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By Steve Davis | May 15th 2009 01:37 AM | 52 comments | Print | E-mail | Track Comments
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About Steve Davis

gadfly: noun (1) fly that stings horses and cattle. (2) (derog) annoying person, esp one that provokes others into action by criticism, etc.... Full Bio

David Sloan Wilson is doing a great job in his blog at The Huffington Post, exposing the contradictions that lie at the heart of selfish gene theory.

He’s run a series of articles under the umbrella title of Truth and Reconciliation for Group Selection, singling out Richard Dawkins as a target for some pretty heavy broadsides. Like this; “Richard has become unaccountable, in part by becoming a public icon. That disqualifies him as a spokesperson for science.” And this; “when it comes to semantic confusion, you can't beat selfish gene theory.”

It just makes me feel warm and tingly all over.

But like most critics of the gene-centric view of evolution, DS Wilson has somehow fallen into the trap of accepting some of the assumptions and rhetoric that underpin the gene-only view. An example can be seen in the following passage; “Selfishness beats altruism within groups. Altruistic groups beat selfish groups.” This was the conclusion that David Sloan Wilson and EO Wilson reached in their paper Rethinking The Theoretical Foundations of Sociobiology, but David Wilson is repeating it so frequently, (albeit in the very good cause of undermining selfish gene theory) that it’s becoming a mantra. The problem with this particular mantra is that the proposition contains a flaw.

That flaw flows from the acceptance of the validity of the Hamiltonian interpretation of altruism, whereby an altruistic act decreases the fitness of the altruist and increases the fitness of the receiver. (That acceptance was not just implied, but actually stated in one of the series articles.) But acts that decrease biological fitness are rare in nature, because such acts present a significant risk to the survival of the altruist. Where such acts occur, it’s highly likely that the altruist makes a mental note to avoid a similar situation in future or to approach it differently.

How many of us help others to the point where our personal well-being is at risk? It just does not happen often enough to be a factor in evolution. So what form of altruism is it that is well known, accepted, even admired? It’s that form of giving in which the giver has no expectation of receiving in return. And in most of these acts, altruists give that which they have in excess, whether that be an excess of goods, time, labour, or in the case of humans, money. Because they give what they have in excess, their fitness is not affected.

But we still help others; our daily lives are an endless round of helping. How do we achieve that without adversely affecting our fitness? By constantly receiving help from others. While we are giving, we’re receiving. It’s that cycle of mutual aid that defines social life. It is therefore the quality of the system of mutual aid, and the quality of the commitment to mutual aid, that determines the fitness of particular groups. And of course that fact applies at all levels of selection. For example, it’s the quality of the system of cooperation between cells, between organs, and other physiological features such as the immune system, that determines the fitness of individual organisms.

This matter of mutual aid was recognized by the Wilsons in their paper, but misinterpreted; “It is simply a fact of social life that individuals must do things for each other to function successfully as a group, and that these actions usually do not maximize their relative fitness within the group.” They overlooked the important point that givers are also receivers. They again came close to solving what they referred to as “the fundamental problem of social life” with this; “Traits that are “for the good of the group” are seldom selectively advantageous within groups.  At worst, they are highly self-sacrificial. At best, they provide public goods at little cost to the actor, making them close to selectively neutral, or they constitute a stable local equilibrium.”

Their “at best-at worst” comparison is the problem here. It seems to imply a balance or equality of frequency and value between the differing forms of altruism, but if it was possible to draw a graph there would be no horizontal line or bell curve, it would show a curve or straight line heading up towards neutral with highly self-sacrificial hovering just above zero.

So unfortunately David Sloan Wilson, for whom I have the greatest respect, in his noble quest to analyse group selection in a big-picture manner, has been diverted from the big picture. He has focused his attention on a perverted and irrelevant version of altruism when the principal element in group fitness, and hence group selection, is clearly cooperation in all its forms, of which altruism is but a part.

Let’s alter his proposition to see how it reads in light of those facts. We now have; “Selfishness beats cooperation within groups. Cooperative groups beat selfish groups.”

The second sentence is obviously true, and actually summarises a well-known yet much neglected passage from Darwin’s "The Descent of Man", as the Wilsons acknowledged. The first sentence is too concise to be meaningful. It could refer to selfishness, or cooperation, as a single act or as a concept, a way of life, or a state of mind. But importantly, it brings into focus the inconsistency contained in the original proposition. If selfishness beats cooperation within groups then there is little likelihood of the development of group cooperation to a level that is effective. Likewise for the original proposition. If selfishness beats altruism within groups, then altruistic groups are unlikely to develop. It’s doubtful that a split personality makes for biological fitness.

So it’s time for David to ditch the mantra. It’s served its purpose. It’s shown a lot of people that selfish gene theory is not the only game in town. Now he should concentrate on cooperation as the principal factor in evolution.

Cooperation is the principal factor in evolution? Now that’s a mantra! And here’s a challenge to all the gene-centrics out there: Find a flaw in it!

Comments

“Richard has become unaccountable, in part by becoming a public icon. That disqualifies him as a spokesperson for science.”

Yes... Spokespeople should avoid the public entirely. But how logical are people that think ad hominem attacks belong in scientific debate?

The word "mantra" does not mean what you think it means.

" If selfishness beats altruism within groups, then altruistic groups are unlikely to develop. "

Don't the Wilson's propose mechanisms by which altruists groups could arise despite being susceptible to invasion by selfish individuals? If it's possible to recognize other altruists and then selectively interact and form groups with them, then we would perhaps expect to see cooperation arise, right? I believe they propose another mechanism as well.

