Three years late, Scientific American is reacting to a paper on evolution of cooperation  with a hint that something is rotten in the state of Denmark. This is what my wife used to call an FGIO, a faint glimpse into the obvious. The paper was by interlopers from physics, mathematically adept senior luminaries. The message of their mathematics was that if you put together a group of selfish individuals what evolves is (I’m sure you’ll be shocked to hear this) a selfish community. A non-communal community. The authors use the colorful word “extortion” to describe the dysfunctional social relations.
The reason this is important is that the standard evolutionary theory of cooperation says that global cooperation can emerge from behaviors that are individually selfish. It can’t.
(We may be tempted to dismiss the whole topic as esoteric or peripheral, but in fact it pokes a hole in a framework of evolutionary thinking that has become sclerotic, and is overdue for rethinking from the ground up. At stake is the way that we think about evolution, and by extension, the way we understand purposes and mechanisms in all of biology. Aging is one particular case in point.)
Diverse forms of Altruism in Nature
Human soldiers offer up their lives all too readily in wars of questionable legitimacy. Amoebas stop competing among themselves and form a “fruiting body” that behaves like a single organism. Worker bees spend their entire lives serving a colony whose only reproduction is by a queen bee that may not even be a blood relation to the worker. And yeast cells will commit suicide to digest themselves as food for their cousins when they detect that the local region has run out of food. Fish of two different species work together to hunt more effectively.
When we look at the biosphere, the ubiquity of grand alliances is striking. Everywhere, we see extreme examples of individuals sacrificing their lives and their legacy for the good of the community. Not infrequently, the sacrifice is complete, in that the individual leaves no offspring at all; and yet the benefit to the group seems modest and indirect. What theory could account for the mismatch between individual cost and communal benefit?
In stark contrast to this broad observation, we have the dominant version of evolutionary theory that purports to explain how all this came about. This is the “selfish gene” model. The great majority of papers published in evolution today are still framed within this model. This mode of thinking has infected and distorted scientific understanding of aging, which is why I became interested in the problem originally, and why this subject belongs in an aging blog.
Of course, adherents of the dominant school feel responsible to provide some account of cooperation, but they look at the world with tunnel vision. There are just two narrow windows through which the broad spectrum of cooperative phenomena are permitted to be viewed. One is kin selection: a gene that causes altruistic behavior in its bearer may survive if the altruism is directed toward close relatives who are likely to have copies of the same gene. Second is Tit for Tat reciprocal altruism. Higher animals that recognize each other as individuals may learn who they can trust, and on the basis of a network of pairings, broad cooperation can spread through a group. It is reciprocal altruism that has been called into question by the 2012 paper referenced above, by William Press and Freeman Dyson.
I find it curious that both these windows were opened by William D Hamilton, early in his career, before Hamilton became a convert to group selection. He died untimely of malaria after a trip to Africa observing and collectins specimens in 2000.
Kin selection is understood in terms of Hamilton’s Rule, which says C < rB. In order for an altruistic behavior to evolve, the cost to the altruist (C) must be less than the benefit to the recipient (B) multiplied by their mutual relatedness (r). Relatedness r is ½ between siblings or for parent and child, and it goes rapidly down from there. “I wouldn’t jump into the river to save my brother,” quipped J.B.S. Haldane, “but I’d lay down my life for two brothers or eight cousins.”
Reciprocal altruism is understood in terms of the “Tit for Tat” strategy, made famous in a paper by Hamilton with Robert Axelrod in 1981. Based on a simulated tournament implemented with the crude computer facilities available at the time, Hamilton and Axelrod concluded that the most successful selfish strategy could be a basis for global cooperation. This was validation of the “invisible hand” that Adam Smith had described two centuries earlier. Emergent order in a system where everyone was only looking after himself. Emergent cooperation in a system where everyone is behaving in a manner wisely selfish.
“What looks like cooperation is really selfishness” is a theme that pervades the evolutionary literature, in guises both direct and quite subtle. It was the basis of the sociobiology movement a generation ago, and it thrives today in fields as diverse as evolutionary ecology and the medical biology of aging.
Origin of the Current Crisis
This way of thinking took it on the chin in 2012 when Dyson and Press published their paper, demonstrating a form of selfishness that Axelrod and Hamilton had not considered. Their diabolical strategy took evolutionary learning into account, and gradually trained opponents over time to be submissive, using a purely mathematical form of “extortion”.
This is but the latest scream in the chorus of voices telling us that the “selfish gene” version of evolutionary theory doesn’t work. The strength of cooperative groups is that they win out in competition against other cooperative groups. The idea that the power of a cooperative group is just a lucky side-effect of selfish behavior self-organizing never was credible, and finally it may be biting the dust. The truth is that the “selfish gene” version of evolution has outlived its usefulness. It is long past due for an overhaul. Evolution does not work “one gene at a time”. Cooperative relationships have evolved not by accident, but because cooperative groups are very successful competitors, and they can spread and take over areas where individuals are unorganized and selfish.
