COMMENTARY

It’s time to consign the “selfish gene” to the history books

The evolutionary idea that all organisms are singularly wired to ruthlessly pass on their genes is flawed

Published May 30, 2021 7:30PM (EDT)

Greedy (Getty Images)
Greedy (Getty Images)

There's an unforgettable moment in the movie "Wall Street" when financier Gordon Gekko tells the shareholders of Teldar Paper why his buyout proposal, incorporating massive layoffs, is not only profitable, but morally legitimate. With his slicked-back hair and custom-tailored suit, he struts to the front of the hall and proclaims that there is a "new law of evolution in corporate America." It's a simple law, he explains:

The point is, ladies and gentlemen, that greed—for lack of a better word—is good.

Greed is right.

Greed works.

Greed clarifies, cuts through, and captures the essence of the evolutionary spirit.

Greed, Gekko is declaring, is the basis of evolution and all that's arisen from it — including human supremacy. Gekko's speech was unleashed on moviegoers in 1987 as the world was reeling from an early encounter with the excesses arising from global financial deregulation. His signature claim — "Greed is good!" — has since become the stuff of legend, strikingly capturing the ethos of unrestrained, free market capitalism that has come to dominate mainstream thinking.

The idea that selfishness and greed are drivers of evolution, and therefore possess underlying virtue, has been around for over a century, ever since Charles Darwin's theory of evolution became widely accepted. The archetypal robber barons, Andrew Carnegie and John D. Rockefeller, both argued that the "survival of the fittest" principle justified their cutthroat tactics. But the publication in 1976 of Richard Dawkins's bestseller, "The Selfish Gene," adroitly repackaged the notion for modern times, reducing the complexities of evolution to a brutally elemental simplicity. As Dawkins summarized it:

The argument of this book is that we, and all other animals, are machines created by our genes. Like successful Chicago gangsters, our genes have survived, in some cases for millions of years, in a highly competitive world. This entitles us to expect certain qualities in our genes. 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. . . . 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.

With the notion of the "selfish gene" as the ultimate driver of evolution, Dawkins helped forge the moral framework of his age. Influential thought leaders have since infused this supposed biological truth into economics, politics, and business. "The economy of nature is competitive from beginning to end," writes sociobiologist M. T. Ghiselin, coeditor of the Journal of Bioeconomics.

It's difficult to overstate the pervasiveness of Dawkins's selfish gene theory in popular culture. In a nutshell, the underlying story goes something like this: All organisms in nature are simply vessels for the replication of the selfish genes that control us. As such, all living entities — including humans — are driven to compete ruthlessly to pass on their genes. This struggle for reproduction is the underlying engine of evolution, as occasional positive random mutations in genes give an entity a competitive edge to beat out weaker rivals. Any apparently altruistic behavior is merely a convenient tactic for a concealed selfish goal. Since nature works most effectively based on selfishness, human society should be similarly organized, which is why free market capitalism has been so successful in dominating all other socioeconomic models.

However, pervasive as it has become throughout our culture, the story of the selfish gene is based on fundamental misconceptions. In recent decades, researchers in evolutionary biology have overturned virtually every significant assumption in the selfish gene account. In its place, they have developed a far more sophisticated conception of how evolution works, revealing the rich tapestry of nature's dynamic interconnectedness. Rather than evolution being driven by competition, it turns out that cooperation has played a far more important role in producing the great transitions that led to Earth's current breathtaking state of diversity and beauty.

The trouble with the selfish gene story is not just that it is scientifically flawed; it's also that it presents such an impoverished view of life's dazzling magnificence. The discoveries of modern researchers showing how life evolved to its current state of lavish abundance reveal a spectacle of awe-inspiring complexity, mind-boggling dynamic feedback loops, and infinitely subtle interconnections.

