Cooperating can often be in one's self interest in the long run (you give charity because you know you may need it yourself oneday). Interesting article but seems to knocking down a straw man.

An interesting thing about the model is its prediction of a "much fairer wealth distribution" in the communities with high diversity and acts of cooperation. Obviously, USA is not such a community.

However, the model is intuitively correct as far as economics is concerned.

The article discussed is very interesting, but suggesting that it "provides a way out from the question that has puzzled academics for centuries" is overstating it, I think...this model provides one particular solution to one particular social dilemma, using one particular type of analysis, namely evolutionary game theory. There are hundreds of other models that predict cooperation in certain social dilemmas under certain conditions; the challenge for social scientists is to figure out which of these conditions actually drive cooperation in reality (or defection, for that matter; let's not forget cooperation also fails very often...).

The problem with "altruism" is that it is based on the assumption that humans are competing individually in an evolutionary sense. However, this is incorrect.

While tigers may compete individually, the most primitive of humans has been subject to groups (ie: tribes) while modern civilizations are much more closely related to "ant colonies". Therefore it would seem that from an evolutionary sense, one has to weigh the selection pressures to promote group cooperation versus those that affect individuals. Similarly, the group, itself, has become a sort of organism in its own right and therefore the survival of individuals has become significantly less important than survival of the group itself. This will wreak havoc on the idea of selection pressures.

In part, the problem of "altruism" has already been seen in ant colonies and bee hives. A human being that does not possess cooperative tendencies to be a member of the "group" has virtually no survival value at all.

I don't think the point is cooperation, but rather does it support the survival of the larger group or not.

All this hand wringing over the origins of cooperation when its clear even from reading selfish gene theory that life only began when molecules first came together in cooperation. Cooperation preceded evolution.

Hi Chuck, thanks for your comment. You might recall that in The Selfish Gene Dawkins went to some trouble to explain to readers that the title was a metaphor, yet throughout the book and particularly p 157-8 from memory, he refers to cooperation between molecules and between genes (complex molecules) with no suggestion that this is metaphorical, nor does the text imply this. Cooperation is so fundamental to life that I've argued elsewhere (see The Secular Web) that cooperation between molecules can legitimately be regarded as the definition of life. As is clear from your comment cooperation within and between organisms is now so complex, requiring mathematical explanations as above, that it can be a shock to think of molecular cooperation. Even Dawkins concedes that this was the origin of life itself, so logic demands that cooperation prededed evolution. Steve Davis

I side with Chuck on this one... I think it's just pollution of terminology to call chemical reactions coopertion. In my book cooperation requires choice, and I wouldn't say that molecules choose what to do. In the end it's all a matter of definitions, but your definition of "cooperation" is so liberal that it renders the term almost useless.

I'm sorry but you might need to reconsider this definition. Cooperation, while it appears to be a choice, is actually a selection pressure on a species. There are numerous species that have no need to cooperate, however, humans do. From the earliest tribal societies, cooperation has been a selection criteria to enable survival, as a requirement for membership in a group.

Members that fail to cooperate, tend not to survive outside the group. Some members may cheat to exploit advantages, while others may actually go "overboard" in their level of cooperation. However in all cases, the cooperation is as much a selection pressure as the development of speed to chase down prey. Consider that herds of prey animals gain advantage by providing a predator with more choices and thereby improve their own chances of surviving. Contrast this with the animal that elects to be uncooperative. It has little to do with choice, but rather adaptation. Similarly, the wolf that cooperates and follows the rules of the pack has a better chance at surviving than one that will go it alone and it allows for larger prey without the need to increase the size of the wolf.

Therefore you could use this definition to conclude that molecules that "cooperated" gained some survival advantage (ultimately leading to multi-cellular organisms) than those that didn't.

The confusion regarding altruism occurs precisely because it is interpreted as being a completely voluntary act which is consciously made, rather than considering it from the perspective that it is a selection criteria for being part of the group.

While it is true that molecules can't choose to cooperate, it is equally true that humans can't either. Their choice is strictly limited to the degree of cooperation if they are to survive.

I think the problem is that the words we're using are too "loaded" with other meanings. "Cooperation" implies a specific type of social interaction, and "choice" implies a specific intellectual action.

In truth what is really being discussed is the concept of "cooperation" as a tendency rather than a conscious act. In this way, cooperation would be reinterpreted to mean something akin to "a mutually beneficial interaction". While this could include conscious decisions, it doesn't require them.

Similarly, the general definition of choice really needs to be constrained by the boundary conditions of survival or failure. In this way, it doesn't restrict the meaning, but it clearly outlines the consequences and why selection pressures would tend to favor one over the other. In humans, suicide is certainly a choice, but from a biological perspective it is a dead-end.