What does Selfish Gene Theory Leave Out?
Now that we see the truth, we can go back and ask about the Selfish Gene theory, what is the reasoning that supports it, and what is the flaw in that reasoning?
There are two lines of reasoning relied on by Selfish Gene adherents:
- All mutations originally appear in just one individual. If a mutation for altruism appeared in one individual surrounded by selfish behavior everywhere, that individual would be left in the dust. Everyone around her would accept her generosity, thank you very much, and then use the advantage she had given them to out-compete her and leave her behind. Genes for altruism could never spread in a selfish community.
- Conversely, in a community where everyone was cooperating, if a gene for selfishness arose in one individual, she would be able to benefit from the help given by all the others around her without herself bearing any of the cost of helping others. Her star would rise, her progeny would prosper, and her selfish gene would spread through the community, poisoning it from the inside out.
This is a compelling perspective, and not obviously wrong. However, it does not describe the world that we see. It predicts that cooperation should be rare in nature, when in fact we find cooperation wherever we look, and not just among close kin.
So we know that the argument for the Selfish Gene must have a catch somewhere…what can the problem be?
In my theoretical work, I have offered a general answer to this question that I derived from the work of Michael Gilpin in the 1970s. It goes like this:
All animals depend on an ecosystem. There is a common reservoir of food in the form of living plants or animals on the next trophic level down. The community of animals dependent on the same food stock is tied together in their fate by a need to conserve that resource. It is all too easy to eat everything in sight, to deplete the food species and to use all that extra food energy to reproduce like crazy. But the collective consequence of this behavior is rapid disaster. Once the food species is gone, it takes a long time to grow back. The next generation will starve, and the ecosystem will take a long time to recover, if it ever does.
There is a powerful tendency for population dynamics to fluctuate wildly, leading to extinction. The extinction is rapid, and can occur in a single generation. Therefore, it constitutes a very potent force of natural selection, one that can easily counterbalance selection for pure selfishness, and defeat the Selfish Gene.
So when Selfish Gene adherents think about a gene for selfishness taking over a community, what they’re leaving out is that the community would likely be destroyed in the process–the community would die out before the selfish gene could become dominant. This is true of the easiest and most powerful form of selfishness, which is overconsumption. And computer simulations show that once this most powerful form of selfishness is tamed, it ties communities together in a way that makes it easier for other forms of cooperation to evolve.
This is the mother of all cooperation, the glue that binds communities and makes selfishness a dead end. Selection for population homeostasis opens a door that permits all other forms of cooperation to evolve.
The Bottom Line
Let me conclude with this bold statement of my radical new thesis: If it looks like cooperation, it probably is.
> “The strength of cooperative groups is that they win out in competition against other cooperative groups.”
You can also add “Fortuna” here. They also win out “against” nature. For example, after a disaster, helping others will increase fitness. (conserve the reproduction energy spent)
Btw, doesn’t farming wheat and cows already invalidate this narrow view of the selfish gene? =)
I mean, if even farmers can invalidate one’s theory, and do so without even uttering a word for argument… There must be a flaw in one’s theory 😉 Although, I think we can expand this “narrow selfish gene” theory to account for this..
For instance, what is the difference between a bundle of thousands and thousands of genes evolving in an individual and mostly the same genes evolving somewhere else. We call one an organism but the whole population can also be seen as a bigger organism. In fact 1 little (selfish) gene has ZERO % kinship with most if not ALL its neighbors, so it should be much harder to explain than population selection. In other words, populations should have even more diversity than we see in genomes themselves.
Otherwise, why not draw the line of “individuals” at the stem cell level? What about that cooperation? Cancer is another example of what you’re saying there, selfishness being selected out. True, there are costs to be surmounted but, that is true at EVERY reproduction checkpoint. What’s the agenda behind stopping at the “individual”? They can’t deny multicellular organisms evolved, or can they? (I bet they could) What about ribosomes suspiciously folding all those proteins for the rest of the cell? Nope. Can’t have that. Oh wait, can’t evolve anything without that.. or can we, why aren’t we still living in an RNA world?
