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    Evolutionary Game Theory And The Mathematics Of Altruism
    By Catarina Amorim | July 18th 2008 10:24 PM | 41 comments | Print | E-mail | Track Comments
    About Catarina

    After many years as a scientist (immunology) at Oxford University I moved into scientific journalism and public understanding of science. I am...

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    Why do humans cooperate in things as diverse as environment conservation or the creation of fairer societies, even when they don’t receive anything in exchange or, worst, they might even be penalized? This is a question that has puzzled academics for centuries, especially since in evolution the basis for the “survival of the fittest” is, after all, selfishness.

    But in an article just published in the journal Nature, three Portuguese theoretical physicists developed a mathematical model capable of providing a way out from this conundrum through the introduction of social diversity - a ubiquitous characteristic of modern social networks - and suggesting that that the act of cooperation may depend on one’s social context/ranking. And in fact, when social diversity was taken into account the numbers of those cooperating increased in direct relation to the system diversity.

    Furthermore, cooperation, according to this model, spreads even faster when the act of cooperation is considered more important than the amount given, with these societies presenting also a much fairer distribution of wealth. This new mathematic model for society’s evolution is particularly interesting because not only it shows a logic behind the large numbers of cooperators that we know that exist in all human societies, but also it gives us a glimpse of the principles that can help “pushing” them into a better, fairer, path.

    Evolutionary game theory is a mathematical approach used to study (and predict) the evolution of social interactions, in which the study of conflict and decision-making is treated – like its name indicates – as a game. One such example are public good games (or PGG), which are frequently used to study cooperation as they look into social behaviour towards public goods - such as education, free health or even street lightning – those that every one can benefit from, regardless of how much they contribute (or not) to create it.

    Here because the individual’s benefits are independent of he/she contribution the most rational and selfish strategy (both in the games and real life) is to chose no-cooperation, what we know does not happen in real life. This is a good example of how difficult it has been to understand and create a theoretical model capable of explaining the emergence and prevalence of cooperation not only among humans but many other species.

    What Jorge M. Pacheco and Marta D. Santos (University of Lisbon, Portugal) did - together with Francisco C. Santos (Free University of Brussels, Belgium) - in order to overcome this apparent paradox, was to introduce into PGG, for the first time, a new variant – social diversity – in contrast to the models previously used in which all individuals were equivalent. Social diversity here refers to the characteristics typical of most social networks: the existence of individuals with different numbers and types of social connections, with few very highly connected and most with very few connections.

    Since PGG are represented as a mathematical formula, diversity was introduced as a new variable in the equation. Then Santos, Santos and Pacheco used this new altered formula to calculate the percentage of collaborators in the community, in function of population diversity (in PGG this would refer to the number and type of games each individual participated or, in other words, his/her “popularity”). And in fact, it was found, that in populations with high diversity, as diversity increased also did collaboration levels. The way PGG work is that each individual pays a certain amount to play (defectors play but do not pay/cooperate) and in the end profit, which is the total amount gathered in a game, is divided by all players. The reason why diversity increased cooperation had to with the fact that those few individuals with more connections and playing more games (the cooperators) would also have much higher “profits” and their impressive success would lead the other players to imitate their behaviour (even when the behaviour per se did not seem to improve directly their own life) resulting in an exponential increase of cooperation. In the same way, in real life the more connected/popular individuals are emulated, becoming role models and opinion makers.

    Equally the model also predicted that even when no-cooperators lead to new no-cooperators (as it happens many times in real life where this kind of behaviour can spread within groups) this will result in less profit, less success and eventually their own self-extinction with only a few sporadic ones left to parasite cooperators.

    Furthermore, it was also shown that the increase in cooperation was particularly accelerated when all individuals contributed to the games with the same total contribution, independently of the number of games played. This corresponds, in real life, to saying that if the act of contributing to the public good was seen as more important than the amount contributed, the percentage of collaborators in a community would grew much faster.

    Interestingly, the model, when applied in a more economical perspective, also suggests that these communities, with high diversity and where the act of cooperation is what matters, will also have a much fairer wealth distribution.

    Although this is obviously a very simple mathematical model and reality will never be as linear, Santos, Santos and Pacheco’s results gives us a total new perspective on how to look at ways of increasing cooperation/altruism and, consequently, also on how to create more successful societies, concerning issues as crucial to our survival as the protection of the environment or fairer social relationships, contributing in this way to the construction of a more peaceful world with less conflict and destruction.