Decoding the "book of life"

For nearly a century after Charles Darwin published his theory of evolution by natural selection in 1859, biologists struggled to identify precisely what caused adaptive traits to be inherited by future generations. Early in the twentieth century, they combined the ideas of Darwin with other groundbreaking researchers such August Weismann and Gregor Mendel to construct what became known as the Modern Synthesis, which has been the dominant interpretation of evolution ever since. The central concept that held it together was the gene: a hypothetical unit of natural selection that was passed on through inheritance. It was the gene that somehow specified the form an organism would take. Random mutations in an individual's genes occasionally gave it unique traits that were different from the rest of its species, and those with the best adapted traits passed these genes onto the next generation.

What exactly were these genes, and how did they pass on their specifications? This burning question was finally answered in 1953 when James Watson and Francis Crick, along with Rosalind Franklin, discovered the double-helix shape of the DNA molecule containing an organism's genes, and described how tiny molecular subunits, named bases, paired with each other to specify proteins that would then be constructed within a cell. Now things were becoming clear! It was as though the secret of life itself had been laid bare to scientific understanding. With headlines blaring around the world, a new story of life entered the public consciousness.

Using metaphors from the newly burgeoning field of information theory, Crick and Watson published their findings in a legendary paper in Nature, writing that "It therefore seems likely that the precise sequence of the bases is the code which carries the genetical information." Now the race was on to "decode" what was quickly becoming known as the "book of life."


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As computing sophistication exponentially increased, the possibility arose that combined advances in molecular biology and information processing might allow a complete mapping of the human genome. Planning began in 1984 and the Human Genome Project formally launched in 1990. During this time, information technology and genetics became tightly linked, both technically and conceptually. The genotype was seen as a "program" that determined the exact specifications of an organism, just like a computer program. DNA sequences formed the "master code" of a "blueprint" that contained a detailed set of "instructions" for building an individual. Prominent geneticist Walter Gilbert would begin his public lectures by pulling out a compact disk and proclaiming "This is you!"

Public excitement around the Human Genome Project was effervescent. The promise was intoxicating: If every gene specified a particular protein in a cell, then once we'd mapped them all, we'd eventually be able to identify the genetic cause for every attribute of a person. We'd know the genes for intelligence, for athletic prowess, for longevity—and of course find cures for a wide array of diseases. "Our fate," Watson declared, "is in our genes."

Thanks to dramatic advances in computing power, the human genome was triumphantly mapped in 2003, years ahead of schedule. However, some rather inconvenient facts quickly began to rain on the parade. It turned out that the entire human genome contained about twenty-one thousand genes that coded for proteins. The tiny roundworm C. elegans, rather embarrassingly, had a similar number, while wheat had more than four times as many. These humbling statistics clearly showed there was something more going on in the cell than simple one-to-one coding between genes and proteins. And even without these awkward comparisons, it was abundantly clear that there were more than twenty-one thousand attributes that determined every aspect of a human being. What was wrong with the model?

The language of the gene

At its heart, the model's fundamental flaw was the machine metaphor it was built on. Genetic determinism, as it is sometimes called, was based on the underlying idea that organisms, like machines, are comprised of components with linear relationships that can be precisely determined.

When Crick and Watson laid the foundation for what they called the "central dogma" of molecular biology, they specified that information could only flow one way. It began with the structure of the gene, which was defined as a particular sequence of base pairs that coded for a protein. In everyday life, we think of protein as an undifferentiated substance we need for good health (as in "Are you getting enough protein?"). Within our bodies, however, proteins are a staggeringly diverse group of molecular structures that carry out most of the business of a cell. By some estimates, there are millions of different proteins within a single cell, each one of which is uniquely shaped in wildly complex configurations, a bit like a microscopic self-organized clump of squiggling steel wool. Something Crick and Watson didn't know when they set down their central dogma was that, in addition to all their other tasks, proteins act directly on the DNA of the cell, specifying which genes in the DNA should be activated.

This is a crucial discovery, which has become a mainstay of modern molecular biology, but has not yet made it into broad public consciousness. It means that the relationship between genes and the organism is not one-way but circular. DNA can't do anything by itself — it only functions when certain parts of it get switched on or off by the activities of different combinations of proteins, which were themselves formed by the instructions of DNA. This process is a vibrant, dynamic circular flow of interactivity. Living organisms, it turns out, are complex systems with multiple feedback loops creating nonlinear relationships. As such, they demonstrate far more complexity than even the most complicated computer.

