Ernest Rutherford is supposed to have said that ‘all science is either physics or stamp collecting’. This reflects an enduring perception that, with their test tubes, lab coats, and dense equations, the physical sciences are hard. Biologists, by contrast, have never quite been able to escape the image of themselves as chasing beetles and butterflies—gathering specimens which are arranged in expansive collections, neatly labelled and organised, with as much concern for aesthetics as for scientific advance. Biologists observe and document; chemists and physicists conduct rigorous experiments and take precise measurements.
Among the many great achievements of The Selfish Gene—now re-issued on the fiftieth anniversary of its publication—is that Richard Dawkins conclusively explodes the idea that biology seeks merely to record and classify the natural world.
Living things are the most complex entities in the universe. In The Blind Watchmaker (1986), Dawkins makes the case that physics and chemistry are concerned with fundamentally simple systems, because the mechanics of those systems are reducible to mathematical description. Living things are not like that. We cannot describe the workings of a single cell—let alone a whole complex, multi-cellular organism—in an equation; we are barely able to map out the intricacies of individual metabolic processes within a single cell.
Explaining the complexity of living things—biology’s central conundrum—is such a difficult problem that the passages of the planets and stars across the night sky were precisely described, computed, and predicted millennia before anyone came close to solving it. A bird’s wing, a frog’s leg, a hawk’s eye: these are highly complicated structures that fit an organism to its particular mode of existence, giving every appearance of deliberate design for functionality. Indeed, design was considered the only viable explanation until—nearly two centuries after Newton wrote the Principia and invented calculus—Charles Darwin developed a workable mechanism by which biological complexity could arise from simpler, pre-existing building-blocks.
Like all of biology, that mechanism is deceptively simple.
Offspring resemble their parents. But all individuals also vary. The resources needed for living things to sustain themselves (water, food, space) are finite, meaning that individuals must compete for access to those resources because many more are born than can survive. These three preconditions, identified by Darwin—variation plus competition plus inheritance—inevitably mean that individuals with traits that make them better fitted to their environment are the most likely to survive and reproduce, and therefore most likely to pass those winning traits on to their offspring. Darwin christened this natural selection, and explicitly compared it to artificial selection brought about by man through selective breeding, for example in dogs or crops or livestock. Where man selects based on his own desires and ultimate goals, nature selects blindly for fitted-ness to the environment.
Under natural selection, the most successful forms proliferate because they are the most successful at reproducing themselves. Telescoped over hundreds of millions of years, this slow, methodical, ruthless process gave rise to earthworms, whales, and pterodactyls—organisms that each seem perfectly designed for their respective ways of life.
The Selfish Gene,published in 1976, marks the end, or culmination, of a long scientific revolution that began with Darwin’s own book, On the Origin of Species, more than a century earlier in 1859.
But the idea of evolution didn’t begin with Darwin. Bubbling up throughout the eighteenth and early nineteenth centuries was a growing scepticism of the classical concept of species as fixed and unchanging kinds, and a shift toward a sense that living things might change over time: the ‘transmutation’ of species. Early modern biologists like the French naturalist Jean-Baptiste Lamarck envisioned a ladder of life that ascended, bit by bit, from simple organisms like plants and sponges, to mammals, primates, and man. For Lamarck, all life strove upwards toward perfection, with body forms that were fluid through use and disuse in life—the giraffe gets his long neck by continually stretching it up towards tree canopies, and those incremental changes to its body are inherited and accumulated over the generations.
As bizarre as it might seem from our vantage point, Lamarck’s idea of the ‘inheritance of acquired characteristics’ remained influential well into the nineteenth century, and offered one explanation for the fact that species seem undeniably designed. Biology at the time was in a state of flux, as scientists sought to harmonise new evidence with old ideas. The early evolutionists laid the scientific and cultural foundations for a theory that living things might change—a conceptual bedrock on which Darwin was able to establish natural selection as a far more convincing and powerful mechanism. Marshalling evidence from across the living world, the Origin acted as a lightning rod, breaking the deadlock and offering an effective evolutionary framework for future biological research.