    Article: Francisco C. Santos, Marta D. Santos & Jorge M. Pacheco, Social diversity promotes the emergence of cooperation in public goods games, Nature 454, 213-216 (10 July 2008) doi:10.1038/nature06940

    Comments

    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.

    this does not guarantee that you will benefit from that charity, nor does it discriminate against those who have not contributed to that charity.

    Gerhard Adam
    Doesn't matter.  If enough people support them, there is a sufficiently large number of charities to ensure that one of them will likely be available to help you as needed.  The few individuals that cheat the process by not contributing don't represent a sufficiently large number of offset the effect.  If too many individuals attempted to "cheat" then the charity system would collapse and there wouldn't be anything for anyone.

    So, the entire process works only because a large enough majority believe in its viability to support it and thereby provide benefit.  Not perfect, but it illustrates the point.  There is no requirement that there be a guarantee or a one-for-one correlation between the charity and the individual receiving the benefit.
    Mundus vult decipi
    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...).

    Gerhard Adam
    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.
    Mundus vult decipi
    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.

    It's an absurd metaphor to claim that atoms and molecules "cooperate" in the same sense that living creatures may.

    Chuck is right, atoms have no self-will and thus can not consciously cooperate with each other

    Gerhard Adam
    At the risk of opening up the debate again, there is nothing in the biological sense of cooperation that requires "self-will" nor "conscious" awareness of cooperation.  In fact, game theory suggests that the simplest implementations (i.e. "tit for tat") don't require anything except the ability to react according to past behaviors.  Since this has been done by machines (computers), it can hardly be argued that they are either self-willed or conscious.
    Mundus vult decipi
    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.

    Gerhard Adam
    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.
    Mundus vult decipi
    Molecules have to cooperate because the conditions require their reactions. Is that a choice?

    Our understanding of the chemical kinetics and the thermodynamics involved explains this cooperation. I am hesitant now to drop some technical terms because I am newly-charmed by the possibilities.

    Reactions can be reversible or irreversible. Further analysis is required. :)

    Gerhard Adam
    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.
    Mundus vult decipi
    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).

    Gerhard Adam
    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.
    Mundus vult decipi
    Steve Davis
    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 ;).

    Gerhard Adam
    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.
    Mundus vult decipi
    Gerhard, I don't think it makes sense to argue about some definition by pointing to phenomena in reality (i.e., ant colonies). Before you start studying ant colonies, you should already agree on what you mean with "cooperative", and only then you can find out whether an ant colony is cooperative by actually studying it. I don't think (in the sense that I believe) that cooperation requires deliberation and preference, I define it as such, and I think this is the definition that is most commonly used in social and behavioral science, and is also what is meant by cooperation in the article which started this discussion. And as a sociologist I think it is perfectly conceivable for humans to choose to live outside a society. In fact, some people do. (And also, some people do live in a society but cooperate less than others...)And yes, I think that ants cooperate in their colonies, and yes, I think ants have preferences and some cognition. It is really far-fetched to state that an ant is cognitively closer to a molecule than to a human. Ants have brains, molecules (obviously) don't. Which was really my initial point.

    Gerhard Adam
    I understand what you're saying, but I have to disagree with your conclusions. In the first place we have to consider whether we're discussing "cooperation" inside or outside of the group since the dynamics are quite different. In that sense, cooperation is required for the cohesion for the group's existence, but at the same time a different criteria is going to apply when individuals vie for position within a group. I do agree that one has to be careful that these terms aren't applied to absurdly reductionist levels, but that's part of the controversy when phrases like the "selfish gene" are invoked. My point about humans is precisely counter to what you've indicated, which is that humans have no biological survival value outside the group. You've indicated that some people choose to live outside of society, and I'm saying they are biologically irrelevant since they have no survival value. To have humans living outside of the group, would require an alternative group. For a human to survive outside of the group in a biologically meaningful way would require that have a geographic location which is populated by available mates, and a sufficient social infrastructure to support the raising of offspring and their future mate selections. This requires a human group. It is my contention that even people that supposedly live "outside of society", in reality, do no such thing. Simply considering the issue of geographic domain suggests that it simply isn't possible for such an existence. While I can appreciate your point about ant cognition, I think it is also stretching the definition to suggest that ants reflect preferences and make choices (with respect to participating in the colony). My whole point has been that there are certain evolutionary strategies that have proven useful to species by forming groups to gain advantage. The formation of such groups changes the selection pressures that individuals experience, since they will tend to favor those that are more "cooperative" rather than those that resist the group. As a result, over time we see distinct cooperative behaviors emerge as traits of the group and in some cases, such a high degree of specialization within the group that it can be arguably considered a super-organism. This doesn't change the fundamental nature of the individuals in it, but it certainly changes the path of their evolution from individuals to group members.
    Mundus vult decipi
    Rense,
    I don't think that you're doing sociology any favor by stating that you have to define cooperation before studying the field. You won't mean that cooperation is some platonic idea that exists independent from reality?
    On this blog we talk about reality.
    And for a chemist it is no problem to say that a molecule has a 'preference', but commonly we use the term 'affinity', which is a concrete, measurable property of molecules.