What this means is that there is no such thing as a simple "gene for something." Genes are expressed within the cell as a result of what is going on around them. Rather than "coding" for something, the way a programmer writes code for a computer, a more useful metaphor for what happens might be "language." Think about how language works. Sometimes it can be a simple instruction: I might tell you to "turn right at the stop sign, then go half a mile until you see the store on the left." But it is frequently context dependent. If I call out "Help me!" you might turn to look at what I'm doing. If I'm carrying a tray piled high with glasses, perhaps you'll remove some to reduce the risk of them toppling. If I'm a teacher, and I've written some arithmetic on the board, you might call out the answer. Depending on the context, you'll respond very differently to the same words. In addition, you might engage in conversation with me, expecting me to respond, and together we might come up with a creative approach to a situation that I wouldn't have arrived at alone. Similarly, the gene sometimes gives clear instructions, and at other times engages in an interactive conversation with the rest of the cell. Just as words have an array of different meanings based on their context, syntax, and grammar, so DNA and proteins use their own language, with its own syntax and grammar, to determine what's best at that moment for the cell.

Take, for example, a cute little grasshopper. It walks slowly on long, spindly legs, eating alone and minding its own business. Obviously, a different species than a locust, which has short, crooked legs, and forms terrifying swarms that can darken the sky and aggressively devour an entire region's crops. Right? In fact, it turns out that grasshoppers and locusts have exactly the same DNA. Just like Dr. Jekyll and Mr. Hyde, when a certain kind of grasshopper senses its environment changing, either from food scarcity or overcrowding, it can transform itself within hours to a manically aggressive locust. Its cells switch on different genes within its DNA, it begins shrinking its legs and wings, changing its coloring, and even growing its brain to deal with the social complexities of the swarm. Later on, when the environment improves, its cells will switch their DNA settings and the locust will magically transform back into a quiet grasshopper.

Going beyond the gene

As biologists gain a deeper understanding of the cell's participation in genetic expression, they have also begun to re-examine the cell's role in evolution. An increasing number of prominent evolutionary biologists have become convinced that the new understanding of evolution requires expanding the theoretical framework beyond the Modern Synthesis that has ruled for nearly a century. They're not agitating for a Copernican-style revolution that rejects the entire conventional model; rather, they're calling for an "Extended Evolutionary Synthesis," arguing that the new findings require a broader conceptual framework. They are holding international conferences on the topic and their papers are flooding the most prestigious academic journals. "A profound, radical, and fascinating transformation of evolutionary theory is taking place," writes one of their leaders, Eva Jablonka.

The foundational idea of the new thinking is that evolution is driven, not by genes alone, but by organisms which, in the words of leading proponent Kevin Laland, "play active and constructive roles in their own development and that of their descendants." Animals, they declare, are masters at directing their own evolution by modifying their environment in ways that eventually become an integral part of their species' repertoire. We see examples of this everywhere in nature. Think of a bird making a nest. It's constructing a little niche in the environment that keeps its eggs safe, protecting them both from predators and temperature fluctuations. Those eggs don't need to be as resilient against cold temperatures as they otherwise would be. And when the chicks grow up, the ones that build better nests will be more successful at rearing their own offspring. Through their niche construction (as this process is called), birds direct their own evolution as a species. The same is true for spiders, whose reliance on web construction favors offspring who can produce sticky threads and react to vibrations on the web. Plants are equally effective in producing their own niches: they have learned to change the acidity, salinity, and other characteristics of the soil to make it more nutritious for their roots and those of their neighbors.

Organisms, then, play a crucial role in looking after themselves and their offspring in sophisticated ways. But how about the "selfish" question? Have we just relocated the "selfishness" of the gene to that of an individual organism? In fact, one of the most important findings in modern biology has been that cooperation, not selfish competition, has been the foremost driving force in each of life's major evolutionary transitions since it began on Earth billions of years ago.