When considering the evolutionary consequences, we must avoid the tendency to examine individuals and look at the overall effects to derive meaning. This is why cooperation is an evolutionary selection pressure, since there are no instances of human populations surviving without it.

Similarly there are many species that could be described as cooperative, but whether they can choose their condition would also stretch the bounds of conventional word usage.

Changing the meanings of words is not going to help us to understand the world better. We invented dictionaries to settle those meanings ;-). Moreover, I think you are also stretching the meaning of "selection pressure". I am not a biologist, but as far as I know this refers to properties of the environment, not to the individual traits or behavior (such as cooperative behavior).

OK, but the dictionary definition of cooperation is "joint operation or action". There's nothing in there about choice.

Similarly the word choice means "to select freely or to have a preference for", while the first portion certainly implies conscious thought, the second definition requires no such restriction.

Selection pressure may involve the environment, but it is primarily whatever processes help a species survive to be reproductive. Cooperation is most definitely influenced by the environment, since many mechanisms employed by species could not function without cooperation.

If we take a simple example of humans, the fact is that human populations cannot survive without mutual cooperation between other humans. Therefore there really is no choice. However, if we consider what would happen to someone that was uncooperative, it doesn't take much imagination to see that their survival value has gone to zero. Regardless of whether they may have the skills to survive, their problems would extend to finding suitable mates, and then ensuring the survival of offspring. These are most definitely selection pressures and will not only dictate the direction human cooperation will take, but it will also be quite specific in ensuring that uncooperative individuals will not be part of the gene pool in the future.

How would you characterize ants or bees? Doesn't "cooperative" apply? How about wolf packs or herds of horses?

This isn't manipulation of the meanings, but rather it is recognizing that evolution doesn't occur with individuals, but rather it is determined by the direction the species takes. It is unnecessarily restrictive to define cooperation and choice as pertaining only to human beings.

Gerhard, I like your reasoning.
Rense, all I can suggest is that you check out Dawkins, nowhere does he state or imply that molecular cooperation is a metaphor. The key here is that it's all about increasing complexity. That first cooperation was basic in the extreme, so basic you don't like the concept, but that's because the entities were basic. As the organisms became more complex, so did the cooperation. Steve

I'm not saying that the concept of cooperation can only apply to humans, but it does require deliberate action and preference. I think it's a little far-fetched to say that molecules have preferences. You might say that we may then have to reconsider the definition of "preference", but just redifining our terminology isn't going to get us anywhere. Let's stick with the terminology that we have and try to figure out the mechanisms. If chemical reactions show patterns similiar to those in human cooperation, that's beautiful, but let's not be anthropomorphic about it.
Nor did I, by the way, dispute that cooperation might be a selective advantage. I just think that we should reserve the term selective pressure to the conditions that make something an advantage, or not. Whether group selection works is controversial, but that's another issue. I was just trying to save the terminology of social science from polution by imperialistic natural science ;).

I'm sorry, but I have to dispute your claim. I don't believe there's an argument to suggest that an ant colony is not cooperative. It certainly has all the appearances of cooperation. So if its not cooperative, then choose the word to describe it and we can use that.

If it is cooperative, you cannot make the argument that the choices are either "deliberate" or reflect a "preference". The cognitive abilities of ants would be more analogous to molecules than humans, so by that standard, we will have demonstrated that no conscious process is required to achieve it. Often the argument uses vague terms like "instinct" to describe behavior, but that really only suggests some autonomic response, just as we are attributing to molecules.

It would also be worth exploring what makes you think that cooperation is "deliberate" or requires a "preference". I don't think there's any dispute that human society is a cooperative venture. However, I would dispute your claim that participation is "deliberate" or reflects a "preference". You may choose to participate in a variety of manners, but you cannot choose to not participate at all. The portions for which deliberate choices and preferences can be used are a relatively small part of the entire society.

Similarly, it cannot be disputed that the formation of a wolf pack conveys advantage, because it enables smaller animals to prey on larger animals that would otherwise be unattainable. Also the formation of herds increases the choices a predator has, so by probability alone, the survival advantage to individual members is increased over being isolated in an encounter. Once again, by any criteria you care to use, if advantage is conveyed and it affects the probability to reproduce over others, then it must be a selection pressure. The only criteria that is required for something to be selective is the ability to be passed on to subsequent generations. There really isn't even any requirement that it be genetic if it can be conveyed as an idea, or taught. Consider that birds are taught their songs which, in part, are used to identify themselves and attract mates. Any bird that fails to learn it properly will have less chance to reproduce and therefore tend to be "selected" out.

I'm not sure what your definition of "group selection" is, but it would be an extreme viewpoint to suggest evaluating animals on an individual basis when they don't exist as such (ie: ants and bees, even humans). For example, using ants again, it is meaningless to ask what the selection pressures on an individual ant are, since they have no reproductive capacity outside the colony.