    Steve Davis
    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.
    Gerhard Adam
    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.
    Mundus vult decipi
    Steve Davis
    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.
    Gerhard Adam
    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.
    Mundus vult decipi
    Steve Davis
    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.
    Gerhard Adam
    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.
    Mundus vult decipi
    Gerhard,
    As a biochemist I see no problem in using the terms cooperation, preference or affinity on the level of molecules and atoms. When oxygen and hydrogen react, they both have to pay the cost of ionization. But what they get for it is the ionic bond, which only has meaning on the level of the H2O molecule. But the total energy is lowered.
    And I fully agree there is no intent or self-motivation needed for cooperation, preference or affinity.
    But on some other posts, you persistingly refuse to accept the concept of "intelligence without intent". What's the problem? If a chess program beats Kasparov, I would easily say that the program is pretty intelligent. Of course I know that the computer has no self motivation, but I know that the intent to win the game is built into the program by the programmers. Boy did they sweat.
    I'm aware I'm writing this 2 years after your post. Nowadays science is much more careful in using words like "free will" and such.

    Gerhard Adam
    I have a problem with the intelligence argument because using your example of the computer, where does one draw the line?  A calculator that can perform calculations faster than a human?  A can that can beat a runner?  These are obviously all tools that perform better than we do, so simply because one of them does better at us in chess is no basis for suddenly claiming "intelligence". 

    Intelligence is a slippery enough term as it is, so I try to distinguish between something that is intelligent versus something that simply emulates intelligence. 
    Mundus vult decipi
    Steve Davis
    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.
    Nicholas Horton
    Rense said:
    I was just trying to save the terminology of social science from polution by imperialistic natural science ;).
    I realize this statement was said (at least half) tongue in cheek.  But, I think it's important not to get too bogged down by the fact that different fields use the same words in sometimes wildly different ways.  I'm in Mathematics, and lord knows I had to lose my English-intuition a long time ago! 

    "Cooperation" in the Social Sciences implies far more than it does in Biology.  That's OK.  Each field has it's definition, and so long as we know what field we're in, we're cool. 

    Though, for my part, I prefer the Biology definition.   Cooperation, like "Rationality", is a too romanticized word in Social Science.  A hobby horse of mine is trying to remind Social Scientists that humans are only one more animal on the planet.  We aren't that special.  As such, studying us the way we study other animals is not a strange idea.  It's the right idea.   There are limits, obviously, but not as many as most Social Scientist would claim. 

    But, that would require using much of the same terminology of Biology to describe human action.  This is good.  The Biological definition of cooperation, I think, is more appropriate to how humans actually cooperate than the Social Science version (or at least the implied rationality that goes with the Social Science version). 

    [I don't want to sound like I'm harping on Social Science.  I just want it to be more firmly based on biology.]

    As for Dawkins' "The Selfish Gene" book, I always read it as a work of philosophical advocasy.  But, then, he may have promoted it that way after the fact.  I don't know.  But, from that perspective, it seems perfectly fine.  It's viewpoint is not ideal.  The word "selfish" is certainly prone to potential misinterpretations.  And, he, as a person, can be jarringly "agro".  But, the premise was/is powerful (especially for the time it was written).   Kin selection becomes more plausible when we consider the genes point of view.  If I die to protect my brother (who shares half my genes), the genes "win".   It was "selfishly" in their interest to get me to sacrifice myself if he lives to breed.  The particular copy of the gene in me "dies", but the copy of it in my brother lives.

    A gene can't be selfish the way we can, it's not alive.  A copy of me is not me.  But a copy of a gene (an exact copy) IS the same as the original.  I think this is why kin/group selection arguments, from the gene point of view, can make sense where from the organism point of view they don't.  Organisms are alive, can die, and are not equal to their copies/twins/clones.

    Again, it's the terminology problem.  "Selfish", like "cooperation" or "rationality", can have a weighted meaning.  For my part, I try not to take people too literally.  The "jist" is what's important. 
    Gerhard Adam
    " Kin selection becomes more plausible when we consider the genes point of view.  If I die to protect my brother (who shares half my genes), the genes "win".   It was "selfishly" in their interest to get me to sacrifice myself if he lives to breed."

