Linking brain structure and activation in temporoparietal junction to explain the neurobiology of human altruism

Human altruism shaped our evolutionary history and pervades social and political life. There are, however, enormous individual differences in altruism. Some people are almost completely selfish, while others display strong altruism, and the factors behind this heterogeneity are only poorly understood. We examine the neuroanatomical basis of these differences with voxel-based morphometry and show that gray matter (GM) volume in the right temporoparietal junction (TPJ) is strongly associated with both individuals' altruism and the individual-specific conditions under which this brain region is recruited during altruistic decision making. Thus, individual differences in GM volume in TPJ not only translate into individual differences in the general propensity to behave altruistically, but they also create a link between brain structure and brain function by indicating the conditions under which individuals are likely to recruit this region when they face a conflict between altruistic and selfish acts.     

Selfishness and altruism can coexist when help is subject to diminishing returns

Altruism and selfishness are 30-50% heritable in man in both Western and non-Western populations. This genetically based variation in altruism and selfishness requires explanation. In non-human animals, altruism is generally directed towards relatives, and satisfies the condition known as Hamilton's rule. This nepotistic altruism evolves under natural selection only if the ratio of the benefit of receiving help to the cost of giving it exceeds a value that depends on the relatedness of the individuals involved. Standard analyses assume that the benefit provided by each individual is the same but it is plausible in some cases that as more individuals contribute, help is subject to diminishing returns. We analyse this situation using a single-locus two-allele model of selection in a diploid population with the altruistic allele dominant to the selfish allele. The analysis requires calculation of the relationship between the fitnesses of the genotypes and the frequencies of the genes. The fitnesses vary not only with the genotype of the individual but also with the distribution of phenotypes amongst the sibs of the individual and this depends on the genotypes of his parents. These calculations are not possible by direct fitness or ESS methods but are possible using population genetics. Our analysis shows that diminishing returns change the operation of natural selection and the outcome can now be a stable equilibrium between altruistic and selfish alleles rather than the elimination of one allele or the other. We thus provide a plausible genetic model of kin selection that leads to the stable coexistence in the same population of both altruistic and selfish individuals. This may explain reported genetic variation in altruism in man.     

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.     

Learning and stabilization of altruistic behaviors in multi-agent systems by reciprocity

Optimization of performance in collective systems often requires altruism. The emergence and stabilization of altruistic behaviors are difficult to achieve because the agents incur a cost when behaving altruistically. In this paper, we propose a biologically inspired strategy to learn stable altruistic behaviors in artificial multi-agent systems, namely reciprocal altruism. This strategy in conjunction with learning capabilities make altruistic agents cooperate only between themselves, thus preventing their exploitation by selfish agents, if future benefits are greater than the current cost of altruistic acts. Our multi-agent system is made up of agents with a behavior-based architecture. Agents learn the most suitable cooperative strategy for different environments by means of a reinforcement learning algorithm. Each agent receives a reinforcement signal that only measures its individual performance. Simulation results show how the multi-agent system learns stable altruistic behaviors, so achieving optimal (or near-to-optimal) performances in unknown and changing environments. Received: 1 August 1997 / Accepted in revised form: 28 November 1997     

WHEN HAWKS GIVE RISE TO DOVES: THE EVOLUTION AND TRANSITION OF ENFORCEMENT STRATEGIES

The question of how altruism can evolve despite its local disadvantage to selfishness has produced a wealth of theoretical and empirical research capturing the attention of scientists across disciplines for decades. One feature that has remained consistent through this outpouring of knowledge has been that researchers have looked to the altruists themselves for mechanisms by which altruism can curtail selfishness. An alternative perspective may be that just as altruists want to limit selfishness in the population, so may the selfish individuals themselves. These alternative perspectives have been most evident in the fairly recent development of enforcement strategies. Punishment can effectively limit selfishness in the population, but it is not free. Thus, when punishment evolves among altruists, the double costs of exploitation from cheaters and punishment make the evolution of punishment problematic. Here we show that punishment can more readily invade selfish populations when associated with selfishness, whereas altruistic punishers cannot. Thereafter, the establishment of altruism because of enforcement by selfish punishers provides the ideal invasion conditions for altruistic punishment, effectively creating a transition of punishment from selfishness to altruistic. Thus, from chaotic beginnings, a little hypocrisy may go a long way in the evolution and maintenance 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.     

