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By Massimo Pigliucci | August 11th 2008 07:39 AM | 3 comments

About Massimo Pigliucci

Massimo Pigliucci is Professor in the Departments of Ecology & Evolution and of Philosophy at Stony Brook University, NY.

His research is on the evolution of genotype-environment interactions... Full Bio

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Researchers have tackled the problem of cooperation (and the related one of the evolution of altruism) for some time now. Initial game theory models suggested that cooperative animals would quickly be supplanted by selfish ones because of a cost to cooperation, as in the the so-called simple prisoners’ dilemma.

In these situations, typically an individual can choose between an action that benefits everyone, but at a cost to oneself, and an action that clearly benefits the individual, but comes at the risk of greater loss if everyone adopts that strategy. It turns out that most of us would rather lose than share, a sad but to many not surprising commentary on the human condition.

However, people eventually realized that cooperation makes the most sense in social groups, where phenomena such as kin and reciprocal altruism can take place. Sure enough, if the prisoners’ dilemma game is played iteratively, instead of in just one round, and if the players are allowed to keep track of what other players are doing (i.e., to build “reputations”) then it turns out that cooperating, even at a certain cost to oneself, is the winning long-term strategy.

Cooperation is therefore possible, but only if it aims at helping our close relatives (kin altruism) or if there is an expectation of reciprocity (I’m going to scratch your back now, but I fully expect you or someone else to return the favor, eventually).

Nature magazine recently published a fairly hard to read but fascinating paper entitled “Social diversity promotes the emergence of cooperation in public good games” (10 July 2008 issue). The authors, Francisco Santos, Marta Santos, and Jorge Pacheco (the first at the University of Brussels, in Belgium, the other two at the University of Lisbon, in Portugal) argue that cooperation in human groups can evolve as a result of social diversity, an outcome that would make concerted efforts at increasing diversity not only ethically good, but practically useful as well.

What Santos and collaborators did was to investigate what happens when another simplifying assumption of game theory models is relaxed: the players are usually considered equivalent in all respects, something that clearly doesn’t happen in the real world. So they simulated groups where there are different “neighborhoods” with individuals characterized by distinct degrees of “connectivity” to other individuals. So not to run into confounding effects from the already well known phenomena of kin and reciprocal cooperation, their model excluded those possibilities, focusing only on the possible effects of diversity.

The results were rather spectacular: Santos et al. observed a significant boost to cooperation in their simulated groups, which they attributed to the fact that the cost of cooperating was no longer fixed, as in the standard models, but depended on the social context of each individual. The more connected you are, the less the cost of cooperation, and as a result your fitness as a cooperator tends to increase. Moreover, the fitness of cheaters tends to diminish, because they breed other cheaters, to the point that their neighborhood is made mostly of cheaters, in a continuous negative feedback loop. If these scenarios remind you of well known human environments, you got the point.

It will be interesting to see what happens when one combines all three known sources of cooperation among individuals: kin selection, reciprocal altruism and increased diversity/connectivity. The point remains, however, that we now have an additional mechanism to explain an otherwise puzzling human (and some other primate’s) trait. Not all news is good from the Nature paper, however: it turns out that connected neighborhoods also create as a byproduct another new phenomenon not observed in standard models: unequal wealth distribution. Oh well, we really don’t live in the best of all possible worlds.

Comments

Gerhard Adam's picture

Cooperation is therefore possible, but only if it aims at helping our close relatives (kin altruism) or if there is an expectation of reciprocity (I’m going to scratch your back now, but I fully expect you or someone else to return the favor, eventually).

I'm sorry but this is much too narrow a focus and it simply isn't true. There is no such thing as a human (with a biological future) that doesn't cooperate. In fact,it can be argued that human beings will go insane if left alone. They absolutely depend on other human beings for their psychological well-being. A key point missing is that all animals will tend to cooperate against something outside the group. This will foster cooperation when it proves mutually advantageous over "going it alone". Humans like all other animals will cooperate (regardless of "kin" relationships) when threatened. There is little doubt that many species have understood the adage "there is safety in numbers" regardless of whether one is hunting or defending oneself (forgive the anthropomorphism). Humans cannot live by themselves, so in order to have mates, offspring, and mates for their offspring there is an absolute requirement for cooperation between different groups. In addition, kin relationships can't be absolutely known, so to suggest that evolution would provide such a specialized view, wouldn't make any sense. The notion of selfishness being presented isn't simply self-interest, but its almost pathological. There aren't many instances of where humans or other animals are militantly selfish. The iterated prisoner's dilemma indicates that cooperation is generally the result because the consequences of selfishness may be greater than the immediate benefit of defecting on any given trial. This becomes self-enforcing when the "Tit for Tat" strategy is employed by the competing parties. This is why cooperation tends to be the "glue" that holds groups together while competition is normally exercised within the group when vying for position. However even internal competition is tempered by the realization that nothing is gained if the group cooperation is lost. When predators cooperate they have the ability to side-step their individual characteristics and derive benefit from larger kills. It doesn't take much imagination to see that lone predators will tend to be less represented in the gene pool against their cooperative mates, and therefore cooperation will tend to be a characteristic of future generations. Similarly prey animals derive advantages by being in larger groups because they have additional warnings for predators and they primarily gain advantage in the sheer number of choices available to the predator. This will tend to result in the weakest members being taken, which would not happen in isolated encounters with individuals. (It's the old joke, "I don't have to outrun the bear, I only have to outrun you") In most cases, the cooperation is quite casual since it costs little and conveys much benefit. Most human beings certainly look out for their own self-interests (and their families), but they would hardly consider themselves actively selfish. It is equally important to note that no animal group coerces cooperation, so when it exists it clearly indicates that there is more benefit than cost involved in the relationship.
Steve Davis's picture
Nicely put Gerhard, and it's worth noting in respect of the predators you mention, that those that hunt alone, such as tigers, are the ones facing a greater threat of extinction. Once again we see in this article that the concept of cooperation is being analysed within the boundaries of selfish gene theory. Which is why we have the so-called "problem of cooperation." Cooperation is not a problem. It is the essence of life itself.
Gerhard Adam's picture
I think another point of confusion is the availability of resources. When resources are abundant cooperation has virtually no cost, so it is easy to see how it can arise. However as resources become more constrained, the competition within the group will escalate since the costs are correspondingly higher. This is what gave rise to the concept of the "pecking order",but this promptly disappeared when resources were abundant. One of the other elements in cooperation that is rarely considered is the role of "forgiveness" with respect to defectors in the prisoner's dilemma. There may be many circumstances where a freeloader can cheat the system and gain advantage from the cooperation to the apparent loss of the provider. However, if the long view is taken, it may be that there are varying degrees of "forgiveness" against future benefit which allows the freeloader to continue to be part of the group. Even in this situation, one would have to assess what the cost of driving off such an individual is versus simply tolerating them. I suspect that if resources become tight, the tolerance for freeloaders will also diminish.

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