I really like your take on this. I think considering "cooperation" instead of "altruism" is a great idea. Have you published the thoughts that you outlined about? I'm not wildly familiar with this literature.

Gerhard Adam's picture

"Don't the Wilson's propose mechanisms by which altruists groups could arise despite being susceptible to invasion by selfish individuals?"

I think such a position does not represent reality.  Among social animals, I have never seen a single instance where "strangers" were automatically admitted to the group, or granted any "privilege".  This would suggest that groups are highly selective in who they allow in, and therefore would be extremely difficult to penetrate by "selfish" individuals.


In fact, it is an easy argument to make that it is the "selfish" individual that runs the highest risk since they can't survive as individuals, therefore there is tremendous pressure to conform and gain acceptance by the group.  Even under those circumstances it may take a long time before such acceptance is assured.



In the above post I had intended to say, "....then we would perhaps expect to see altruism arise, right?" Although I think the same argument would hold for cooperation.

Gerhard Adam's picture

“Selfishness beats cooperation within groups. Cooperative groups beat selfish groups.”

In my view both statements are incorrect without establishing whether or not the possibility of future encounters is included (game theory).  Clearly the greater the number of encounters between individuals or groups, the greater the pressure to cooperate, so that some degree of mutual benefit may be achieved.


I'm also not sure on why there seems to be a tendency to immediately jump to the conclusion that cooperation or altruism creates a "risk" for the giver?  In the first place, we need to determine what the actual risk to the individual is anyway.  Obviously predatory animals have different risks than prey animals and consequently the role that cooperation or altruism plays would be very different.

However, the concept of "fitness" is too often abused in these discussions because it really means "how successfully one reproduces".  Therefore, short of death, you would be hard pressed to demonstrate that being altruistic is anything but beneficial if it attracts more potential mates and garners the support of the group by your actions.  I haven't seen ANY described scenarios whereby selfishness garners more potential mates.

As we've discussed before, there is no question that there is competition within the genders for mates, but to simply ascribe it to the group would be missing the point.  I don't believe it is a coincidence that humans have developed many philosophical points regarding "honor" and "obligation".  Both of these strongly suggest that human psychological evolution recognizes that altruism incurs a debt from the receiver which would make it a much more positive force in group dynamics than a liability.

Group identity is probably one of the strongest forces that brings individuals together and there is plenty of evidence to suggest that among many of the "higher" animals that strangers are not readily accepted.  This would suggest that an element of trust is necessary for inclusion in the group from which the concept of "altruism" can more readily evolve since the presumption would be that one wouldn't be exploited by extending such cooperation. 

I don't think there's any stronger example of "cooperative/altruistic" behavior than "elite" military units where each member is an indispensible component of the entire apparatus.  Any sense of "selfishness" would destroy such a unit and it seems highly unlikely that we could even have such things if "selfishness" were the rule of the day.



"I don't think there's any stronger example of "cooperative/altruistic" behavior than "elite" military units where each member is an indispensible component of the entire apparatus. Any sense of "selfishness" would destroy such a unit and it seems highly unlikely that we could even have such things if "selfishness" were the rule of the day."

So you are saying that cooperation cannot result from selfishness? Taking your military example, I think I am going to fight with fellow soldiers instead of going it alone, because that increases my chances of survival. I will also give you a boost into an apple tree if it helps me get food. So I am finding it difficult to see how any sense of altruism could NOT be framed as selfish behavior in some regard.

Gerhard Adam's picture
I'm replying to this now, because I must've missed it earlier.

Taking your military example, I think I am going to fight with fellow soldiers instead of going it alone, because that increases my chances of survival

The problem with that example is ... why did you join the military in the first place, an altruistic organization?


I will also give you a boost into an apple tree if it helps me get food.

Why should motivation come into it?  If you perform a cooperative act, then you're cooperating.  Whatever is going on in your brain is irrelevant.  It's like grumbling if your boss orders you to do something.  If you do it, then what difference does it make?  Your grumbling is irrelevant to the result.

I'm also not sure on why your desire for food is elevated to the level of selfishness.  Self-interest, yes.  Selfishness?  Why?

No one is suggesting that self-interest isn't part of biological behavior, but it doesn't rise to the level of selfishness (since that is the opposite of altruism), this isn't just an idle semantic distinction.

It is nearly 4am and I have been drinking, which is perhaps why I spent felt the need to reply. Anyway here is a reply from a gene-centric.

Firstly the author seems to misunderstand the concept of fitness and consequently fails to see the costs involved in altruistic acts. Fitness is not primarily concerned with the survival of the individual, but with that individuals potential to contribute evolutionary to future generations. Obviously survival is important in so far as it allows the individual the opportunity to reproduce in future, but survival is not the only thing that can affect reproductive potential.

Time, food, labour, money or anything given in an altruistic act could have potentially been invested in reproduction. There is an opportunity cost to any altruistic act, and that can be measured in the reduction in reproductive success of the individual, no matter how small this may be. Dawkins refers to this cost as being definable in terms of "gonad equivalents". Any excess could be directed towards producing offspring and even though the exchange of resources for offspring may be inefficient, it would still improve the fitness of the individual.

