Let’s get real about Selfish Genes, Group Selection and Altruism. Towards a less impoverished view of humanity.


Last month journalist David Dobbs wrote an article that blew up in evolution circles: “Die, selfish gene, die“. It’s a takedown of Richard Dawkins’s excellent 1976 bestseller The Selfish Gene, which consistently and deservedly ranks as one of the best pop science books ever written. As an example of its influence, it’s where the word meme comes from. Dawkins uses the “selfish gene” as a powerful metaphor to explain genetics and evolution. Genes (only as a metaphor, since obviously genes don’t think) do their best to selfishly replicate into subsequent generations. Dobbs argued the selfish gene metaphor was out of date, since we now know much of the action lies in gene regulation, interaction and expression. In short, we have social genes. The title was provocative enough that Dawkins himself replied. The nerd highlight came when ECOLOGIST HULK brought the awesome by debating Graham Coop live on twitter. And while I think Dobbs generally didn’t fair that well, he was game and it ended amicably enough.

Yet I think The Selfish Gene itself (the book) got off too easy. Though not for the reasons Dobbs wrote about. Everyone involved in the debate, Dobbs included, was careful to note that of course everyone knows selfish genes don’t imply selfish people. As Dawkins himself has said many times. But let’s get real. Dawkins’s book influenced the public into making exactly that superficial mistake. To be fair, the real issue was a larger 1970’s social science movement portraying humans as primarily selfish. But Dawkins’s book was part of this. And it led to an impoverished view of humanity from which we’re still recovering. Writing about how selfish genes tie to social science unfortunately made this a long post. In particular the middle section bogs down a bit with a historical crash course on group selection and the evolution of altruism. But if you stick with it (or skip down to the second smiley face many paragraphs below smiley-face-smile), we’ll come back to this larger theme at the end. And even touch a bit on business culture.

Why is the evolution of altruism a puzzle?

To a biologist, altruism is defined as reducing your own fitness (chance of reproducing) to help another. But if populations evolve by favoring those who reproduce more successfully, then at first glance altruistic creatures shouldn’t evolve. They would be outcompeted by those who are not. And yet altruism clearly has evolved. So it’s a puzzle. What are the exact conditions under which altruistic genes get passed down?

To be clear, don’t make the common mistake of confusing altruism with cooperation. Cooperation is NOT a puzzle. If a school of fish avoids predators by swimming together, this is not altruism. Each individual benefits from the cooperation. Most social activity in animals can be explained by simple cooperation. The evolution of altruism is more subtle.

To help illustrate modern views on this topic, I’ll use the people below as stand in representatives: 1. Steven Pinker – mainstream on evolution, and fine with human altruism (under the right conditions). 2. Richard Dawkins – also mainstream on evolution, but more skeptical of human altruism. 3. David Sloan Wilson – non-mainstream on evolution, but similar to Pinker in being fine with human altruism (though his conditions differ from Pinker).


Deep dive on selfish genes, group selection, and the evolution of altruism

In last month’s selifsh gene dust up Dawkins quoted himself saying “Another good alternative to The Selfish Gene would have been The Cooperative Gene. It sounds paradoxically opposite, but a central part of the book argues for a form of cooperation among self-interested genes.” Dawkins added this to the introduction of the 30th anniversary edition of his book precisely because so many people claimed he was saying selfish genes implied selfish people. And he was getting annoyed by how he was misunderstood. So I’m not in any way original in accusing him. Nonetheless. In the quote above he’s talking about genes inside a single organism. Later he talks about cooperation between organisms (altruism) within the gene-centric framework. This reflects Dawkins’s considered scientific position. Here’s Dawkins doing a TV special in 1986 making his case. It’s worth a look, for the amazing 1980’s big hair if nothing else. In it Dawkins says cooperation and altruism are just fine. But in his book and later TV special he’s very passive/aggressive. Trashing altruism while claiming he’s being balanced. He’ll say altruism is so rare we can ignore it, then leave in a tiny caveat. Then claim selfish genes imply selfish people. But pull back to “cooperative genes” when challenged. Here’s a quote from Dawkins’s book via David Sloan Wilson illustrating the style:

I shall argue that a predominant quality to be expected in a successful gene is ruthless selfishness. This gene selfishness will usually give rise to selfishness in individual behavior. However, as we shall see, there are special circumstances in which a gene can achieve its own selfish goals best by fostering a limited form of altruism at the level of individual animals. ‘Special’ and ‘limited’ are important words in the last sentence. Much as we might wish to believe otherwise, universal love and the welfare of the species as a whole are concepts that simply do not make evolutionary sense. Let us try to teach generosity and altruism, because we are born selfish. Let us understand what our own selfish genes are up to, because we may then at least have a chance to upset their designs, something that no other species has ever aspired to do.

As David Sloan Wilson points out, people are also born with an evolved capacity for altruism. Not just selfishness. Dawkins goes too far when he claims we are “born selfish” and need to be taught altruism (overriding our apparently one-sided instincts). The misunderstanding that selfish genes make selfish people is not a misreading of Dawkins. It’s right there in his original and very carefully worded text. 