Selection for altruism through random drift in variable size populations

ABSTRACT: BACKGROUND: Altruistic behavior is defined as helping others at a cost to oneself and a lowered fitness. The lower fitness implies that altruists should be selected against, which is in contradiction with their widespread presence is nature. Present models of selection for altruism (kin or multilevel) show that altruistic behaviors can have 'hidden' advantages if the 'common good' produced by altruists is restricted to some related or unrelated groups. These models are mostly deterministic, or assume a frequency dependent fitness. RESULTS: Evolutionary dynamics is a competition between deterministic selection pressure and stochastic events due to random sampling from one generation to the next. We show here that an altruistic allele extending the carrying capacity of the habitat can win by increasing the random drift of "selfish" alleles. In other terms, the fixation probability of altruistic genes can be higher than those of a selfish ones, even though altruists have a smaller fitness. Moreover when populations are geographically structured, the altruists advantage can be highly amplified and the fixation probability of selfish genes can tend toward zero. The above results are obtained both by numerical and analytical calculations. Analytical results are obtained in the limit of large populations. CONCLUSIONS: The theory we present does not involve kin or multilevel selection, but is based on the existence of random drift in variable size populations. The model is a generalization of the original Fisher-Wright and Moran models where the carrying capacity depends on the number of altruists.     

Primate Nepotism: What is the Explanatory Value of Kin Selection?

Kin selection theory (KS) is widely invoked to account for the preferential treatment of kin—nepotism—in primate societies. Because this idea is so pervasive the role of KS is often unquestioned and optional mechanisms are often ignored. I first examine the potential role of some other nepotism-generating mechanisms by concentrating on the effect of the proximity correlate of matrilineal kinship. This correlate of kinship may bias the development of mutually selfish interactions among relatives—kin-biased mutualism—and that of reciprocally altruistic interactions—kin-biased reciprocal altruism—two mechanisms that have been given little weight compared to KS and whose impact on the evolution of nepotism is therefore unknown. However, these two options to KS cannot account for the existence of unilaterally altruistic interactions among kin, which provide, therefore, the best type of evidence to test KS. But such evidence is difficult to obtain because many behaviors considered altruistic may in fact be selfish, and because kin altruism is seldom unilateral; it is most often bilateral, as expected by reciprocal altruism theory. For these reasons, one should be extremely cautious before equating nepotism exclusively with KS. Next, I examine the predictions of KS regarding the deployment of altruism according to degree of kinship by considering, in addition to the variables of Hamilton's equation, the duration of behaviors, the size of kin classes and their differential availability. In general, altruism is expected to be allocated at a fairly constant rate among kin categories and to drop markedly past the degree of relatedness beyond which altruism is no more profitable. Very little data allow one to test conclusively this prediction, as well as some other significant predictions. Overall, there is ample evidence for the role of KS in shaping mother-offspring interactions in various areas. But the evidence for kin-selected altruism beyond the mother-offspring bond (r < 0.5), though qualitatively solid, is much less abundant. Kin altruism drops markedly beyond r = 0.25 (half-siblings and grandmother-grandoffspring dyads).     

Could sexual selection have made us psychological altruists?

Psychological altruism (being motivated by the needs of others) has a tendency to produce behaviour that is costly in evolutionary terms. How, then, could the capacity for psychological altruism evolve? One suggestion is that it is the result of sexual selection. There are, however, two problems that face such an account: first, it is not clear that the resulting behaviour would be altruistic in the relevant sense, and second, it does not seem to fit with key features of our actual helping behaviour. I will argue that both of these problems can be avoided if we adopt a modular account of desire formation.     

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.     