Secondly, there is a significant difference between altruism and cooperation. An act is altruistic where it helps a second party at a cost to the individual. But in the case of cooperation there need not be a cost to the individual, there is likely to be a benefit. This is why altruistic acts between cells, organs, related organisms are quantitatively different. The fitness of both individuals involved can increase in these transactions, and it therefore isn't in fact altruistic.

Reciprocal altruism is also different from simple altruism. If I know I am getting something in return for my altruistic act, the benefit I receive may in fact outweigh the cost, and therefore my act is no longer altruistic. Of course it is impossible to "know" that I will receive such a benefit, the receiver may benefit from my act and then refuse to reciprocate in future. A number of mechanisms can be found in nature to counter such a situation, ranging from punishment of the defaulter, to withholding of altruism in future. These responses can be explained simply in a gene centric view, but are difficult to account for from a group selectionist standpoint.

As to "Selfishness beats cooperation within groups" being too concise to be meaningful, read the literature if you want an explanation, is this not meant merely as a one line summary of an argument, a "mantra"?

Gerhard Adam's picture
It sounds like the meaning of altruism and cooperation are being parsed a bit too precisely.

Despite the overarching definition, can you provide one or two scenarios that would illustrate what you mean by "altruism" versus "cooperation".  The reason I ask, is because the narrow definition I'm reading doesn't sound like it actually exists anywhere in nature (except in those individuals that might be on the psychological fringes and not representative).

Gerhard Adam's picture
I think another point to consider is that "altruism" could also play a role in the same fashion as the peacock's feathers do.  They convey no direct advantage beyond demonstrating the fact that the male with the "best" display is the better mate because he can "afford" the display.

Similarly, some "altruistic" behavior may be in the same category by indicating that the individual is better fit for survival, even when performing actions that appear to "cost" something.

but Dawkins make it clear in TSG that the "selfishness" was of the gene (the whole book is about genetics from the gene's point of view) not the individual

Gerhard Adam's picture
Except that it isn't.  Dawkin's point was that the gene influences all the other elements of behavior and evolution because it is "selfish".  This is a preposterous claim, but otherwise it would be even worse since a gene can't be anything, let alone "selfish".

Steve, you seem to have absolutely no understanding of Dawkins' argument. Even a cursory reading of "The Selfish Gene" reveals that his theory EXISTS to explain why cooperation (actually, altruism, to which I will return) exists in nature, when selfish acts would seem by their definition to be selectively advantageous. His explanation is that cooperation and altruism serve to increase the likelihood that genes shared between members of a group will be passed on, since genes (as Luke perceptively noted above) are the stuff of reproduction. It is genes, Dawkins argues, not organisms, that are being selected for - enabling individual organisms to perform seemingly anti-evolutionary acts. You have neglected to discuss genes at any point in your criticism of Dawkins here, instead focusing on the behavior of organisms and groups, and on the semantic qualities of "altruism," "cooperation," and "selfishness" - thereby failing utterly to engage your opponents' argument.

Also, I'm no zoologist, but Dawkins' book is replete with examples of genuine, self-sacrificing altruistic acts performed by organisms on a daily basis, so your argument that true altruism is too rare to be relevant ignores not just the argument of Dawkins' book, but its substance. It's fine if you don't find his examples compelling, but you can't simply dismiss them with a facile analogy to human society - arguing that by and large we only give what we have in excess, so shouldn't nature operate the same way? That statement doesn't become correct by virtue of being internally logical. You need to confront the case evidence - to say nothing of the fact that even social humans have the capacity for truly self-sacrificing behavior, which almost certainly happened regularly enough to influence evolution at some point in our species' past (if, indeed, it has ceased to do so today).

Gerhard has made some attempt to include Dawkins' argument in his refutation of it, but in so doing has slashed it completely to ribbons. Dawkins' point is not that some magical, anthropomorphic quality of genes causes them to influence evolution - that's the kind of nebulous nonsense in which real biology does not engage. It is rather that when we speak of "natural selection" we think that the relevant factor is the survival of an organism, when in reality it is the survival of a gene. Genes, after all, are the mechanism for inheritance, and may be present in that organism's kin. Again, without such a theory, Dawkins' numerous examples of genuine altruism (a bird crying out to alert its flock to the presence of a predator, and thereby drawing the predator's attention to itself) would be wholly inexplicable. I don't see a single line of the debate above that is pertinent to that thesis, and to argue that Dawkins' argument is wrong because genes are not truly "selfish" in the emotional sense is to make an argument he addresses in his introduction - and therefore to reveal, again, ignorance of his work.

And enough ink has been wasted on this preposterous inverse correlation between Dawkins' popularity and his scientific credibility. That's the worst, least founded form of elitism. I wonder what Stephen Hawking and Albert Einstein would have to say about it.

Gerhard Adam's picture
"It is rather that when we speak of "natural selection" we think that the relevant factor is the survival of an organism, when in reality it is the survival of a gene."

Sorry, but this is precisely why I said it was nonsense.  Might just was well argue that the purpose of genes is to advance the survival of carbon atoms.

The gene cannot direct outcomes and therefore whatever organism the gene produces will survive on the traits that it has.  The gene cannot benefit, because there is nothing to benefit.  To suggest that the gene has a reason to survive is ludicrous, since the gene has enough trouble just maintaining its integrity without mutation.  It is the agent of change, not the benificiary of conservation.

