The Selfless Gene: Evolutionary Theory Reconsidered

Harvard University myrmecologist—that’s right, he studies ants—Edward O. Wilson has created a deep schism in the field of evolutionary biology. His latest book, The Social Conquest of Earth [1], released this April, advances a theory of group selection, at odds with the field’s former consensus and, indeed, Wilson’s own fifty-year corpus of Pulitzer Prize-winning books and groundbreaking research. Opposing Wilson stands Oxford University evolutionary biologist Richard Dawkins, famous (before his radical atheist days) for his theory of “inclusive fitness,” also known as kin selection, which he proposed in his landmark 1976 book The Selfish Gene. Now, the two are duking it out in an article war that started with Dawkins’s biting review of Wilson’s recent traitorous book.

After thirty years of relative unanimity, the field of sociobiology is split between its two fathers, and the resulting debate has profound implications for our understanding of human nature. From Dawkins’s perspective, human society is nothing more than an accumulation of selfish individuals. From Wilson’s, it’s a sea of ambivalent individuals perpetually torn between serving self and society. I daresay we hope the latter to be the case.

Both theorists agree on the concept of natural selection (evolution) as an explanation for biological diversity, but they disagree on the level at which it acts. Dawkins, as per the tradition of sociobiology, pioneered the idea that it works at the smallest possible level of selection: that of genes. His so-called “Selfish gene theory” has become a cornerstone of modern evolutionary biology, though many contest its hazy empirical grounding. Though the term “selfish” sounds cold and individualistic, selfish gene theory does provide an explanation for group solidarity and altruistic behavior known as “kin selection theory,” by which individuals act to benefit others who share their genes. To explain the cost/benefit calculus that altruistic acts entail, Dawkins proposes formula suggested by evolutionary theorist W.D. Hamilton years ago: r[relatedness of the actor to the recipient] x B[benefit to the recipient] > C[cost to actor]. By this logic, individuals––who, after all, are only their genes––act in ways that favor the propagation of their genes, even if those genes lie in other, related individuals. For Dawkins, once we’ve accepted that genes are the sole vehicle of genetic inheritance, they must also be the sole target of natural selection.

But kin selection theory is insufficient for Wilson, who has observed altruistic animal societies his entire life––especially those of ants, for which success depends upon the self-sacrifice of sterile worker castes. Even though worker ants don’t directly reproduce, they still have reproductive fitness by both Wilson’s and Dawkins’s accounts. Since all the ants in a colony are genetically related, their labor does continue their genes’ existence, though it does so indirectly. But, for Wilson, biological life’s “conquest of Earth” is fundamentally a competition between groups of the same species. He thus considers the social group an additional unit on which natural selection acts, as if it were one organism. Dawkins criticizes Wilson’s theory:

Wilson now rejects “kin selection”… and replaces it with a revival of “group selection”—the poorly defined and incoherent view that evolution is driven by the differential survival of whole groups of organisms.

Nobody doubts that some groups survive better than others. What is controversial is the idea that differential group survival drives evolution, as differential individual survival does. The American grey squirrel is driving our native red squirrel to extinction, no doubt because it happens to have certain advantages. That’s differential group survival. But you’d never say of any part of a squirrel that it evolved to promote the welfare of the grey squirrel over the red. Wilson wouldn’t say anything so silly about squirrels. He doesn’t realise that what he does say, if you examine it carefully, is as implausible and as unsupported by evidence…

To borrow from Dorothy Parker, this is not a book to be tossed lightly aside. It should be thrown with great force. And sincere regret. [2]

However, Wilson has shown Hamilton’s (and Dawkins’s) deceptively simple forumla to be statistically less accurate than his own multilevel models, which explain individual actions in terms of both individual and group fitness. Wilson then responded, also in Prospect, with a brief word for Dawkins:

Richard Dawkins’s review of The Social Conquest of Earth makes little connection to the part he criticizes. The central issue in the book, which he urges others not to read, is the replacement of inclusive fitness theory (kin selection theory) by multilevel selection theory (ie, individual and group selection combined), with a new and major role assigned to group selection in the origin of advanced social behavior…. We demonstrated that while inclusive fitness theory sometimes works, its mathematical basis is unsound, and inclusive fitness itself is an unattainable phantom measure. Multilevel selection in contrast is mathematically sound, analytically clear, and works well for real cases—including human social behavior.

The science in our argument has, after 18 months, never been refuted or even seriously challenged—and certainly not by the archaic version of inclusive fitness from the 1970s recited in Prospect by Professor Dawkins.

Then comes Wilson’s real slam. Dawkins continually points out how lonely Wilson’s side of the evolutionary aisle looks, and Wilson responds with a lightly veiled charge that Dawkins has stopped analyzing scientific data and succumb to peer pressure by defending his mere personal beliefs––much the way of the religious bigots that Dawkins has dedicated his life to correcting:

While many have protested [multilevel selection]… many others of equal competence are in favor of the replacement proposed. In any case, making such lists is futile. It should be born in mind that if science depended on rhetoric and polls, we would still be burning objects with phlogiston and navigating with geocentric maps.