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Darwin gave us the process, but the underlying mechanics remained opaque. DNA and even chromosomes—the cross-shaped packages into which DNA organises itself within cells—were yet to be discovered, and Darwin himself believed that heredity—the means by which offspring resemble their parents—was a process of parental blending. It was left to Gregor Mendel, an Austrian friar and Abbot of St. Thomas’s in Brno, working at the same time as Darwin but unknown to him, to deduce from botanical experiments that inheritance was particulate. Traits are passed on through discrete hereditary units, later termed genes, from one parent or the other, such that at the aggregate level offspring might appear superficially to be a blend of characteristics.
Mendel’s work initially fell into obscurity, with its significance only being fully appreciated when it was rediscovered in the early twentieth century. What followed were decades of trying to knit together Darwin’s ideas and the new science of genetics. Mutations at the gene level and the mixing up of genes through mating created variation within a population, and natural selection acted on that variation to promote the survival and reproduction of the fittest individuals. These then passed on their particular (and particulate) genes to the next generation. Over time, this process led to the evolution of complex biological structures from simpler, pre-existing ones.
Everything seemed to be coming together, but persistent paradoxes remained.
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Most conspicuously, if evolution selects the fittest individuals, how could we explain the occurrence of unselfish, altruistic behaviour in nature? Why do some animals share resources within a group—like wolves or wild dogs sharing meat with the rest of their pack—when it would make sense for an individual to monopolise a resource for their own benefit? Why should a bee sting a predator in defence of its hive, when that act means certain death?
If natural selection is all about survival, why sacrifice anything that could increase your chances in the high-stakes game of life?
Perhaps, through so-called ‘group selection’, bands of individuals within a species competed against one another, propagating the attributes of successful groups into the future. A co-operating, unselfish group might well defeat a group of treacherous, back-stabbing individuals. The question became one of what natural selection was actually selecting. The individual or the group? Or something else?
Group selectionists were in good company. Even Darwin himself occasionally drifted into group-based reasoning—the subtitle of the Origin talks not about the survival of the fittest individuals, but the ‘preservation of favoured races in the struggle for existence’. But even as a group-level framework seemed to solve some evolutionary problems, it raised others.
Consider a group of meerkats—small, burrowing African mammals. Individual meerkats take turns serving as a look-out for predators, while the rest of the group gets on with their daily tasks of foraging, feeding, and so on. Such a behaviour could feasibly evolve via group selection, because co-operative groups might out-compete non-cooperative groups. But suppose we introduce a genetic mutation—a cheating gene—which makes an individual actively avoid sentry duty. Our cheat relies on the alarm calls of other meerkats, but never takes the risk of serving as look-out himself. A gene that promoted such cheating would likely be very advantageous, and a cheat might survive longer and sire more offspring than an honest meerkat, passing that gene on. Before long, we would have more cheats than honest meerkats in our group and, with no reliable system of sentries and alarms, the social system of co-operation would break down: degenerate into every-meerkat-for-himself.
A similar problem arises in the sterile workers of ants and bees. On a straightforward group selection account, colonies with more co-operative workers should outcompete less co-operative colonies because helping behaviour enhances the efficiency and survival of the whole group. But such an explanation is vulnerable to the same sort of cheating we saw in our meerkat group: if a mutant worker can gain a direct reproductive advantage by withholding effort or by diverting resources toward its own reproduction, then that variant should spread within the colony. In that case, individual-based selection within groups would erode the co-operative system from the inside.
The vulnerability of co-operating groups to invasion by cheats posed a major challenge to early group selection explanations. It is not enough to say individuals work ‘for the good of the group’, let alone, as some of the more extreme group selectionists argued, that they regulate their own population size ‘for the good of the species’. There must be something more.
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The key to solving the problem of altruism came from looking within individuals, at the genetic information that acts as the recipe for building every organism.
A child shares half of all their genetic information with each parent. And with a grandparent, one quarter. By aiding a relative, therefore, an individual might help to ensure that some of that shared genetic information flows down the river of time and into the next generation, even if not through its own offspring. Although worker ants do not reproduce directly, they each share half of their genes with the queen, who does reproduce. A hypothetical cheat worker ant who could breed might be able to mate, and pass its genes on directly. However, if helping the queen produce large numbers of relatives carries more copies of a worker’s genes into future generations than direct reproduction, sterile ‘helping-only’ behaviour could evolve as a stable strategy. A co-operating worker, in other words, could end up with more (proportionately closer) relatives in the next generation than the cheat. This idea was pithily summarised by British biologist J. B. S. Haldane, who allegedly said he would ‘lay down [his] life for two brothers or eight cousins’.