The example of the birds is NOT what Dawkins says.  In fact Dawkins specifically says:

"There is no suggestion that the caller is 'trying to draw the predator's fire' away from his colleagues.  .... Nevertheless the act of calling seems, at least at first sight, to be altruistic, ..."

He then goes on to make his point which is:

"There are many ways in which the caller could gain selfish benefit from warning his fellows."

The primary point is TSG is to point out how even altruistic behavior can ultimately be linked to selfish intentions.  The seriously flawed model of kin selection is another aspect of this.

Gerhard, I can see where you would take me to have implied that the bird's INTENT was to draw the predator, which is certainly not the case, nor is it what I meant - the attraction of the predator's attention is a result, not a goal. It was intended as an example of a behavior of substantial risk to the actor, which has nonetheless not been eliminated by natural selection, not an example of suicidal altruism (though the latter is not unprecedented).

You're again attacking Dawkins' central point by erroneously implying that he anthropomorphizes genes, giving them a "reason to survive." Of course they don't WANT in any conscious or directed way to be passed on to the next generation of organisms - it is simply that, by definition, they will pass on if they are built to do so and will not if they are not. Let's take for example a hypothetical gene that causes a bird, when it notices a predator, to call out to its flock and alert them of the predator's presence. Attracted by the noise, the predator kills that particular animal, but the rest of the flock has a split-second head start and escapes. So that particular instance of the calling gene has been removed from the pool - there are, however, ten direct relatives of the caller in the flock, all callers or carriers of the gene, and they survived. The gene will be found in the next generation of birds; not because it wanted it really hard, not because its PURPOSE was to survive, but simply because survive it did.

Let's take the same flock, but the bird who notices the predator does not call out. Armed with a split-second head start, the bird who noticed the predator escapes - the predator, however, has time to kill three animals, all direct relatives of that bird. The noticer's genes will now certainly appear in the next generation of birds, but instances of those same genes present in his relatives will not - even though he is the fitter ORGANISM to survive, the next generation will be less like him than they would have been had he been less fit. Though logical from a survival-of-the-organism perspective, his actions have altered the gene pool, the mechanism for inheritance and therefore BOTH the engine AND the "beneficiary" of natural selection.

To be clear, I've warped Dawkins' bird example for my own purposes, but I think it's a serviceable demonstration of the principle. Genes don't WANT to survive, they don't have an emotional PURPOSE to survive - but sometimes they do survive and sometimes they don't, and that is what drives evolution. The primary point in TSG is to point out how even altruistic behavior on the part of an ORGANISM can be linked to the survival of the GENE. It's not a moral treatise.

Gerhard Adam's picture
"...have implied that the bird's INTENT was to draw the predator, which is certainly not the case, nor is it what I meant - the attraction of the predator's attention is a result, not a goal."

There are only two possible ways to interpret this and therein lies the problem.  Either it WAS the bird's intent, or it was the gene's intent.  Consider that if the bird didn't intend it, then it would suggest that the bird's response was "hard-wired" by the genes.  If the latter is being suggested, then it is ascribing intent to the gene.

"There are only two possible ways to interpret this and therein lies the problem. Either it WAS the bird's intent, or it was the gene's intent. Consider that if the bird didn't intend it, then it would suggest that the bird's response was "hard-wired" by the genes. If the latter is being suggested, then it is ascribing intent to the gene."

No, there are not only two possible ways. Neither the bird nor the gene intended that the bird be eaten by the predator. The bird intended to save the other birds. The gene didn't intend anything.

I'm not uncomfortable, however, suggesting that the bird's response is "hard-wired" by the genes. I think there are ample reasons to suppose "nature vs. nurture" applies more to human beings than to other animals, who in the first place are less capable of learning behaviors than humans are.

Gerhard Adam's picture
You're being far too liberal in your interpretation here.  There is no way it has ever been determined what a bird's intention was beyond the act of crying out a warning.  In the second place, the sound of such a bird is quite difficult to locate so it appears that it is hardly a forgone conclusion that the bird would be eaten.

As for hardwiring behavior in the bird.... I think that's simply too far a stretch.  You're attributing far to much to the genes.  There is ample evidence to suggest a significant amount of learning by birds (just to stay with this species).

"It sounds like the meaning of altruism and cooperation are being parsed a bit too precisely."

Didn't you say that earlier?

"the sound of such a bird is quite difficult to locate"

For humans, perhaps.

"You're attributing far to much to the genes. There is ample evidence to suggest a significant amount of learning by birds"

And the learning is handled by pixie dust, instead of genes?

Steve Davis's picture

the attraction of the predator's attention is a result, not a goal. It was intended as an example of a behavior of substantial risk to the actor,
I'm glad you brought up one of Dawkins' favourite themes; animals that give warning calls when a predator is sighted. (I must point out here that the above article concerned DS Wilson, not Dawkins, but if you wish to discuss Dawkins that's fine.) The example you quoted illustrates beautifully a point I made in another article, that biologists should refrain from lecturing on animal behaviour and natural selection until they have spent a considerable period of time in the field. Dawkins' ignorance, and yours in following him blindly, is quite apparent. The last thing a predator wants to do, although it can happen, is to pursue prey that has a good start in escaping. Predators, in general, like the element of surprise. The animal giving the warning call is therefore not increasing its own risk.


As for your other points in support of Dawkins, I've dealt with and disposed of all of them in other articles, do you want me to bore readers to death? 