Fairness expectations and altruistic sharing in 15-month-old human infants

Human cooperation is a key driving force behind the evolutionary success of our hominin lineage. At the proximate level, biologists and social scientists have identified other-regarding preferences--such as fairness based on egalitarian motives, and altruism--as likely candidates for fostering large-scale cooperation. A critical question concerns the ontogenetic origins of these constituents of cooperative behavior, as well as whether they emerge independently or in an interrelated fashion. The answer to this question will shed light on the interdisciplinary debate regarding the significance of such preferences for explaining how humans become such cooperative beings. We investigated 15-month-old infants' sensitivity to fairness, and their altruistic behavior, assessed via infants' reactions to a third-party resource distribution task, and via a sharing task. Our results challenge current models of the development of fairness and altruism in two ways. First, in contrast to past work suggesting that fairness and altruism may not emerge until early to mid-childhood, 15-month-old infants are sensitive to fairness and can engage in altruistic sharing. Second, infants' degree of sensitivity to fairness as a third-party observer was related to whether they shared toys altruistically or selfishly, indicating that moral evaluations and prosocial behavior are heavily interconnected from early in development. Our results present the first evidence that the roots of a basic sense of fairness and altruism can be found in infancy, and that these other-regarding preferences develop in a parallel and interwoven fashion. These findings support arguments for an evolutionary basis--most likely in dialectical manner including both biological and cultural mechanisms--of human egalitarianism given the rapidly developing nature of other-regarding preferences and their role in the evolution of human-specific forms of cooperation. Future work of this kind will help determine to what extent uniquely human sociality and morality depend on other-regarding preferences emerging early in life.     

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.     

Genomic Imprinting As a Window into Human Language Evolution

Humans spend large portions of their time and energy talking to one another, yet it remains unclear whether this activity is primarily selfish or altruistic. Here, it is shown how parent‐of‐origin specific gene expression—or “genomic imprinting”—may provide an answer to this question. First, it is shown why, regarding language, only altruistic or selfish scenarios are expected. Second, it is pointed out that an individual's maternal‐origin and paternal‐origin genes may have different evolutionary interests regarding investment into language, and that this intragenomic conflict may drive genomic imprinting which—as the direction of imprint depends upon whether investment into language is relatively selfish or altruistic—may be used to discriminate between these two possibilities. Third, predictions concerning the impact of various mutations and epimutations at imprinted loci on language pathologies are derived. In doing so, a framework is developed that highlights avenues for using intragenomic conflicts to investigate the evolutionary drivers of language.     

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.     

Altruism or solidarity? The motives for organ donation and two proposals

Proposals for increasing organ donation are often rejected as incompatible with altruistic motivation on the part of donors. This paper questions, on conceptual grounds, whether most organ donors really are altruistic. If we distinguish between altruism and solidarity--a more restricted form of other-concern, limited to members of a particular group--then most organ donors exhibit solidarity, rather than altruism. If organ donation really must be altruistic, then we have reasons to worry about the motives of existing donors. However, I argue that altruism is not necessary, because organ donation supplies important goods, whatever the motivation, and we can reject certain dubious motivations, such as financial profit, without insisting on altruism. Once solidaristic donation is accepted, certain reforms for increasing donation rates seem permissible. This paper considers two proposals. Firstly, it has been suggested that registered donors should receive priority for transplants. While this proposal appears based on a solidaristic norm of reciprocity, it is argued that such a scheme would be undesirable, since non-donors may contribute to society in other ways. The second proposal is that donors should be able to direct their organs towards recipients that they feel solidarity with. This is often held to be inconsistent with altruistic motivation, but most donation is not entirely undirected in the first place (for instance, donor organs usually go to co-nationals). While allowing directed donation would create a number of practical problems, such as preventing discrimination, there appears to be no reason in principle to reject it.     

Give unto others: genetically unrelated cotton-top tamarin monkeys preferentially give food to those who altruistically give food back

Altruistic food giving among genetically unrelated individuals is rare in nature. The few examples that exist suggest that when animals give food to unrelated others, they may do so on the basis of mutualistic or reciprocally altruistic relationships. We present the results of four experiments designed to tease apart the factors mediating food giving among genetically unrelated cotton-top tamarins (Saguinus oedipus), a cooperatively breeding New World primate. In experiment 1 we show that individuals give significantly more food to a trained conspecific who unilaterally gives food than to a conspecific who unilaterally never gives food. The apparent contingency of the tamarins' food-giving behaviour motivated the design of experiments 2-4. Results from all three experiments show that altruistic food giving is mediated by prior acts of altruistic food giving by a conspecific. Specifically, tamarins do not give food to unrelated others when the food received in the past represents the by-product of another's selfish actions (experiments 2 and 3) or when a human experimenter gives them food (experiment 4) as did the unilateral altruist in experiment 1. By contrast, if one tamarin gives another food without obtaining any immediate benefit, then the recipient is more likely to give food in return. Overall, results show that tamarins altruistically give food to genetically unrelated conspecifics, discriminate between altruistic and selfish actions, and give more food to those who give food back. Tamarins therefore have the psychological capacity for reciprocally mediated altruism.     