The network of life

Earliest life consisted of single cells called prokaryotes, very similar to the bacteria that have thrived on Earth ever since. Prokaryotes contain relatively simple genomes, which they pass on to the next generation by dividing themselves into two, each daughter cell containing the identical DNA as the parent.

It was a billion years or so before a new type of cell arrived on the scene. Called a eukaryote (Greek for "true kernel"), this cell contained a nucleus that housed all its DNA material. Eukaryotes found a novel way to get their nutrition: they took advantage of their more flexible cell walls to engulf other bacteria and ingest them, breaking their parts down to use as food.

Except that something rather strange happened—and probably more than once. A eukaryote engulfed a prokaryote, and instead of digesting it, they started working together. This particular prokaryote was a tiny powerhouse, specialized in taking oxygen and turning it into energy. Called a mitochondrion, it formed a relationship with eukaryotes that could lay claim to be the most successful partnership on Earth. Every organism that you see around you—every plant, every insect, every animal—is comprised of eukaryotic cells containing hundreds, and sometimes thousands, of mitochondria within them (in plants they're called plastids), producing the energy that allows the cell to go about its business. To this day, mitochondria still carry their own DNA with them, which they use to replicate separately from the rest of the cell.

This startling hypothesis was proposed by biologist Lynn Margulis in 1967, and she underwent years of ridicule from the mainstream scientific community until it was finally accepted. It is now recognized as an indisputable part of Earth's evolutionary history — and not just any part. The cooperation initiated between eukaryotes and mitochondria is viewed by many biologists as one of the most important events ever to have occurred in the history of life on Earth.

Prokaryotes, meanwhile, may have remained relatively simple in their design, but they developed their own striking form of cooperation which has enabled them to flourish even in a world of energy-guzzling eukaryotes. They learned to share their genes with each other, in much the same way that neighbors in a tight-knit community might share their tools, favorite books, or useful gardening tips. They've been doing this now for billions of years, and it may be the single most important skill that has enabled them to thrive over the eons, allowing them to survive even in the most inhospitable places, such as hydrothermal vents, oil slicks, or radioactive dumps. Most of us know about this powerful networking trick of bacteria through their antibiotic resistance, which they transfer to each other through gene sharing.

In the early years of life on Earth, it's likely that gene sharing (known officially as horizontal gene transfer) was the predominant way evolution worked. In fact, researchers now believe that the eukaryote genome was itself the result of a fusion of two prokaryotic genomes. Instead of a Darwinian "tree of life," biologists are offering alternative metaphors such as a "bush" or "net" of life to better describe how we are all intricately connected. In the memorable words of Lynn Margulis: "Life did not take over the world by combat but by networking."

As eukaryotes, boosted by their mitochondrial energy packs, developed larger and more complex cellular structures, horizontal gene transfer became less important for them. While they still continue to engage in it, the size and complexity of their genomes make it more difficult. Eukaryotes, however, gradually devised another form of cooperation that led to the full unfolding of the prodigious grandeur of life on Earth as we know it today—multicellularity.

Better together

Almost every manifestation of nature that we can see with the naked eye is multicellular: a daisy, a rhinoceros, and a tiny mite, are all composed of a multitude of cells doing different things, yet working together for the greater interest of the organism. The evolutionary step from a single-celled eukaryote to the emergence of multicellular life took a long, long time—about a billion or so years, during which so little changed on Earth that it's been called the "boring billion."

Why did it take so long? No-one knows for sure, but a clue may reside in the staggering intricacy of gene expression that multicellular life had to master. It's one thing for a cell to organize itself and then divide into two. It's quite another to figure out how a newly born cell can split into two new cells different from each other but working together toward a shared goal, and then continue this differentiating process time and time again.

A fundamental distinction had to be made between cells that specialized in passing on genes to the next generation (germline cells) and the somatic cells that took care of everything else. Here again, we see a major evolutionary breakthrough in the scale of nature's cooperation. Somatic cells had to give up their own ability to reproduce in order to become part of something bigger than themselves. Without this momentous accord, there would be no complex life on Earth.