I think it is safe to assume that predators prefer to pursue prey that does not have a head start escaping. But assuming this alone, the act of giving a warning call can be seen to increase the risk to the individual. In the case of offering a warning the individual will be part of a group who are all attempting to escape and will likely have an equal chance of being individually attack as any other member of the group. If an individual however chooses not to give a warning call, and selfishly escape alone, then the predator will choose to attack one of the unwarned, non escaping members of the group. There is a cost to giving a warning even if the warning doesn't necessarily attract specific attention to the individual; the opportunity cost of not throwing the other members of the group "to the wolves" to buy time for your own escape.

Gerhard Adam's picture

I think this particular example is beginning to be warped quite badly.  In the first place, a warning cry is only an issue, if the prey were hidden and therefore undetectable before such a cry is issued.  If they weren't, then a warning cry is irrelevant and serves a purpose in generating a higher level of awareness to others in the group and potentially escalating the activities to become more chaotic to facilitate an escape.  After all, if you're not hidden, then any number of potential strategies may be workable to provide "cover" for an escape. 

This is the flaw in the example, because clearly the predator already knows where the prey is, so the warning cry does NOT increase anyone's risk because they were all at risk to begin with.  There is also no reason to believe that giving such a cry increases one's risk.



It is not an example I would have personally chosen to demonstrate altruism due to its propensity to be misunderstood, but it is a behavior found widely in nature and one requiring explanation.

As to there being no INCREASED risk to an individual from issuing a warning cry we need to consider two possibilities, the individual either issues a warning or it doesn't. In the case where it doesn't it will be one of a minority escaping, with the majority unaware of the approaching threat. Now as Steve points out "The last thing a predator wants to do, although it can happen, is to pursue prey that has a good start in escaping", so it is extremely likely that the predator will ignore the escaping individuals and attack the unaware members of the group. The risk to the individual if it doesn't issue a warning cry is then very low.

If instead the individual does issue a warning cry all members of the group can be assumed to attempt to escape and despite the predator preferring not to chase escaping targets he no longer has an alternative (He could of course break off the attack, but there is little incentive to do this). The predator will attack the escaping prey, and even if we assume the warning cry doesn't attract particular attention to the individual, the individual is now part of the group the predator will likely attack. A situation that the individual was not in in the case where no warning cry was issued. The risk to the individual is therefore higher in this situation than in the previous.

Now there are two things to note about this. Firstly when the warning cry is issued the chance of the predator catching any prey at all is diminished, but the chance of it catching the individual issuing the warning cry is increased to a level higher than if there was no warning cry. Secondly in this like any other example there are many other factors which could potentially affect the situation, but in the simple case outlined above there is an increased risk to issuing a warning cry.

Gerhard Adam's picture
I understand, but the point is largely irrelevant because it places too much incidental emphasis on the gene, which is largely subject to random chance.

In other words, the gene must first produce an organism, after which there is a considerable range of variable expressions that the gene may experience.  One of the problems, of course, is the "nature" versus "nurture" argument which could completely overrule anything to do with the gene (there is no reason to believe that it is purely a human phenomenon).

Another problem is that far too many genes are similar or even identical across many species, so there is no intrinsic benefit in the trivial traits attributed to them over the much larger conserved processes that operate on the large scale (fundamental structures similar in most species).  Therefore a strong argument can be made that individual species are simply a minor variation in gene expression without improving or neglecting the specific gene in either case.

I realize that there is no "intent" or "emotional" connection to the gene, but that is precisely what makes the argument about a selfish gene so absurd.  Since there can be no intent, we are only examining chemistry and regardless of the survival or reproduction of any organism, the passing on of a set of genes is incidental and not a direct objective of the process.  That may sound odd, because it is these traits that are passed on to subsequent generations, but in truth, the gene is largely along for the ride, rather than directing events in any fashion (even chemically). 

Since the gene must express itself first, there is no way to know whether any particular gene (or it's expression) will be beneficial or not.  I also realize that this is something of a "chicken and egg" problem, which is another reason why I think Dawkin's really went over the edge in describing the gene as the fundamental unit of evolution and using it to illustrate how evolution was basically a means of ensuring the survival of genes into the future.  While this is certainly what happens, it is wrong to attribute anything specific to that result (such as selfish behavior).  As I mentioned before, such an argument could be extended to the absurd point of suggesting that genes only exist as a means of propagating carbon atoms into the future.  While it is certainly true, it is hardly the reason for evolution.

Genes rarely result in explicit expressions, but rather they produce a range of possibilities for which an organism can respond.  While a gene may produce muscles, it doesn't produce a static muscle but rather one that may respond to exercise or neglect.  Similarly, genes don't explicitly connect up the nerves and blood vessels, but rather they provide a general "schematic" against which the cells operate to regulate the process.  Therefore to suggest that the gene itself is responsible at such a specific level seems like a real stretch.

In particular, this search into altruistic behavior is erroneous because the way it is described relies on intent which makes no sense.  Recent predictions from game theory have successfully predicted the behavior of ravens (which some call altruistic) in foraging for food.  It is precisely an independent model like this which supercedes either chemistry or intent that is necessary to understand the workings of cooperation and altruism. 

"I realize that there is no "intent" or "emotional" connection to the gene, but that is precisely what makes the argument about a selfish gene so absurd."

That statement is humorous on numerous levels. Why do you think you have written so many posts about something that you think you understand? Much like this statement, you are failing to put yourself outside of your preconceived notions and looking merely to validate your beliefs. Of course, you are not at fault; it is your genes. They have predisposed us to accepting things that support our beliefs. It makes life happier. Do you see the irony? So you do not have a scientific mind... It is ok. But much like I will not be changing your mind, you cannot change facts.