    This is precisely where the problem comes in though.  Given the length of time (in humans especially) to raise offspring to the stage where they are self-sufficient and are able to reproduce would suggest that it is sacrificing the young that would be the biologically important thing to do, since they are more readily replaced.  An adult that sacrifices themselves for a child condemns them both to death.  The only reason this position isn't accepted in today's society, is because society itself has largely come to become the surrogate parent to try and ensure the survival of an individual's offspring.

    Even if we consider the example stated where I sacrifice myself for my adult brother, the "selfishness" argument fails since it can't know if my brother is capable of reproducing.  It would always make more sense to go with the sure thing (our own ability to reproduce) than to run the risk that the life we were saving was sterile.  In addition, the question arises; "which genes"?   Since my brother would not possess a complete suite of my genes, (and this is exacerbated with other kin arguments), the point becomes more muddled since there may only be a small percentage of genes I would be sacrificing to protect.

    The only point at which kin selection makes sense is when the adult making the sacrifice is beyond the age of reproduction (or has already contributed to the gene pool) and now wants to protect adult offspring to ensure the propagation into the next generation.  In my view, this is a much more specific example than the general explanation of kin selection offers.

    When other animals are considered, it isn't surprising when one observes that the young are the one's that are often the most expendable. 
    Mundus vult decipi
    Steve Davis
    Nicholas, I think there was not only a problem with terminology associated with selfish gene theory and kin selection. At the risk of offending a mathematician (I don't want to do that because I enjoy reading your comments) I think their attempt to introduce math to explain behaviour was inexcusable, an act of desperation. There are simply too many variables to even begin to work out who should save whom and why, and why did they introduce such a concept in the first place? It rarely occurs in real life, so why agonise over it in theory? It's possible that it all flowed from Haldane's view of the issue, but he had such a well developed sense of humour that I suspect he presented it in jest. It did happen in a pub, (from memory) after all. Here's my view of kin selection. If two people are drowning, a rescuer will rarely swim past a drowning stranger to rescue a relative. That's a scenario in which the variables are almost infinite. It highlights the folly of trying to use mathematical models or equations as evidence for theories of behaviour.  
    Nicholas Horton
    Gerhard,
    you make some good points.   The first one, in which an adult sacrifices themselves for a child would indeed be a problem.  The child would then only have one parent.  Raising children takes a long time for our species, and for years kids are "worthless" in terms of their ability to survive on their own.  Given an extreme situation where it's either you or your child who must die, and you have to choose, I can see how sacrificing the child makes sense.   The child will die either way, you might live to breed again.  With other animals, it may not be a problem at all, if their children don't take long to rear. 

    I think a better position on altruistic vs selfish behavior is a neutral one.    In some cases, like with the elderly, it makes sense to sacrifice yourself for your younger kin.  Other extreme cases may also hold, but generally, an individual is not looking to die and will try and avoid dying. 

    My point was simply that Dawkins' selfish-gene view has the potential to add to the discussion a perspective that may be useful, sometimes.  But, I'm not a Dawkins-fundamentalist.  The individual organism as the unit of selection still rings truer to me than the gene as the unit of selection.


    Steve,
    No worries.  I don't offend easily.  I think the first lesson you learn in math is that you are nearly always wrong!  And the person who points out where you are wrong is doing you a favor.  If my logic is flawed somewhere, I want to know it so I don't make the mistake again.  I think the only significant difference between math people and everybody else is their ridiculously high tolerance for failure.  :)

    But, this time I'll defend modeling with mathematics.

    Since ALL models in the sciences omit "real life" information out of necessity, are ANY of them useful?  Why?  If so, what are the criteria of a good model?  These are central questions for those of us doing applied mathematics.  And they aren't trivial.

    Sometimes the models we have are way too simplistic to be of any value.  BUT, sometimes the world is far too complicated for us to discern the underlying and central patterns involved (like your example of the swimmer).  A simple (in mathematical terms!) model can "cut through the fat" and highlight a cause, or a key effect.  Not all the information in the real world is relevant to the cause and effect relationship, it's just "noise" (like how many fish are in the river, or if they are within 50 miles of a town).  Sometimes the extra information that we omit IS relevant, but only in minor ways that would not significantly change the relationship (like the current of the river, or if it's light or dark, if the swimmer had been drinking).  In those situations, we accept that the model is not 100% accurate, but that its predictions will be probabilistically relevant and be central to the discussion at hand (is kin selection valid). 

