terms of their averages. Here we use the methods of Multitype Evolutionary Game Theory to analyse the role of individual differences in direct reciprocity interactions. Multitype analysis shows that non-idealized populations possess an ESS profile wherein individuals who cannot afford reciprocity (low-quality) defect, while individuals who derive net benefits from reciprocity (high-quality) cooperate. Furthermore, this cooperation is implemented via unmodified tit-for-tat (TfT) strategy. Hence, our results may help resolve a long-standing problem concerning the evolutionary stability of TfT in direct reciprocal altruism. Finally, this difference between idealized and real populations is not restricted to direct reciprocal cooperation. Previously (Lotem et al., 1999) we have demonstrated evolutionarily stable indirect reciprocal cooperation among high-quality individuals in heterogeneous populations.     

Kin recognition and the evolution of altruism

In 1964, Hamilton formalized the idea of kin selection to explain the evolution of altruistic behaviours. Since then, numerous examples from a diverse array of taxa have shown that seemingly altruistic actions towards close relatives are a common phenomenon. Although many species use kin recognition to direct altruistic behaviours preferentially towards relatives, this important aspect of social biology is less well understood theoretically. I extend Hamilton's classic work by defining the conditions for the evolution of kin-directed altruism when recognizers are permitted to make acceptance (type I) and rejection (type II) errors in the identification of social partners with respect to kinship. The effect of errors in recognition on the evolution of kin-directed altruism depends on whether the population initially consists of unconditional altruists or non-altruists (i.e. alternative forms of non-recognizers). Factors affecting the level of these error rates themselves, their evolution and their long-term stability are discussed.     

Population viscosity and the evolution of altruism

The term population viscosity refers to limited dispersal, which increases the genetic relatedness of neighbors. This effect both supports the evolution of altruism by focusing the altruists' gifts on relatives of the altruist, and also limits the extent to which altruism may emerge by exposing clusters of altruists to stiffer local competition. Previous analyses have emphasized the way in which these two effects can cancel, limiting the viability of altruism. These papers were based on models in which total population density was held fixed. We present here a class of models in which population density is permitted to fluctuate, so that patches of altruists are supported at a higher density than patches of non-altruists. Under these conditions, population viscosity can support the selection of both weak and strong altruism.     

A behavioral analysis of altruism

Altruistic acts have been defined, in economic terms, as “…costly acts that confer economic benefits on other individuals” (Fehr and Fischbacher, 2003). In multi-player, one-shot prisoner's dilemma games, a significant number of players behave altruistically; their behavior benefits each of the other players but is costly to them. We consider three potential explanations for such altruism. The first explanation, following a suggestion by the philosopher Derek Parfit, assumes that players devise a strategy to avoid being free-loaders—and that in the present case this strategy dictates cooperation. The second explanation says that cooperators reject the one-shot aspect of the game and behave so as to maximize reward over a series of choices extending beyond the present situation (even though reward is not maximized in the present case). This explanation assumes that people may learn to extend the boundaries of their selves socially (beyond their own skin) as well as temporally (beyond the present moment). We propose a learning mechanism for such behavior analogous to the biological, evolutionary mechanism of group selection. The third explanation assumes that people's altruism is based on a straightforward balancing of undiscounted costs to themselves against discounted benefits to others (social discounting). The three proposed explanations of altruism complement each other.     

Multilevel selection, cooperation, and altruism

Unto Others (Sober and Wilson 1998) shows how the general principles of Multi-Level Selection (MLS) theory apply to selection at multiple levels of the biological hierarchy. It also argues for the existence of "genuine" evolutionary and psychological altruism. The authors’ views on altruism do not follow logically from principles of MLS, and their failure do disentangle these two themes undermines their otherwise excellent presentation of MLS theory. Rebuttal of the view that human nature is completely selfish depends not on the prevalence of altruism but on the importance of group-advantageous traits that benefit both self and other group members without necessarily inflicting direct costs on outsiders.     