"I also realize that this is something of a "chicken and egg" problem, which is another reason why I think Dawkin's really went over the edge in describing the gene as the fundamental unit of evolution and using it to illustrate how evolution was basically a means of ensuring the survival of genes into the future."

If indeed you were thinking, you would realize that the egg came before chickens; reptiles predate birds.

"In particular, this search into altruistic behavior is erroneous because the way it is described relies on intent which makes no sense."

Another great statement... I am gathering you just do things randomly to see what happens? You had no intent in writing this post? See this is funny; you think your intent is altruistic, while failing to see that there is intent there.

"Recent predictions from game theory have successfully predicted the behavior of ravens (which some call altruistic) in foraging for food. It is precisely an independent model like this which supercedes either chemistry or intent that is necessary to understand the workings of cooperation and altruism."

Could you explain how proving behavior to be predictable clearly shows that there is no intent? Because I am sort of getting the opposite out of that...

I find it ridiculous to the point of hilarity that you should question Dawkins' qualification to discuss animal behavior. I think you've used the general term "biologist" to give his work a clinical, microscopic feel; in truth, he's a zoologist, with a specialty in animal behavior. As to my following Dawkins blindly, well, I'm not any sort of biologist myself, and am therefore forced to principally weigh two factors in deciding my position on a biological debate: a) credentials, experience, and other trivia like decades of specialized research; b) the argument that I find most logical. If you call that blind faith, I call it more forgivable than were I to fancy myself an expert.

It would certainly be specious to pretend that Dawkins and his arguments were not involved in this post, whether or not they were the central focus, and to be honest it's not important whether or not I'm right about the relative risk of bird behavior (again, I'm not a zoologist); you seem not to concede that ANY behavior found in nature constitutes substantial, deliberate risk to the actor, and I find that totally preposterous. I see human beings engaged in such behavior on a daily basis, and in ANY case of verifiable risk the Dawkins model works for the reasons indicated above. If you don't concede that substantial risk exists in nature, then again, you're not attending to even the most basic anecdotal evidence.

Hank's picture
Appeal to authority arguments don't count for much here - while there aren't many here as well-known as Dawkins, for reasons outside science, the credentials of the research community here are way, way beyond his.   Like Stephen Jay Gould, you can be famous and still be mostly wrong.    Darwin would have a hard time winning an argument among modern world-class biologists too because he knew what he knew - this is the plight of Dawkins as well.

Were he to be less sanctimonious and inflammatory in his approach, Dawkins would get more respect for writing a decent book for its time.  Instead, the people who support him tend to be people outside science who like his cultural agenda.     Which is lovely, we just happen to be a science site.

Hank, I didn't make any appeal to Dawkins' fame, and certainly wasn't arguing that his fame LENT him credibility. I'm also less than certain the comparison to Darwin holds muster - by modern standards, Darwin simply isn't a trained biology professional. He's a self-taught observer with no knowledge of the mechanism which underlies his theory. That doesn't apply to Dawkins - disagree with his theories if you want, but he's a modern, trained professional in his field, not a backwards Victorian with some bright ideas (to downplay my great respect for Charles Darwin).

I can't help but take some umbrage at your use of the phrase "outside science." Professional application of the scientific method doesn't make you a professional in every field of science, any more than I'd be permitted to start a site called Humanities Blogging and remark with confidence on the work of linguists. Anyone on this site, inestimable though its overall credentials may be, who is commenting on an area in which they are not a trained professional with personal research experience, is relying on the authority of their own preferred professionals. These bloggers and commenters might be scientists, but they are just as autodidactic in the field of ethology as I am. Let's try to keep that in mind, and refrain from disdainful suggestions that I agree with Dawkins' science because I'm an uneducated Bright who likes his Youtube videos, shall we?

Gerhard Adam's picture
Another point to consider .... if there was such a thing as a "selfish" gene (not assigning any intent), then there would be no lifeforms above single-celled organisms.  The very nature of multi-celled organisms requires cooperation between the cells and the genes.  Cooperation at the gene level is necessary to ensure sexual reproduction, and since there is no "fighting" over the genes that are passed on to eggs or sperm, it would suggest that the genes just quietly go along with the chemical program and don't exert any influence in an attempt to be passed on to future generations. 

Genes that don't cooperation invariably prevent cell reproduction and don't survive.

Steve Davis's picture
Anyone on this site, inestimable though its overall credentials may be, who is commenting on an area in which they are not a trained professional with personal research experience, is relying on the authority of their own preferred professionals.
That would be one of the sillier points you've made, and it shows why you cling so hard to Dawkins. Some people need an authority figure as a crutch, others can think, reason, analyse for themselves. Try it. It's quite invigorating.

You've been notably unwilling to engage my points instead of just being snotty, which I admit I find somewhat heartening. In this case, there's no need for it, since my remarks are rather self-evident. Certainly everyone is capable of thinking, reasoning, and analyzing for themselves - everyone is equally qualified to apply logic, although not everyone is equally competent at it. But whose data are you analyzing, your own? Are your arguments based exclusively on your own personal field work on bird behavior, your own sociological studies of human interaction, your own laboratory research on the human genome and its permutations? Those can't all be true, and your conclusions are based on the studies of authorities you deem qualified and whose logic you deem sound. Your agency was in choosing your information sources, but you can't get away with portraying me as an ill-informed rube because I admit I get my data from others. It's arrogant, high-handed and patently ridiculous - if your arguments against Dawkins don't quote or paraphrase Robert Ardrey or Peter Kropotkin, I'll eat my hat. Do you believe in the existence of black holes? If so, why? Is it because you reasoned them into being independently, or because you read a book that convinced you?