Even that formula there is wrong, because sex is already cooperation at a lower level: “Here, I’ll nurture half of your genes if you nurture half of mine”. Carrying all the eggs in one bag of skin (I mean the “individual”, probably a Selfish Gene adherent) is the best evolution can do? WOW. Really? O.o” Mwahaha
> There are two lines of reasoning relied on by Selfish Gene adherents:
Point 1 is also easily invalidated by Game Theory:
> “and then use the advantage she had given them to out-compete her and leave her behind. ”
But then again, it would give an advantage against the rest of the selfish community to have such help. Thus a niche for a form of cooperation to evolve… more like vampirism but.. =) I mean, “parasitism”. But not so different from the ecosystem example you mention — or the farming one I use — if it fully drains the host, it would lose that edge. Case in point: Scumbag politicians/businessmen.. perhaps sending the “next trophic level down” to “rackets of questionable legitimacy”, heh? 😉
It’s known that humans don’t usually think more than 1 or 2 levels deep on the Game tree like this. Selfish Gene “adherents” can’t even think 1 ahead it seems, LOL.
> Conversely, in a community where everyone was cooperating, if a gene for selfishness arose
Point 2 can be invalidated similarly. Protecting that edge, specially when we add Fortuna as I said earlier. Can’t “adherents” conceive of more realistic scenarios where there isn’t simply 2 individuals or even 2 populations competing? Entropy is not to be trifled with and they simple shrug it off like it’s an idealized sports game (not even a team sport! they probably grew up when chess was still a badge of ultimate complexity and brainzzz).
In fact you already wrote about the bacteria that police against free riders, right? Perhaps the deeply ingrained justice instinct is even more ingrained than we think, it’s probably ancient… unicellularly ancient.
The thing about about selfish gene theory is that it is Neoliberalism applied to biology. The same theory of selfishness was being applied to economics and politics at the same time as Dawkins was cooking up the selfish gene, and to a large extent it was done in the same environment – in elitist universities tasked with the job of rationalising elitism.
As we’ve seen in economics, in the form of “free markets”, it leads to disaster on a large scale as banks given free reign to create money via debt will keep doing so until the financial system collapses. Wealthy people given the chance to opt out of making social contributions, through e. tax, will gladly do so. And so on.
Dawkins is simply part of a wider movement in British intellectual life – with roots in the thought of Jeremy Bentham, John Stuart Mill and Herbert Spencer. Survival of the ruling classes depends on them finding rationalisations for their selfishness.
As Friedrich Engels wrote almost 150 years ago:
How do you guys quote one another? Are there  tags? I’m using email notation.
Funny though, it shows the quote on the email notification but not here..
Josh – the reply form for your latest post does not seem to work.
Yes – you’re the second person to complain, but you seem to have gotten your comment through. Is there a trick?
It seems like a WordPress thing. Probably depending on arbitrary words it seems to misclassify some posts as spam.
Another possibility is if our computers or browsers are infected with malware that tries to post commercials/spam (I caught one right on the act!) the site may filter our IPs for a while after.
The trick would be for WordPress to improve this stuff. Maybe Josh has access to some configuration he can tweak tho, but probably not much or he would get swamped by bots. The only trick seems to be typing stuff on an editor before posting here?
I’ve given up posting or replying on several occasions already. =( Sometimes it seems it won’t even present the post to Josh to moderate.
Sometimes (like now!) after I try once and it doesnt show up, the page says: “Click back and type in the correct password. (Spam Free WordPress)” — The password doesnt change though! And nothing seems to happen (maybe it detected I’m posting a “duplicate”) I try refreshing the page for a new password and sometimes it works, sometimes the post doesnt show up again.
If Josh sees duplicates, that’s probably the cause.. people trying to circumvent the spam protection which is misclassifying their posts and wasting their times.
All I can say is that “a faint glimpse into the obvious” is a perfect way of describing this. The ultimate goal is not to pass on a specific set of genes at any cost to the community, but to contribute to the survival of said community. After all, no indvidual would be able to survive without its community and ecosystem, no matter how perfectly adapted or powerful in terms of selection its genes make it.
With the extinction of 99.999% of all species that ever lived (including their fittest individuals), then what you are saying is it is not survival of the fittest individual, it is survival of the species, the selfish gene leading to extinction.
My point exactly. If we consider reproduction to ultimately be the goal of life (whether to pass on one set of genes or to continue the species), we have to take into account that the vast majority of species is unable to reproduce on their own. Exclusively selfish genes therefore do lead to extinction. Not to say that any degree of selfishness has the same effect though.
Evolution seeks to maximise gene replication within the local reproducing group in the context of the environment. It does so by distributing genes among vehicles (organisms) which are each motivated to maximize their inclusive fitness. Both genes and individuals are free to compete or cooperate, but they are always selfish in the sense that genuine altruism (compromising the goal of maximizing inclusive fitness) is impossible. Dawkins got this right 40 years ago–the ‘selfish gene’ is an excellent metaphor.
Yes – you’ve accurately stated what is still the mainstream view. I ask you to look around you at the biosphere and tell me – does this theory describe all that you see?