Even as they followed their unique evolutionary pathways to become redwood trees, whales, or worms, virtually all creatures on Earth have continued to share about a third of their genes from the collective ancestral pool. Because of our deep common ancestry, even though animals and bacteria have very different lifestyles, our cells are able to communicate using the same genomic language. Even as species differentiated, they developed ways to trade their own specialized skills for the unique skills of other species that could help them thrive. This process, known as mutually beneficial symbiosis (or mutualism) is so widespread throughout nature that it forms a bedrock of every ecology on Earth. The prevalence of mutualism means that life is rarely a zero-sum game, where a species can only gain at the expense of another. On the contrary, by working together, species have co-created ecosystems everywhere in which the whole is far greater than the sum of the parts.

These deeply intimate symbioses are everywhere in nature, forming the foundation of the living world. It's impossible take a walk in the woods, eat a meal, or dip in the ocean, without participating in the deep symbioses that have nourished life's plenitude. On the most fundamental level, plants have specialized in transforming sunlight into chemical energy that provides food for other creatures, whose waste then fertilizes the soil that the plants rely on. If you hike in the woods, you may notice how the trees provide shade that maintains moisture for creatures on the ground. Below you, mycorrhizal fungi maintain underground networks allowing "guilds" of trees to exchange carbon and nutrients among each other in a sophisticated interplay of resources that's been dubbed the "wood-wide web."

These symbiotic relationships are frequently so intimate that we rely on them without even knowing about it. We share our bodies with a vast multitude of bacteria—more than the number of cells we call our own. We need them to help us perform biochemical tricks that we can't do ourselves, such as producing enzymes to digest food that our own enzymes can't manage. These symbionts are so important to us that, after birth, a mother's milk contains special sugars that the baby can't digest but provide nutrition for the newborn's symbiotic bacteria.

In countless instances, over hundreds of millions of years, life has decided time and again, that things work better together.

Cooperation, competition, and harmony

Where, then, does competition fit into the picture? It seems clear that the gene's supposed "ruthlessly selfish" drive to replicate is not the sole explanatory factor of evolution. But surely competition has nevertheless had a significant part to play? What about all those spectacular nature documentaries showing cheetahs sprinting to catch gazelles? Male chimpanzees fighting rivals for sexual dominance? Bacteria that make us sick by overpowering our immune systems? There is no question that ruthless competition also has a central role to play in the drama of life. How can we reconcile pervasive competition with the forces of cooperation?

Let's imagine a spectrum with extreme competition at one end and extreme cooperation at the other. We can think of an organism as an ecosystem where the different parts have agreed to coexist at the cooperative end of the spectrum. Outside the organism, however, relationships exist all along the spectrum. An ecosystem can be understood as the emergent creation of organisms acting together in different degrees of competition and cooperation. In fact, the creative tension that arises from the confluence of both competition and cooperation is itself a driving force of evolution.

A pair of prominent evolutionary biologists, David Sloan Wilson and E. O. Wilson, have developed a sophisticated theory they call multilevel selection, tracing the dynamics between cooperative and competitive behavior at different scales of life. E. O. Wilson, a world leader in the study of social insects, has shown how colonies of ants that cooperated closely were more evolutionarily successful than those that experienced internal competition. The same is thought to be true of human evolution, when early hominids developed deeply felt values such as compassion, altruism, and fairness, which enabled them to live complex lives together in community. The groups in which these attributes predominated were more successful at hunting, foraging, and defending themselves from attack.

We can conceive of evolution, now, as a multidimensional force acting through both competition and cooperation at multiple levels—within the organism, in symbiotic relationships, within a species, between species, and within an ecosystem. At each level, competitive and cooperative forces create their own dynamic tensions, while simultaneously impacting other levels.

With this in mind, we can move beyond a sterile debate of whether evolution is a result of competition or cooperation. After all, these are concepts created by humans to establish neat categories. Living systems, whether they're genomes, cells, organisms, or ecosystems, have no interest in sticking to a category. We know that trees rely symbiotically on animals to spread their seeds. However, nut trees would have a problem if the squirrels ate all their nuts before they could germinate. To overcome this, in a phenomenon known as mast fruiting, they cooperate as a species, refrain from producing nuts for several years, and then collectively decide one year to produce an overwhelming number of nuts, so the squirrels will be unable to devour them all. Who's competing? Who's cooperating?