Why the severe skepticism of altruism? Thirty years ago George C. Williams had just published his book Adaptation and Natural Selection, following W. D. Hamilton in establishing Kin Selection as a key evolutionary explanation for altruism. Kin selection means helping your closely related kin, and their closely related genes. In a word, nepotism. Parents altruistically help children. Uncles and Aunts help cousins. Around the same time, Robert Trivers published his paper on Reciprocal altruism. Reciprocal altruism means scratch my back and I’ll scratch yours. Conditional tit-for-tat altruism can evolve as a result. Combined together, kin selection and reciprocal altruism provided a rigorous, gene-centric mathematical framework for the evolution of altruism that’s stood the test of time. This framework came to be called Inclusive Fitness. It completely destroyed the earlier (and honestly not mathematically rigorous) work of people like Vero C. Wynne-Edwards, who had been arguing that altruism evolved via a naive form of group selection. What’s important here is both kin selection and reciprocal altruism require special conditions, making altruism theoretically rare. And in his zeal to debunk the muddled thinking of old fools like Wynne-Edwards, young Dawkins overplayed altruism’s rarity. This was a career defining move. It deservedly put him on the map. He hasn’t looked back. And like a retired general, he enjoys refighting battles already won, continuing to bash altruism and group selection with gusto to this day. We can with only slight exaggeration (ok large exaggeration) place Richard Dawkins in the “altruism is so rare it I’m not sure it even exists, except for bees” school of thought. His response to accusations of overplaying the rarity of altruism is to question if people have even read his book, which of course has a chapter precisely on that topic (using bees as an example). I’ve read his book several times. It’s brilliant. But regarding altruism, he passively/aggressively says in meticulous and precise language exactly what he intends.

Moving forward in time David Sloan Wilson and others recast the old naive group selection into a modern form compatible with the gene centered approach. This “group selection 2.0”, now labelled multilevel selection theory, ultimately became compatible with inclusive fitness. The theories are mathematically related via the Price Equation. Technically, multilevel selection partitions genetic fitness by population structure (groups). While inclusive fitness partitions fitness by relatedness (kin). Think of this as the groupish lens versus the relatedness lens. If you know physics, a good analogy is how Hamiltonian Mechanics is mathematically equivalent to Newtonian Mechanics, but uses a different formulation. You pick your formulation based on the problem at hand. David Queller compares inclusive fitness and multilevel selection theory to two different languages: say English and Spanish. While English can be used to solve all evolution problems, some are better understood in Spanish. If you have a lot of free time, David Sloan Wilson walks through the details in his 13 part series on the topic here. But to be fair, group selection even in modern form remains disparaged. Though quite publishable. Until 2012 that is. When Martin Nowak and E. O. Wilson published a highly controversial paper in Nature saying group selection was NOT compatible with existing theory (trumping the David Sloan Wilson angle). It was better and a replacement. You have to understand E. O. Wilson is a giant in the field of evolution, though at age 84 near the end of his career. So people got extremely annoyed.

Steven Pinker is a good example of the mainstream response to Nowak and E. O. Wilson’s paper. He wrote a widely commented on essay The False Allure of Group Selection, where he says “Group Selection has no useful role to play in psychology or social science.” In his usual careful manner, Pinker then talks about altruism arguing that that “The huge literature on the evolution of cooperation in humans has done quite well by applying the two gene-level explanations for altruism from evolutionary biology, nepotism and reciprocity, each with a few twists entailed by the complexity of human cognition.” That is to say, Pinker thinks the David Sloan Wilson multilevel selection formulation and the newer Nowak/E.O. Wilson variant add nothing useful to the standard inclusive fitness model. Except confusion. But in contrast to Dawkins, Pinker is far less skeptical of selfish genes evolving (contingently) altruistic humans via the accepted pathway of inclusive fitness. Needless to say, I’m a huge fan of Pinker. He’s gracious with people he disagrees with, and is fine with human altriusm/social cooperation evolving via the standard model.

Let’s summarize views in the table below. I include our three main protagonists mentioned at top, plus a couple others. Blue color shows mainstream thinkers. Note E.O. Wilson is not blue, but has an asterisk because he’s had a brilliant and distinguished career. It’s only his group selection views which are non-mainstream. In the table, the 1960’s version of Naive Group Selection from Wynne-Edwards is right out. Inclusive Fitness is fine with everyone, except for E.O. Wilson. And the reformulation of group selection as multilevel selection is a mixed bag. Dawkins thinks it’s completely wrong. Pinker finds it equivalent but adds nothing useful to our understanding, and hence is also a no. Queller and David Sloan Wilson find it equivalent and useful. This is obviously my view for what it’s worth. I’m clearly a David Sloan Wilson partisan, so read accordingly. I didn’t include a column for the Nowak/E.O.Wilson model since it’s uniformly disliked, even by David Sloan Wilson.