While I understand what you're sayihg about words like "cooperation" coming from social science, my point is that if we don't like the common usage of a word, then we shouldn't use it to describe scientific principles that may describe phenomenon outside the common behavior. This is something that is routinely encountered in quantum physics and, frankly, biology can't be that squeamish about the words we have to work with.

Confusion over terminology came to the fore with the selfish gene. I know, I know, if I can have cooperating genes surely we can have selfish genes. We certainly can, but not in the sense that Dawkins used the word. It's clear from this discussion and the work of all biologists on the matter including Dawkins that the genes that combined and cooperated survived, while those that did not, that acted alone, that were self-ish, did not survive or develop. It's the cooperators that Dawkins describes as selfish, for the sole reason that they survived. The word was inappropriately used, he was describing a useful concept, but the only entity in the scenario presented that could be labelled as selfish was the loners. This was a disservice to evolution theory because it pushed cooperation almost entirely out of the debate for decades, and it's only in recent times that its importance has begun to be rediscovered.

I think we're placing too singular a definition of these words instead of viewing them as concepts that explain or describe larger principles.

Selfishness is not mutually exclusive with cooperation in the same sense that we can employ a phrase such as "the enemy of my enemy is my friend". Cooperation can be driven by completely selfish motives (which it clearly is when survival is at stake).

Instead of viewing these attributes as absolute, we need to consider them as opposing forces towards a particular goal, each of which may have many manifestations and contexts in which they can occur. Human society is cooperative, but that doesn't preclude selfish behavior for competition within that society. Its a question of balance; if one is too selfish, then one may be viewed as uncooperative and lose support within the group. If one is too altruistic, it may leave one open to exploitation by the group. There are numerous combinations which can occur however, these various strategies would only have meaning within an already cooperative setting. In other words, cooperation provides better assurance of survival, so now we can compete for position.

During the turn of the last century there was a case in Africa of two lions apparently cooperating in attacking humans building a railroad (they made a movie out of it). From the information available, this was successful, but in truth, it was too successful. It caused a response from the species being hunted (humans) in a fashion that killed both lions. Therefore, "cooperation" in this case was "selected out". This doesn't mean that it was a bad idea, or that it couldn't occur somewhere else, but rather than in this particular case, "cooperation" resulted in a response that made it a less desirable survival strategy.

My only point in this story is that it provides a means of visualizing exactly the kind of "cooperative" and/or "selfish" behavior we've been talking about and how consequences may derive from them.

While some people may balk at the words being used in this fashion, I think they convey exactly the concept we want them to. The only real problem is if one assigns a "willfullness" to them which doesn't exist.

Exactly so Gerhard, and you have (possibly inadvertantly) illustrated my point that it is a disservice to biology to assume as Dawkins does that because forms of selfishness exist that they are the basis of evolution.
I think we could make a valid case for your great idea that "cooperation is for survival, competition is for position." It fits well with a point I made elsewhere that we are so dependent on cooperation that we take it for granted like the air we breathe. In such a cooperative world, competition stands out for its novelty.

Actually Dawkins emphasizes the Evolutionarily Stable Strategy (ESS) as the criteria for determining whether cooperation or conflict will dominate. He even introduces the idea of "cheating" as a vehicle where some members of a species may gain advantage, but that it isn't a global strategy that would remain effective.

Essentially, Dawkins is actually proposing a form of game theory to explain behaviors using the ESS as the basis for determining what strategies are possible.

In general, no animal will take a risk that outweighs the benefit to its survival, so the need to cooperate, or at least be neutral, well tend to be the norm unless the risk to survival is high enough to overcome all other risks. In this sense, a member of a species is selfish since they won't run a risk for other species members that don't ultimately benefit themselves.

Part of Dawkin's point is to illustrate that evolution doesn't cooperate solely for the survival of the species, but rather gains that objective by behaving selfishly for its own survival (similar to economic theory).

Dawkin's also recognizes that there are cooperative species like bees where sacrificial (or altruistic) behaviors will advance the survival of the colony, but because the individuals have no reproductive capacity outside the hive, they are basing their risks on hive rather than individual survival.

I would certainly agree that competition in the traditionally aggressive manner in which we think of it is not a trait of most species, since it simply wastes too much energy and has too high a risk for little benefit. Therefore the trend is to be neutral or cooperate to whatever degree is necessary for survival.

Clearly there are many nauances that exist in describing such a set of attributes, but I would agree that the world is fundamentally cooperative with competition representing the exception when risks to survival are high or for "position" within a cooperative group.

And this is where I (and you I think) differ from Dawkins. Where you conclude that bees "base their risks on hive rather than individual survival" a conclusion I think most people would agree with, Dawkins would disagree, saying they base risks on survival of their own genes, which by an accident of evolution happen to be the genes of the hive. It's such a narrow view, a gene's-eye view, (that metaphor is more telling than its proponents would like to admit) that it does not allow for, in fact excludes, the nuances that are found in the natural world.