By building individual bodies that co-operate, genes can increase their chances of surviving into the next generation—even if they do so by proxy, helping a relative that shares much of their genetic information to survive and reproduce in their stead.
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Darwin’s mechanism of natural selection, then, acts not at the level of species or groups, nor even, in truth, at the level of individual organisms, but on the invisible genetic units that are the building blocks of all life. Genes work together to build bodies: automata in which they ride, do battle, and engineer their own proliferation. Genes endure—the ‘immortal replicators’—passed down from individual to individual through the generations, and we are merely their ‘survival machines’.
In The Selfish Gene, that vision—the gene-centric view of evolution—is painted in vivid prose, and with such clarity of reasoning that it has become far more than a classic of popular science. Just as evolution did not begin with Darwin, the gene’s-eye view did not begin with Richard Dawkins. One finds it being painfully pieced together throughout the mid-twentieth century, by R. A. Fisher (The Genetical Theory of Natural Selection, 1930), W. D. Hamilton (‘The evolution of altruistic behavior’, 1963), G. C. Williams (Adaptation and Natural Selection, 1966), and many others. Indeed, these names are among those most frequently cited in Dawkins’s oeuvre, particularly in The Selfish Gene.
And yet, more than any other single work, The Selfish Gene crystallises the synthesis of natural selection and genetics, making the most coherent extended explanation of the fundamental, gene-based mechanics underlying evolution. With a gift for crafting a turn of phrase, Dawkins coined expressions and concepts in the book that have since proven highly successful replicators of their own, spreading vigorously in the public imagination. Through clarity of reasoning and metaphor, Dawkins not only popularised evolutionary theory but also solidified a genuine shift in the conceptual paradigm of the field.
Fifty years after its publication, The Selfish Gene remains a cornerstone of modern evolutionary biology and continues to be read by students around the world. In the epilogue to the new edition, Dawkins himself reflects on this startling fact: ‘In 1976, the very word ‘genomics’ had not yet been coined. The human genome’s presidentially celebrated completion was a quarter century in the future. The polymerase chain reaction (PCR) technique had not been invented, nor DNA fingerprinting […]’. While the world around has changed, the book’s central argument remains as relevant, and as invigorating, as when it was first written.
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I have tried to set The Selfish Gene within the context of the growth and development of evolutionary theory more broadly. Some might ask why tracing this path through biological history matters: science is about progress, they may say, not ruminating over the errors of long-dead predecessors. But to understand the importance of Dawkins’s contribution we must see how, together with On the Origin of Species, it book-ends one hundred years of striving towards a coherent understanding of evolution. Even today, the jigsaw is not complete; there are still holes scattered across the puzzle. But posterity will surely say that Richard Dawkins let us see, more clearly, the true nature of the picture we are working towards.
And that picture is one of breathtaking scope—the grand theoretical implications of the science of biology. The story of The Selfish Gene spans the history of life on Earth, from the simple self-replicating molecules that swarmed in prehistoric slime, to the evolving cultural replicators (‘memes’) of complex human societies. It is a tale of such power it must surely break free from the parochial confines of our own planet. ‘Not only could it have been published one hundred years ago: The Selfish Gene, or at least its message shorn of detail, could be published on any planet in the universe where life exists.’
Wherever there is life, there is Darwinism.
Related reading
‘An animal is a description of ancient worlds’: interview with Richard Dawkins, by Emma Park
‘The Genetic Book of the Dead’: A Dawkinsian Medley, by Daniel James Sharp
A farewell to Dawkins? by Samuel McKee
Why Design Arguments Necessarily Fail, by Samuel McKee
Linnaeus, Buffon, and the battle for biology, by Charles Foster
Consciousness, free will and meaning in a Darwinian universe: interview with Daniel C. Dennett, by Daniel James Sharp
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