Multilevel selection: the evolution of cooperation in non-kin groups

Hamiltons (1964a, 1964b) landmark papers are rightly recognized as the formal basis for our understanding of the evolution of altruistic traits. However, Hamiltons equation as he originally expressed it is simplistic. A genetically oriented approach to studying multilevel selection can provide insights into how the terminology and assumptions used by Hamilton can be generalized. Using contextual analysis I demonstrated that Hamiltons rule actually embodies three distinct processes, group selection, individual selection, and transmission genetics or heritability. Whether an altruistic trait will evolve depends the balance of all of these factors. The genetical approach, and particularly, contextual analysis provides a means of separating these factors and examining them one at a time. Perhaps the greatest issue with Hamiltons equation is the interpretation of r. Hamilton (1964a) interpreted this as relatedness. In this paper I show that what Hamilton called relatedness is more generally interpreted as the proportion for variance among groups, and that many processes in addition to relatedness can increase the variance among groups. I also show that the evolution of an altruistic trait is driven by the ratio of the heritability at the group level to the heritability at the individual level. Under some circumstances this ratio can be greater than 1. In this situation altruism can evolve even if selection favoring selfish behavior is stronger than selection favoring altruism.     

The adaptive dynamics of altruism in spatially heterogeneous populations

Abstract.— We study the spatial adaptive dynamics of a continuous trait that measures individual investment in altruism. Our study is based on an ecological model of a spatially heterogeneous population from which we derive an appropriate measure of fitness. The analysis of this fitness measure uncovers three different selective processes controlling the evolution of altruism: the direct physiological cost, the indirect genetic benefits of cooperative interactions, and the indirect genetic costs of competition for space. In our model, habitat structure and a continuous life cycle makes the cost of competing for space with relatives negligible. Our study yields a classification of adaptive patterns of altruism according to the shape of the costs of altruism (with decelerating, linear, or accelerating dependence on the investment in altruism). The invasion of altruism occurs readily in species with accelerating costs, but large mutations are critical for altruism to evolve in selfish species with decelerating costs. Strict selfishness is maintained by natural selection only under very restricted conditions. In species with rapidly accelerating costs, adaptation leads to an evolutionarily stable rate of investment in altruism that decreases smoothly with the level of mobility. A rather different adaptive pattern emerges in species with slowly accelerating costs: high altruism evolves at low mobility, whereas a quasi-selfish state is promoted in more mobile species. The high adaptive level of altruism can be predicted solely from habitat connectedness and physiological parameters that characterize the pattern of cost. We also show that environmental changes that cause increased mobility in those highly altruistic species can beget selection-driven self-extinction, which may contribute to the rarity of social species.     

Competitive altruism: from reciprocity to the handicap principle

Current work on cooperation is focused on the theory of reciprocal altruism. However, reciprocity is just one way of getting a return on an investment in altruism and is difficult to apply to many examples. Reciprocity theory addresses how animals respond dynamically to others so as to cooperate without being exploited. I discuss how introducing differences in individual generosity together with partner choice into models of reciprocity can lead to an escalation in altruistic behaviour. Individuals may compete for the most altruistic partners and non-altruists may become ostracized. I refer to this phenomenon as competitive altruism and propose that it can represent a move away from the dynamic responsiveness of reciprocity. Altruism may be rewarded in kind, but rewards may be indirectly accrued or may not involve the return of altruism at all, for example if altruists tend to be chosen as mates. This variety makes the idea of competitive altruism relevant to behaviours which cannot be explained by reciprocity. I consider whether altruism might act as a signal of quality, as proposed by the handicap principle. I suggest that altruistic acts could make particularly effective signals because of the inherent benefits to receivers. I consider how reciprocity and competitive altruism are related and how they may be distinguished.