Altruism: a research program

Phoebe: I just found a selfless good deed; I went to the park and let a bee sting me.
Joey: How is that a good deed?
Phoebe: Because now the bee gets to look tough in front of his bee friends. The bee is happy and I am not.
Joey: Now you know the bee probably died when he stung you?
Phoebe: Dammit!
- [From Friends, episode 101]

Altruism is a lively research topic. The evolutionary foundations, neural substrates, psychological mechanisms, behavioral manifestations, formal modeling and philosophical analyses of cooperation constitute a coherent—although not unified—field of inquiry. See for instance how neuroscience, game theory, economic, philosophy, psychology and evolutionary theory interact in Penner et al. 2005; Hauser 2006; Fehr and Fischbacher 2002; Fehr and Fischbacher 2003. The nature of prosocial behavior, from kin selection to animal cooperation to human morality can be considered as a progressive Lakatosian research programs. Altruism has a great conceptual "sex-appeal" because it is mystery for two types of theoreticians: biologists and economists. They both wonder why an animal or an economic agent would help another: since these agents maximize fitness/utility, altruistic behavior is suboptimal. Altruims (help, trust, fairness, etc.) seems intuitively incoherent with economic rationality and biological adaptation, with markets and natural selection. Or is it?

In the 60's, biologists challenged the idea that natural selection is incompatible with altruism. Hamilton (1964a, 1964b) and Trivers (1971) showed that biological altruism makes sense. An animal X might behave altruistically toward another Y because they are genetically related: in doing so, X maximize the copying of its gene, since many of its genes will be hosted in Y. Thus the more X and Y are genetically related, the more X will be ready to help Y. This is kin altruism. Altruism can also be reciprocal: scratch my back and I'll scratch yours. Tit-for-tat, or reciprocal altruism also makes sense because by being altruistic, one may augments its payoff. X helps Y, but the next time Y will help X; thus it is better to help than not to help. In both cases, the idea is that altruism is a mean not an end. Others argue that more complex types of altruisms exists. For instance, X can help Y because Y already helped Z (indirect reciprocity). In this case, the tit-for-tat logic is extended to agents that the helper did not meet in the past. Generalized reciprocity (see this previous post) is another type of altruism: helping someone because someone helped you in the past. This altruism does not require memory or personal identification. X helps someone because someone else helped X. Finally, Strong reciprocity is the idea that humans display genuine altruism: strong reciprocators cooperate with cooperators, do not cooperate with cheaters, and are ready to punish cheaters even at a cost to themselves. Their proponents argue that it evolved through group selection.

Experimental economics and neuroeconomics also challenged the idea of rational, greedy, selfish actor (the Ayn Rand hero). Experimental game theory showed that, contrarily to orthodox game theory, subjects cooperate massively in prisoner’s dilemma (Ledyard, 1995; Sally, 1995). Rilling et al. showed that players enjoy cooperating. Players who initiate and players who experience mutual cooperation display activation in nucleus accumbens and other reward-related areas such as the caudate nucleus, ventromedial frontal/orbitofrontal cortex, and rostral anterior cingulate cortex (Rilling et al., 2002). In another experiment, the presentation of faces of intentional cooperators caused increased activity in reward-related areas (Singer et al. 2004). In the ultimatum game, proposers make ‘fair’ offers, about 50% of the amount, responders tend to accept these offers and reject most of the ‘unfair’ offers (less than 20%;Oosterbeek et al., 2004). Brain scans of people playing the ultimatum game indicate that unfair offers trigger, in the responders’ brain, a ‘moral disgust’: the anterior insula (associated with negative emotional states like disgust or anger) is more active when unfair offers are proposed (Sanfey, Rilling, Aronson, Nystrom, & Cohen, 2003). Subjects experiment this affective reaction to unfairness only when the proposer is a human being: the activation is significantly lower when the proposer is a computer. Moreover, the anterior insula activation is proportional to the degree of unfairness and correlated with the decision to reject unfair offers (Sanfey et al., 2003: 1756). Fehr and Fischbacher (2002) suggested that economic agents are inequity-averse and have prosocial preferences. Thus they modified the utility functions to account for behavioral (and now neural) data. In Moral Markets: The Critical Role of Values in the Economy, Paul Zak proposes a radically different conception of morality in economics:

The research reported in this book revealed that most economic exchange, whether with a stranger or a known individual, relies on character values such as honesty, trust, reliability, and fairness. Such values, we argue, arise in the normal course of human interactions, without overt enforcement—lawyers, judges or the
police are present in a paucity of economic transactions (...). Markets are moral in two senses. Moral behavior is necessary for exchange in moderately regulated markets, for example, to reduce cheating without exorbitant transactions costs. In addition, market exchange itself can lead to an understanding of fair-play that can build social capital in nonmarket settings. (Zak, forthcoming)

See how this claim is similar to :

The two fundamental principles of evolution are mutation and natural selection. But evolution is constructive because of cooperation. New levels of organization evolve when the competing units on the lower level begin to cooperate. Cooperation allows specialization and thereby promotes biological diversity. Cooperation is the secret behind the open-endedness of the evolutionary process. Perhaps the most remarkable aspect of evolution is its ability to generate cooperation in a competitive world. Thus, we might add "natural cooperation" as a third fundamental principle of evolution beside mutation and natural selection. (Nowak, 2006)

Hence, biological and economic theorizing followed a similar path: they started first with the assumption that agents value only their own payoff; evidence suggested then that agents behave altruistically and, finally, theoretical models were amended and now incorporate different kinds of reciprocity.

So is it good news? Are we genuinely altruistic? First a precision: there is a difference between biological and psychological altruism, and the former does not entail the latter; biological altruism is about fitness consequencences (survival and reproduction), while psychological altruism is about motivation and intentions:

Where human behaviour is concerned, the distinction between biological altruism, defined in terms of fitness consequences, and ‘real’ altruism, defined in terms of the agent's conscious intentions to help others, does make sense. (Sometimes the label ‘psychological altruism’ is used instead of ‘real’ altruism.) What is the relationship between these two concepts? They appear to be independent in both directions (...). An action performed with the conscious intention of helping another human being may not affect their biological fitness at all, so would not count as altruistic in the biological sense. Conversely, an action undertaken for purely self-interested reasons, i.e. without the conscious intention of helping another, may boost their biological fitness tremendously (Biological Altruism, Stanford Encyclopedia of Philosophy; see also a forthcoming paper by Stephen Stich and the classic Sober & Wilson 1998).

The interesting question, for many researchers, is then: what is the link between biological and psychological altruism? A common view suggests non-human animals are biological altruists, while humans are also psychological atruists. I would like argue against this sharp divide and briefly suggest three things:

1. Non-humans also display psychological altruism
2. Human altruism is strongly influenced by biological motives
3. Prosocial behavior in human and non-human animals should be understood as a single phenomena: cooperation in the economy of nature

1. Non-humans also display psychological altruism


A discussed in a previous post, a recent research paper showed that rats exhibit generalized reciprocity: rats who had previously been helped were more likely (20%) to help unknown partner than rats who had not been helped. Although the authors of the paper take a more prudent stance, I consider generalized reciprocity as psychological altruism (remember, it can be both): rats cooperate because they "feel good", and that feeling is induced by cooperation, not by a particular agent. Hence their brain value cooperation (probably thanks to hormonal mechanisms similar to ours) in itself, even if there is no direct tit-for-tat. In the same edition of PLoS biology, primatologist Frans de Waal (2007) also argue that animals show signs of psychological altruism; it it particularly clear in an experiment (Warneken et al, again, in the same journal) that show that chimpanzees are ready to help unknown humans and conspecifics (hence ruling out kin and tit-for-tat altruism), even at a cost to themselves. Here is the description of the experiments:

In the first experiment, the chimpanzee saw a person unsuccessfully reach through the bars for a stick on the other side, too far away for the person, but within reach of the ape. The chimpanzees spontaneously helped the reaching person regardless of whether this yielded a reward, or not. A similar experiment with 18-month-old children gave exactly the same outcome. Obviously, both apes and young children are willing to help, especially when they see someone struggling to reach a goal. The second experiment increased the cost of helping. The chimpanzees were still willing to help, however, even though now they had to climb up a couple of meters, and the children still helped even after obstacles had been put in their way. Rewards had been eliminated altogether this time, but this hardly seemed to matter. One could, of course, argue that chimpanzees living in a sanctuary help humans because they depend on them for food and shelter. How familiar they are with the person in question may be secondary if they simply have learned to be nice to the bipedal species that takes care of them. The third and final experiment therefore tested the apes' willingness to help each other, which, from an evolutionary perspective, is also the only situation that matters. The set-up was slightly more complex. One chimpanzee, the Observer, would watch another, its Partner, try to enter a closed room with food. The only way for the Partner to enter this room would be if a chain blocking the door were removed. This chain was beyond the Partner's control—only the Observer could untie it. Admittedly, the outcome of this particular experiment surprised even me—and I am probably the biggest believer in primate empathy and altruism. I would not have been sure what to predict given that all of the food would go to the Partner, thus creating potential envy in the Observer. Yet, the results were unequivocal: Observers removed the peg holding the chain, thus yielding their Partner access to the room with food (de Waal)