In fairness to Dawkins, one of the initial points in the book is that the prevailing view suggested that an animal's behavior was intended to enhances the survival of the species. This clearly suggested a level of altruism which doesn't exist and misrepresents the idea of species survival.

Instead Dawkins suggested that the perspective be shifted to look at things from the gene's "point of view", so that we could re-interpret events in light of the fact that the gene could only act by (and for) itself and that species survival arose from the fact that each gene acting selfishly is what produced that result and all other interpretations could be derived from it as well. Despite the use of the term "selfish", the gene has no "choice" in the matter, since it cannot interact with anything outside of itself anyway.

My position is that both viewpoints run the risk of anthropomorphism because regardless of how we mean them, these terms will give rise to interpretations based on human interactions. This is precisely why there have been earlier disagreements on how they should be used and what they mean within the context of species survival.

However, I would suggest that cooperation and selfishness are two sides of the same phenomenon and are present in every interaction. Just as physics has the problem of wave-particle duality, so that the particular characteristic being seen is subject to the nature of the observer (or experiment), so it is with cooperation and selfishness. It is not out of the question to be cooperative for completely selfish reasons.

Another problem that Dawkins realized was being too reductionist by considering the selfishness of smaller and smaller components until it became an absurdity (such as thinking that an Oxygen atom would derive benefit by cooperating with a Hydrogen atom and by behaving selfishly assure its survival.

Ultimately all these different attributes have to be assessed from the perspective that they occur independently of intent or evaluation. Individual animals do not assess the value of a particular action, but rather are driven to a particular course of action by their natures. Clearly, their "natures" are genetically derived, so in that sense Dawkins is correct.

However, selfishness isn't an absolute concept and as a result we see much cooperation (or at least neutral behavior) because it is beneficial to do so. If one was inclined to take the comparison down to absurd levels (i.e. the atom), the argument could be made that all chemical interactions will ultimately resolve down to the lowest energy states, or basically, to follow the path of least resistance.

This could be a useful interpretation when examining the basis for other interactions, since a similar criteria would result in the greatest benefit for the least energy expenditure, or following the path of least resistance.

This would suggest that the typical behavior we would expect to encounter is that each individual would behave in a manner that resulted in the highest benefit for the lowest risk (or least expenditure of energy) to achieve its objectives. As different circumstances arise, then different strategies are employed (ie. reproduction versus food). However, in the end, each should illustrate a tendency use the most "efficient" behavior for the desired result.

We should bear in mind that "efficiency", in this context, refers to the particular ecological niche occupied by the species, so it may be a more intensive energy consumer than others, but for its particular environment, it will probably be the most efficient under the circumstances.

We would still expect variations from this central path, primarily from non-adaptive forces (mutation, recombination, and genetic drift), but from an overall perspective, any given species should not deviant very far from its optimal course left unmolested. Once again, if new variables are introduced or conditions change, then chaos will prevail until a new equilibrium is reached.

So to return to our basic terms, competition doesn't mean aggressiveness or confrontation any more than cooperation leaves one open to exploitation. The best choice for any species is the most efficient use of its energy for the lowest possible risk to attain a particular objective. This is precisely why most species are not only tolerant of each other, but will minimize confrontations that risk injury or waste energy.

Gerhard, if Dawkins presented his case in your reasonable, non-dogmatic, non-judgemental way I would be far more sympathetic, but his schoolboy attitude and intolerance leads him to ignore the standards of the scientific method and reach conclusions that are not sustainable.
John Maynard Smith was spot on in his review of The Selfish Gene, "...it reports no new facts. Nor does it present any new mathematical models at all. What it does offer is a new world view." Dawkins clearly missed the implied criticism because he included this review in the 30th anniversary edition.
A world view is exactly what The Selfish Gene was, one person's opinion on how certain data should be interpreted and applied, so I'm afraid I have to differ from you here, we do not need to be fair to Dawkins at all.
Serious, considered, careful work on selfish gene theory had been going on for decades, yet one man suddenly sprang to prominence and convinced the world that this was the full story of evolution. How could this happen? Because he was a wordsmith, a populariser, an embellisher, a journalist of biology, and because his work was not presented in a scientific manner he somehow escaped the scrutiny that we all accept is necessary to prevent science going up blind alleys. He will defend his use of the word selfish until the day he dies, but it cannot be denied that the genes he describes as selfish are the cooperators, the loners were selfish if the word has any technical use at all, and they were the non-starters in the story of evolution. The blind alley he produced was to exclude cooperation from its rightful place in the discussion, to such an extent that now that it is finally being treated seriously, that treatment is only within the framework of selfish gene theory as the article above indicates.