    Models give us a simple way to visualize what we think is going on.  It's like using a story like "The Tragedy of the Commons" in Poli-Sci to highlight the potential for environmental damage caused by a society to a swath of land that is owned by no one.   Poker models are quite accurate in modeling the behavior of poker players, but poker is a very complex "social" interaction with a lot of chance and random variables.   Some of those variables are included, some aren't.

    But, you are dead right to question any attempt to generalize a micro-level model to the behavior happening at the macro-level.  The main reason is the idea of the Emergent Property.  Sometimes the macro-level behavior has properties that are not obviously related to the underlying micro-level behavior.  The classic example is the micro Brain, with all of its neurons, ACh, and goop, and the macro Mind, which we imbue with a near magical quality.  There IS a connection, and the mind cannot exist without the brain.  But, what the connection is is not as simple as knowing the basics of the micro behavior of the brain.

    Strangely, mathematicians tend to agree with your worry, but from a slightly different angle.  They are always convinced that scientists are "dumbing" down their mathematical models to make them simpler to understand, and in the process losing important information.  Scientist counter claiming that mathematicians just like complexity for complexities sake and aren't in a position to say what is and is not relevant information.  (As an example, you should hear what many Mathematicians in private say about Phyisicists.  To the rest of the world, Physics is way too math heavy.  To many mathematicians, it's "naive".)

    I'm actually inclined to agree with the scientists.  Math is useful when it's useful.  It doesn't always have to be complicated.  And it sometimes obscures the point when it is.  But, it's a fine line we walk.

    [a joke often heard by Physicists is, "physics is like sex, math is like masterbation."  Thankfully, Woody Allen unwittingly countered with, "Don't knock masterbation, it's sex with someone I love."]

    math is useful when its useful. keep that in mind. if you're standing on a tower and throw a stone the trajectory follows a parabole. the parabole has two solutions for h=0. one where the stone lands and the other has no meaning. math is useful, but sometimes meaningless.

    Gerhard Adam

    I realized I had failed to extend the idea regarding kin selection in the direction I had originally intended.  The original explanation struck me as too contrived and ultimately "selfish" which didn't make sense in terms of the real world where far too many variables may enter.  For example, we may not realize that an individual is kin, or the issue of age or survival potential, etc.  In all these cases, the issues only make sense when we take the time to evaluate them which is beyond the scope of behavioral considerations.

    It is my contention that the explanation is probably much simpler.  When a species has evolved to become more cooperative (as in pack or herd animals, as well as humans), there is a natural tendency to behave selfishly in the sense of looking out for our own best interests, but recognizing that cooperation is the only means by which this can be achieved.  As a result, it actually becomes a sort of "you scratch my back and I'll scratch yours" (or like "tit for tat" in game theory).

    My comment about the government behaving like a surrogate parent actually tends to lend credence to this perspective.  In other words, the behavior is not altruistic at all, because in a cooperative society, there may be a reasonable expectation that someone else may sacrifice themselves for YOUR child.  Therefore, there's an implied reciprocity (whether it actually occurs or is a reasonable expectation is irrelevant).  What does occur is that there is an implied sense that raising a child is everyone's responsibility to varying degrees.  We can see that this attitude is quite strong in our society, so one has to consider what conditions would have given rise to such a philosophical position within a culture.

    As game theory tells us we must be prepared to make these choices with imperfect knowledge, the force that drives the cooperation is the fact that these are iterative, so the "altruistic" choice must be based on the possibility of an iterative encounter for which a strategy like "tit for tat" may dominate the applied rationale.

    (As for the adult sacrificing for the child, we can easily see that many species recognize this liability which is precisely why they tend to over-produce offspring in the hopes that a few may survive.  The issue of altruism only becomes an issue when offspring are much difficult to produce and raise to maturity).

    Mundus vult decipi
    Steve Davis
    Hey Nick, if you can somehow work that last line into an article headline, you'll get a huge readership!
    Hank
    He's not kidding.   Two publications are still bigger than we are because they write headlines like 

    "Rough Sex at 40,000 Leagues Under the Sea" - Scientific American

    and 

    Bizarre Aquatic Creatures Are Secretly "Lesbian Necrophiliacs" - Discover 

    So we definitely need someone who can linguistically whore it out there for page views!
    It seems like a universe created with algorithms which directly foster altruism is comparable to scientific confirmation of a loving Creator with preferences. Perhaps as our mathematical understandings grow, universal moral principles can be derived directly from mathematical models.