I repeat: everyone who is not a working professional in a given field is relying to some extent on the word of those who are. To deny that is to reveal yourself as charlatan.

Gerhard Adam's picture
It seems to me that too much of this topic in biology is muddled with confused concepts and assumptions that typically aren't true.

The concept of the "selfish gene" was in direct opposition to an equally implausible scenario of "group selection" wherein it was suggested that evolution favored the group by having the individuals be altruistic and sacrificing themselves for the group if necessary ("for the good of the group").

Both of these ideas are nonsensical.  In the first place biology is, at root, a chemical process.  By whatever means the emergent properties manifest themselves, we cannot ascribe any purpose or intent to anything beyond the fact that it is a repeatable chemical event.  These processes are conserved in the same way that any chemical interaction occurs, which is by attaining a level of stabilities and keeps it going. 

Part of the problem is that in using words like "altruism" and "selfishness" we introduce a stronger meaning and emotional component that is unwarranted.  Certainly we can say that the organism has evolved to act in it's own self-interest without necessarily requiring "selfish" behavior as we think of it.  In addition, it should be clearly understood that there is no singular expression of any organism that does not experience variation.  Therefore we would expect that there would be a range of behaviors that are likely within any group (including some behaviors that might be aberrant and exist on the fringes).

We also know that the gene isn't responsible for "hard-wired" behaviors because that would require a specific and consistent response to a particular phenomenon for all members of the species.  In other words, they would have no choice to be behave in a particular fashion, so it would be impossible to single out the behavior of any member within such a group.

It should also be apparent that animals that are not social are more apt to behave in self-interested ways and are certainly not above exploiting their neighbors.  If we consider animals like ants or bees, then their individual behaviors are very much going to be emphasizing the survival of the group, since they ARE the group.  Therefore to call their actions altruistic is as absurd as suggesting that a lymphocyte is being altruistic when it fights an invader.

However, without embracing the original concepts of "group selection" it should also be clear that a significant biological force (ranging from the basis of multicellular organisms to the groups themselves) is "cooperation".   The use of this concept must also be tempered to distinguish between the active business of cooperation between individuals versus the indifference to others that is also displayed.  There are clear benefits that have accrued to cooperative groups that clearly extend the simplistic interpretation of "selfishness" especially when we see herd/pack behaviors where there is restricted access for reproduction, so there must be a significant benefit to being in the group for members to give up their right to reproduce.

In general, I find that too many assertions are made to imply "global" implications from limited species actions.  If we start from the assumption that whatever we observe is clearly from a species that has achieved some degree of success, then we may begin to understand how their specific behavior has advanced that objective.  However, it is equally clear that no simplistic rendering of "selfishness", "cooperation", or "altruism" can be applied on such a scale.

Steve Davis's picture
Free Radical? Sorry to be the bearer of bad news but you're neither free nor radical. All you are interested in doing is repeating what you've read of selfish gene theory. You're not thinking about the subject matter at all, you've decided that some authority has spoken the last word on evolution. I wrote an article on DS Wilson and in response you wrote a defence of Dawkins. Is that logical? Look at it from this angle. You would never accept the findings of a molecular biologist who did not spend years in a laboratory, yet you're quite happy to accept, and defend to the death, the findings of a zoologist who did not spend years in the field. Is that logical? You say that I base my conclusions on the studies of authorities I deem qualified. Sorry to bear bad news again, but I've criticised just about everyone in the field, because the field is a mess. Now that's radical.

You wrote about one of a series of articles by DS Wilson that you yourself claim is "singling out Richard Dawkins as a target for some pretty heavy broadsides", something that makes you feel "warm and tingly all over". Surely that makes a defense of Dawkins pertinent to the discussion at hand? Particularly when you yourself question his qualifications as an (evolutionary) biologist?

Steve Davis's picture
Luke, I questioned his qualifications in the comments section, not in the article. But apart from that, it makes more sense to criticise my articles on Dawkins rather than this one. If someone wants to criticise my rebuttals of selfish gene theory that's fine, but if they just want to repeat the theory then as far as I'm concerned they can go write their own article and not waste my time, and the time of those readers who want the discussion to progress.

What I was referring to was your decision to include the following qoute in the article proper; “Richard has become unaccountable, in part by becoming a public icon. That disqualifies him as a spokesperson for science.” I took your inclusion, and the agreeing comments following it, to indicate your acceptance of this statement.

But yes, this does not furthur the discussion. If you are truly interested in discussion of your own rebuttals to selfish gene theory, please let me know on which of your articles you think most accurately summarizes them and I will be happy to comment there.

This discussion is a wonderful example of how simple semantic confusion, along with a good dose of the naturalistic fallacy, makes for completely pointless conversation. Note the abrupt descent into ad hominem attacks, coupled with half-understood examples from outside any of the speakers' area of expertise.
Actually, you know, this is a good example of why cooperation, where it exists, needs to be explained. Animals within a group have generally competing interests, even those animals who are sort of discussing science online, and the fact that cooperation works well, once it exists, does not explain how it arises in the first place.
I also enjoy the irony of a science blogger talking about how Dawkins is disqualified from talking about science because he's a public figure. Is there a cooperation-centric view of cognitive dissonance?