Wilson is surely one of the greatest biologists who ever lived, and he began his momentous career at the humble age of 9 by observing ants in his back yard in Alabama. By age 18, he undertook an independent project to map the location and behavior of populations of invasive and highly destructive fire-ants, which had just begun to colonize in the American South. His book The Ants chronicles his decades of fieldwork and laboratory research on these notoriously social insects and won the Pulitzer Prize for non-fiction in 1991. He followed this up with his semi-autobiographical novel, The Anthill in 2010. Ants are the creatures that led to Wilson’s founding of the modern field of sociobiology, which was the first systematic attempt to explain social behavior––from ants to human beings––in the empirical context of evolutionary biology. By manipulating social factors like genetic relatedness, environmental conditions, and group competition, Wilson tested ants’ behavior to determine the sociobiological forces that miraculously keep them loyal to their colonies. His work with ants can be seen in the 2008 NOVA documentary Lord of the Ants, which features impressive footage of the complex societies that ants form.

Rebounding from Dawkins’s blow, the 83-year-old Wilson showed that he’s still in the game with his latest article, “Evolution and Our Inner Conflict,” which recently appeared in The New York Times. Wilson is deeply invested in the implications of kin or group selection theory for the history and future of human nature. In his mind, this is far more than a scientific debate––it is a social debate, and a high-stakes one at that. He writes:

Are human beings intrinsically good but corruptible by the forces of evil, or the reverse, innately sinful yet redeemable by the forces of good? Are we built to pledge our lives to a group, even to the risk of death, or the opposite, built to place ourselves and our families above all else? Scientific evidence, a good part of it accumulated during the past 20 years, suggests that we are all of these things simultaneously. Each of us is inherently complicated. We are all genetic chimeras, at once saints and sinners — not because humanity has failed to reach some foreordained religious or ideological ideal — but because of the way our species originated across millions of years of biological evolution…

During the [period of Homo habilis], a conflict ensued between individual-level selection, with individuals competing with other individuals in the same group, versus group-level selection, with competition among groups. The latter force promoted altruism and cooperation among all the group members. It led to group-wide morality and a sense of conscience and honor. The competitor between the two forces can be succinctly expressed as follows: within groups selfish individuals beat altruistic individuals, but groups of altruists beat groups of selfish individuals. Or, risking oversimplification, individual selection promoted sin, while group selection promoted virtue.

So it appeared that humans are forever conflicted by their prehistory of multilevel selection. They are suspended in unstable and constantly changing locations between the two extreme forces that created us. We are unlikely to yield completely to either force as an ideal solution to our social and political turmoil. To yield completely to the instinctual urgings born from individual selection would dissolve society. To surrender to the urgings from group selection would turn us into angelic robots—students of insects call them ants.

The eternal conflict is not God’s test of humanity. It is not a machination of Satan. It is just the way things worked out. It might be the only way in the entire universe that human-level intelligence and social organization can evolve. We will find a way eventually to live with our inborn turmoil, and perhaps find pleasure in viewing it as a primary source of our creativity. [3]

Multilevel selection is the name Wilson places on his theory of combined individual and group selection, and it provides an explanation of human social tendencies that actually fits our highly conflicted understandings of ourselves. Though Dawkins’s neat, clean theory had its time in the limelight, there may be some wisdom in our intuitive aversion to reducing human beings to sacs of selfish genes. Dawkins’s selfish gene theory was attacked by religious leaders as immoral and evil, but Dawkins long defended it as a bitter but truthful medicine that humankind must swallow. However, now that Wilson’s data more accurately places man on a pendulum between group and individual interests, our selfish tendencies are only half the story. They are fundamentally challenged by our innate group morality, and both sides push and pull with considerable weight. These competing forces make us all simultaneously both saints and sinners; moral and immoral; selfish and selfless; and the list goes on.

Wilson’s multilevel selection distinguishes the tendencies of highly social creatures––of which human beings are the most outstanding example––from our less social animal cousins, which are more kin-selected. Though all biological life is subject to genes for reproduction, Wilson’s theory suggests that the genes of social beings (like humans and ants) flourish most when their selfishness is extended to be that of their group. Yet, in a way, this theory is just as disturbing as Dawkins’s: groups may be genetically programmed to exclude not just non-kin, but all individuals outside their social group. Just the way ants and bees form competing, hierarchical colonies, human beings may have in-group/out-group distinctions hard-wired and only act altruistically when they think it benefits their own group. Still, this theory at least allows for a genuine kind of altruism, whereas in selfish gene theory every action the individual performs is intended to directly enhance his or her own reproductive fitness.

“Human beings must have an epic, a sublime account of how the world was created and how humanity became part of it,” Wilson wrote in his landmark text of sociobiology On Human Nature. And he does not leave us hanging: “The evolutionary epic is probably the best myth we will ever have.” The Social Conquest of Earth is certainly a key chapter of that epic, alongside Dawkins’s masterful portrait of biodiversity in The Greatest Show on Earth (which is, of course, evolution). Both theories make for astonishing evolutionary epics––far more awe-inspiring than supernatural creation––but ultimately, Wilson’s is the only one that we can actually believe.


[1] Edward O. Wilson, The Social Conquest of Earth. New York: Liveright, 2012.
[2] Richard Dawkins, “The Descent of Edward Wilson”. Prospect.
[3] Edward O. Wilson, “Evolution and Our Inner Conflict”. The New York Times.

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