So why does this rather arcane debate on the evolution of altruism matter to the general public? (people skipping the gory details above can resume reading here smiley-face-smile)

The great tragedy of this debate is arguments over the mechanism for how altruism evolved have spilled over into claims about how altruistic humans are in the first placeOne of the reasons I’ve done so many posts on Jonathan Haidt’s book The Righteous Mind is because of his focus on in group/out group dynamics as being at the very core of human nature. This was due to the influence of David Sloan Wilson. But to be clear, Haidt could have just as easily come to these conclusions using a more standard Pinker-style inclusive fitness framework. The standard inclusive fitness model emphasizes relatedness, so if we want to explain how human groups go to war, they go to war as fictive kin groups. Soldiers are (fictive) brothers-in-arms. Haidt’s Righteous Mind thesis could have been built on an evolutionary foundation of clans, reciprocal altruism and fictive kin groups.

In fact, given the recent renewed stink surrounding group selection, it would probably be in social scientist’s self interest to avoid it. For example a second paper by E. O. Wilson and Martin Nowak has been utterly ignored, as I think people are tired of debunking it. Group Selection remains disdained, even the consilient David Sloan Wilson multilevel selection version. Yet I think the David Sloan Wilson approach adds something to the mix. This formulation allows human groupishness via more abstract non-kin mechanisms such as ideology and religion. I’ve done a previous post on religion, Dawkins and group selection, so won’t go into that here. But religion and human ultrasociality are an angle where someone like Peter Turchin also finds Pinker’s inclusive fitness views insufficient. Though Pinker and most biologists would obviously disagree. My pet peeve with Dawkins’s otherwise great book is NOT that he uses the standard model. That’s totally legit. Rather it’s that he’s so down on human altruism and human cooperation. He has too narrow a view of our quirky and unique humanity.

There are others besides Haidt influenced by David Sloan Wilson. Adam Grant recently published a business book Give and Take: A Revolutionary Approach to Success. Grant is at the Wharton Business School at the University of Pennsylvania. Here’s my own (hopefully accurate) transcription of an interview with David Sloan Wilson:

Wharton students stop by my office for career advice. And one of the conversations that comes up most frequently is a variation of “I really want to help others and make a difference, so I’m going to be as selfish as possible and maximize my wealth for the next 30 years. And then I can start giving back.” And you hear that story often enough, and you start to wonder. What if that’s backward? What if some of the most successful people started giving first and then succeeded later?

Grant’s book is an extended argument for exactly this point. If you are careful on who you work with, a give-and-take cooperative approach to business is more successful than being narrowly selfish. You don’t need to wait until retirement to give back. As they discuss business and the social sciences in more general terms, David Sloan Wilson says thinking of people as exclusively selfish is an “impoverished view.” Exactly! Both Grant and Wilson agree narrow human selfishness is still a common assumption, especially in business.

Contrast Adam Grant’s book to Francis Fukuyama’s recent The Origins of Political Order. I loved this book. It has Arnold Toynbee level ambitions, attempting to outline world history and the growth of civilizations. And in it Fukuyama properly cites kin selection and reciprocal altruism as standard theoretical reasons for the evolution of human altruism and cooperation. But he has unfortunately picked up on the narrow Dawkins take rather than the more generous Pinker one. He remains far too skeptical of in group/out group dynamics and cooperation as being central to civilization and humanity. His book suffers accordingly. This is the unfortunate legacy of The Selfish Gene, or more generously of improper readings of that book, and the movement of which it was a part.

The selfish genes/selfish humans meme is part of a larger history of viewing people as a one sided and selfish. And this view diminishes us all. It makes companies stack rank employees thinking this will bring out their best. It makes us scoff at religion. It makes us constantly underestimate the seductive tribal power of groups. It leaves us blind to the wellsprings of our passion for sports, which simultaneously bring out our very best and worst. It makes us believe greed is good. As an enthusiastic young reader of Dawkins’s book, I went down that very path myself. It’s time to move on. Regardless of the mechanism by which human altruism and ultrasociality evolved, it is part of us, and we should find ways to harness it to our mutual benefit.

By Nathan Taylor

I blog at http://praxtime.com on tech trends and the near future. I'm on twitter as @ntaylor963.


    1. Radiolab is the best. Such good story tellers. The Price of Altruism which they talk about in that episode is a great book by the way, read it last year. What’s funny is in retrospect I had a good story in this post. The last paragraph is the key. Should have built entire post around it, and cut post by 2/3. Story is: Young Ohio boy misunderstands Dawkins, thinks greed is good, only to learn it’s not, wants to share his hard won lesson. Would have made a much better and much shorter post. Instead I rambled on with too much science history (which I find absolutely fascinating by the way). Learning how to write is learning the basics over and over and over. When in doubt, more darlings must be killed. Faulkner quote. Need to fire my editor. 🙂

      1. A keyword in my Internet writing these days is PII, pronounced “PIE.” As in personally identifiable information. I try to cut PII when it is not essential to the post.

        Saber [google login broken]

  1. Good post but overly winding and wordy.
    You focus too much on the people, the dramatic interaction between them, and too little on the ideas.

    I get this is for quite erudite people, but it would still be nice to keep the prerequisite reading to a minimum.

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