Gerhard Adam's picture
"..the fact that cooperation works well, once it exists, does not explain how it arises in the first place."

Try the iterated prisoner's dilemma and "tit for tat".  There is ample evidence in game theory that cooperation can readily arise from purely selfish interests.

Gerhard Adam's picture
"Animals within a group have generally competing interests ..."

Beyond the competition for mates, what is this based on?  Most social animals have groups that supercede the ability of most individuals to survive without them (that's why they belong to groups).  Therefore, if the group conveys a more significant survival advantage, then those that are cooperative will tend to be better represented in the gene pool.

If a group has a stable existence, it is virtually impossible for outside males or females to interact and reproduce.  Therefore, individual existence or competition has little chance of supplanting a successful group.  This applies to humans, horses, wolves, etc.  In every instance you see very little survival potential in isolated individuals.

Steve Davis's picture
Anonymity is a wonderful thing isn't it? It allows you to indulge in the most inane, lazy, unthinking comments. Some readers are worried by a quote I took from a Wilson article, but are too tired to go to his blog to get the full context. And in this comment above I'm chastised for not explaining the origin of cooperation. If the courageous commenter wants to go back through my articles and the discussions they generated the answer can be found. It really is amazing. Some people expect the mysteries of the universe to be explained to them in a few hundred words. And as for actually working out a few answers for themselves, of which most are quite capable, that's not even considered.   

Rick Ryals's picture
Funny thing happened. I got this email saying that I'd received a message via my contact form, but when I click on it, I get a message telling me that I'm not authorized to access that page. So my email is rickryals at gmail.com if somebody still wants to reach me.

logicman's picture
Richard:  it was probably a correct email when sent, but if the comment was pure spam it would have been deleted by an admin - in which case you get a 'not authorised' error message.  You get the same if you click on a 'what people are saying' link just as the comment is being deleted.  It's not censorship, its a web experience enhancement strategy.  :)

Applying sociobiological thinking to human societies is a perilous undertaking. But the sociobiology of the New World Order must be revealed!!

Gerhard Adam's picture
Actually it's only perilous when neither social understanding or biology is actually involved.  In too many cases, sociobiology is used as a rationale by which social and political agendas are advanced to try to provide a veneer of scientific validity to blatant power grabs.

Biology applies to human societies as it does to any other animal, and many of the ideas of sociobiology are also applicable, provided they are not treated as a engineering blueprint to try and advance any particular cause.

I don't think it's realistic to say, "Sociobiology is legitimate science, but don't try to apply it to anything -- exempt it from engineering applications."

If we know how something works, somebody will figure out how to apply that knowledge to advancing their own self interests.

Gerhard Adam's picture
.... and that's where the problems begin.  Sociobiology can be used to advance understanding, but it would be difficult if not impossible to think that humans (since that's what I was referring to) are amenable to social engineering.  It's no different than understanding evolution.  Such an understanding doesn't mean that we possess the means to direct it.

Such systems are fundamentally chaotic, so regardless of what you think you understand, it is extremely unlikely that the results would mirror something as simply as an engineering problem would. 

logicman's picture
Applied science is not the same as applied technology.
Just because somebody's got an implementation of something
doesn't mean they've got a theory of it.

Gerald Gazdar.

Pff, a lot ado about absolutely nothing. Cooperation and the selection of groups may be more important in the evolution of life than kin and gene selectionists think. But even the selection of cooperating groups is a form of gene selectionism. It doesn't matter whether single genes or whole genepools are selected: in both cases, it are, ultimately, genes that benefit. Not organisms. They are mere and mortal instruments of genes and genepools. People used to be upset by evolutionism because it falsified the idea that man is the purpose of the creation. Now, they are upset by genecentrism because it falsifies the idea that the organism is a purpose in itself. It's not: it's an instrument of the genes. Plus ça change...

Gerhard Adam's picture
Except that it's not.  Might as well argue that genes are simply the vehicle that carbon uses to perpetuate itself through the cosmos.

Yes, and we used to think that it was the other way around: that carbon was the vehicle, used by the genes, to perpetuate themselves. We didn't of course. It's not possible to look at both entities this way, because genes and carbon molecules are not two separate spheres, like genes (potentially immortal - the information encoded in them) and organisms (mortal instruments of genes) are.

Genes are made of carbon (and other) molecules. You can't say that genes are an instrument of carbon, just like you can't say that your body is an instrument of your internal organs. But you cán say that your mortal body is an instrument of the genes that it passes on to the next generation, if you 'reproduce' (you don't reproduce yourself of course: you pass genes on to the next generation, just like the sun doesn't really 'set', and no, it are not 'your' genes, it are they who possess you).

It is of course possible to 'rebel' against your 'selfish' genes. As Dawkins always says. We're not stupid, blind robots, like other animals. But look at him. He just doesn't stop. Why does he attack religions the way he does? Probably because he craves for more power, influence and status. Women like powerful men. Dawkins is currently living with his third wife. He's a serial polygamist. If his genes could talk, they would say: good boy (although they wouldn't like the contraceptives he and she use). Yes, it's possible to rebel. But that requires in the first place that you know what drives you (we're often not aware of what drives us). And it's often not a pleasant option.

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