(image from Warneken et al video)

2. Human altruism is strongly influenced by biological motives

In many cases, human altruism appear as a complex version of biological altruism (see Burnham & Johnson, 2005. The Biological and Evolutionary Logic of Human Cooperation for a review). For instance, Madsen et al. (2007) showed that humans behave more altruistically toward their own kin when there is a significant genuine cost (such as muscular pain), an attitude also mirrored in study with questionnaires (Stewart-Williams 2007): when the cost of helping augments, subjects are more ready to help siblings than friends. Other studies showed that facial similarity enhances trust (DeBruine 2002). In each cases, there is a mechanism whose function is to negotiate personal investments in relationships in order to promote the copying of genes housed either in people of—or people who seems to be of—our kin.

Many of these so called altruistic behavior can be explained only by the operations of hyper-active agency detectors and a bias toward fearing other people’s judgement. When they are not being or feeling watched, peoples behave less altruistically. Many studies show that in the dictator game, a version of the ultimatum game where the responder has to accept the offer, subjects always make lower offers than in the ultimatum (Bolton, Katok, and Zwick 1998). Offers are even lower in the dictator game when donation is fully anonymous (Hoffman et al. 1994). When subjects feel watched, or think of agents, even supernatural ones, they tend to be much more altruistic. When a pair of eyes is displayed in a computer screen, almost twice as many participants transfer money in the dictator game (Haley and Fessler 2005), and people contribute 3 times more in an honesty box for coffee' when there is a pair of eyes than when there is pictures of a flower (Bateson, Nettle, and Roberts 2006). The sole fact of speaking of ghosts enchances honest behavior in a competitive taks (Bering, McLeod, and Shackelford 2005), while priming subjects with the God concept in the anonymous dictator game (Shariff and Norenzayan in press).

These reflections also applies to altruistic punishment. First, it is enhanced by an audience. (Kurzban, DeScioli, and O'Brien 2007) showed that with a dozen participants, punishment expenditure tripled. Again, appareant altruism is instrumental in personal satisfaction. Other research suggest that altruism is also an advantage in sexual selection: "people preferentially direct cooperative behavior towards more attractive members of the opposite sex. Furthermore, cooperative behavior increases the perceived attractiveness of the cooperator" (Farrelly et al., 2007).

An interesting framework to understand altruims is Hardy (no relation with me) & Van Vugt (2006) theory of competitive altruism: "individuals attempt to outcompete each other in terms of generosity. It emerges because altruism enhances the status and reputation of the giver. Status, in turn, yields benefits that would be otherwise unattainable." We need, however, a more general perspective.


3. Prosocial behavior in human and non-human animals should be understood as a single phenomena: cooperation in the economy of nature

All organic beings are striving to seize on each place in the economy of nature - (Darwin, [1859] 2003, p. 90)

With Darwin, natural economy began to be understood with the conceptual tools of political economy. The division of labor, competition (“struggle” in Darwin’s words), trading, cost, the accumulation of innovations, the emergence of complex order from unintentional individual actions, the scarcity of resources and the geometric growth of populations are ideas borrowed from Adam Smith, Thomas Malthus, David Hume and other founders of modern economics. Thus, the economy of nature ceased to be an abstract representation of the universe and became a depiction of the complex web of interactions between biological individuals, species and their environment—the subject matter of ecology. Consequently, Darwin’s main contributions are his transforming biology into a historical science—like geology—and into an economic science.

I take the economy-of-nature principle to be a refinement of the natural selection principle: while it describes general features of the biosphere, it puts emphasis on the intersection between individual biographies and natural selection, and especially on decision-making. On the one hand, the decisions biological individuals make increase or decrease their fitness, and thus good decision-makers are more likely to propagate their genes. On the other hand, natural selection is likely to favor good decision-makers and to get rid of bad decision-makers. Thus, if our best descriptive theories of animal and human economic behavior indicate that all these agents have prosocial preferences and make altruistic decisions, then these preferences and decisions are not maladaptive and irrational. They must have an evolutionary and an economic payoff. Markets and natural selections requires cooperation, even if the deep motivations are partly selfish. Fairness, equity and honesty are social goods in the economy of nature, human and non-human.

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