Evolution,altruism, and the prisoner's dilemma

I first argue against Peter Singer's exciting thesis that the Prisoner's Dilemma explains why there could be an evolutionary advantage in making reciprocal exchanges that are ultimately motivated by genuine altruism over making such exchanges on the basis of enlightened long-term self-interest. I then show that an alternative to Singer's thesis — one that is also meant to corroborate the view that natural selection favors genuine altruism, recently defended by Gregory Kavka, fails as well. Finally, I show that even granting Singer's and Kavka's claim about the selective advantage of altruism proper, it is doubtful whether that type of claim can be used in a particular sort of sociobiological argument against psychological egoism.     

Nominal kinship cues facilitate altruism.

We investigated whether names in common promote altruistic behaviour, predicting that this would be especially so for relatively uncommon names, for surnames (which are better kinship cues than first names), and among women (who, although less willing than men to help strangers, according to prior research, are also the primary "kin keepers"). We solicited help from 2960 email addressees, with the request ostensibly coming from a same-sex person sharing both, either, or neither of the addressee''s first and last names. As anticipated, addressees were most likely to respond helpfully when senders shared both their names (12.3%) and least likely when they shared neither (2.0%), and this was especially true for relatively uncommon names. A shared surname was more effective than a shared first name only if it was relatively uncommon. Women were substantially more likely to reply than men. These results indicate that names elicit altruism because they function as salient cues of kinship.     

Questioning the cultural evolution of altruism

The evolutionary foundations of helping among nonkin in humans have been the object of intense debates in the past decades. One thesis has had a prominent influence in this debate: the suggestion that genuine altruism, strictly defined as a form of help that comes at a net fitness cost for the benefactor, might have evolved owing to cultural transmission. The gene–culture coevolution literature is wont to claim that cultural evolution changes the selective pressures that normally act to limit the emergence of altruistic behaviours. This paper aims to recall, however, that cultural transmission yields altruism only to the extent that it relies on maladaptive mechanisms, such as conformist imitation and (in some cases) payoff‐biased transmission. This point is sometimes obscured in the literature by a confusion between genuine altruism, maladaptive by definition, and mutualistic forms of cooperation, that benefit all parties in the long run. Theories of cultural altruism do not lift the selective pressures weighing on strictly altruistic actions; they merely shift the burden of maladaptation from social cognition to cultural transmission.     

Hidden altruism in a real-world setting

Concerns for reputation can promote cooperative behaviour. Individuals that behave cooperatively stand to benefit if they gain in influence, status or are more likely to be chosen as interaction partners by others. Most theoretical and empirical models of cooperation predict that image score will increase with cooperative contributions. Individuals are therefore expected to make higher contributions when observed by others and should opt to make contributions publicly rather than privately, particularly when contributions are higher than average. Here, however, I find the opposite effect. Using data from an online fundraising website, I show that donors are more likely to opt for anonymity when making extremely low and extremely high donations. Mid-range donations, on the other hand, are typically publicized. Recent work has shown that extremely generous individuals may be ostracized or punished by group members. The data presented here suggest that individuals may hide high donations to avoid these repercussions.     

Models of the evolution of altruism

There are two ways of calculating the spread of a gene for altruism. One, originally proposed by Hamilton, is to allow for the effects of the gene on the survival and reproduction of collateral relatives of the individual carrying it (i.e., “inclusive fitness”); this leads to the condition k > 1/r for the spread of the gene, where k is a benefit/cost ratio. The other is to count only the direct offspring of a carrier, but to allow for the altruistic acts performed toward the carrier by its relatives (“neighbour modulated fitness” or “personal fitness”). A recent personal fitness model (L. L. Cavalli Sforza and M. W. Feldman, 1978, Theor. Pop. Biol.14, 268–280) analyses parent-offspring and sib-sib altruism and concludes that k > 1/r is applicable only when fitness components are combined additively. The present paper analyses some simple models in which the phenotypic effects are carefully specified. It is concluded that it is sometimes, but not always, appropriate to combine fitness components additively. The relative roles of inclusive and personal fitness models are compared. The former have the virtue of being easier to think about in causal terms; and the latter of incorporating the evolution of altruism into the corpus of population genetics as an example of frequency-dependent selection.