Maybe there's another way to describe the elegantly complex interweaving of natural processes that comprise an ecosystem: harmony. In music, harmony arises when different notes sound at the same time in such a way that an emergent, more complex and pleasing sound is produced. The notes aren't competing or cooperating with each other, but the way in which their differences act upon each other creates a blended experience that is richer, and more beautiful, than any of them alone. Could it be that the best description of how nature works is, in fact, a harmonic meshwork of life?

Mind the metaphor

The mainstream metaphors used to describe the evolution of life do a great disservice to its awesome majesty, while influencing us to think about our own lives, our planet, and our society in harmful and destructive ways. Metaphors are more than just techniques to communicate ideas—they form foundational structures of thought in the human brain that we unconsciously use to construct our worldview and shape our value system. Metaphors matter.

The most pervasive mistaken metaphor of life in common currency is that it's merely a very complicated machine. This goes back to the seventeenth century philosophical musings of Descartes and Hobbes, but it has fused with the bedrock of modern thought ever since Crick and Watson defined the gene in terms of coded information. At this stage, it's difficult to read any popular discourse about life without being bombarded by this misconception.

This metaphor becomes even more dangerously pervasive when it's used to describe the human mind which, according to a pair of evolutionary psychologists, is "a set of information-processing machines that were designed by natural selection to solve adaptive problems faced by our hunter-gatherer ancestors." The brain, we are told, is "a computer that is made of organic (carbon-based) compounds rather than silicon chips."

And then, of course, there's the Gordon Gekko crowd, justifying exploitative free-market global capitalism on the basis that it's what evolution intended. The metaphor of "life as a market" pervades public discourse so extensively that it seems like it must be nature's own way.

How would it change our conception of nature, and of our own social norms, if we instead framed these symbiotic relationships in terms of mutual consideration?

Frequently, when cell biologists describe the mind-boggling complexity of their subject, they turn to music as a core metaphor, making statements like "the music of life is a symphony." However, a symphony is a piece of music written by a composer, with a conductor directing how each note should be played. The awesome quality of nature's music arises from the fact that it is self-organized. There is no outside agent telling each cell what to do.

Perhaps a more illustrative metaphor would be an improvisational jazz ensemble, where a self-organized group of musicians spontaneously creates fresh melodies from a core harmonic theme, riffing off each other's creativity in a similar way to how we've seen evolution work. A related metaphor—and perhaps even more compelling—is a dance. Cell biologists increasingly refer to their findings in terms of "choreography," and philosopher of biology Evan Thompson writes vividly how an organism and its environment relate to each other "like two partners in a dance who bring forth each other's movements."

If our mainstream media and commentators began using these metaphors in place of the selfish gene, before long we might begin to perceive our world in a fundamentally different way. What might happen if we applied this new understanding of nature's harmonic dance to establish different norms for our own society? Imagine if, instead of our socioeconomic system constructed on the presumption that "the economy of nature is competitive from beginning to end," it was structured instead on the basis of symbiosis—an ecological civilization.

Ecosystems have developed tremendous resilience from these internal dynamics, sometimes existing for millions of years, continually adapting yet remaining stable and robust. Widespread symbiosis means there are no waste products—what one species expels is nutrition for another. Healthy ecosystems embrace both competition and cooperation at multiple levels, but always within a context of harmony for the entire system. The possibility of applying these ecological principles to our own society, and using them as an alternative way for humanity to organize itself, could potentially offer a powerful new navigating principle to steer our civilization on a course to survive the existential challenges of this century.


By Jeremy Lent

Jeremy Lent is an author and speaker whose work investigates the underlying causes of our civilization’s existential crisis, and explores pathways toward a life-affirming future. His recently published book is The Web of Meaning: Integrating Science and Traditional Wisdom to Find Our Place in the Universe. Find his website at jeremylent.com.

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Biology Commentary Evolution Richard Dawkins Science Selfish Gene