IN 1973, strike action by the National Union of Mineworkers led to a crisis in which the Conservative government of Edward Heath imposed a so-called ‘three-day week’ in Britain. In order to conserve fuel stocks, electricity for non-essential purposes was rationed. We were limited to three days per week, and there were frequent power cuts. My cricket research depended on electricity but writing didn’t – in those days I wrote with a portable typewriter on, of all outlandish surfaces, sheets of flat white stuff called paper. So I decided to call a temporary halt to my cricket research and begin work on my first book. This was the genesis of The Selfish Gene.
Selfishness and altruism and the whole idea of a ‘social contract’ were much in the air at the time. Those of us on the political left tried to balance our sympathy for the miners on the one hand with hostility to what some saw as their strong-arm tactics, holding society at large to ransom, on the other. Did evolutionary theory have anything to contribute to this important dilemma? The previous decade had seen a string of popular science books and television documentaries, gallantly attempting to apply Darwinian theory to questions of altruism and selfishness, collective versus individual welfare, but actually getting the theory flat-out wrong. The error was always a version of what has been called ‘evolutionary panglossism’.
As reported by my friend and mentor the late John Maynard Smith, his own mentor the formidable J. B. S. Haldane satirically coined three erroneous, or at best unreliable, ‘theorems’. Aunt Jobiska’s Theorem (from Edward Lear) was ‘It’s a fact the whole world knows . . .’ The Bellman’s Theorem (from Lewis Carroll) was ‘What I tell you three times is true.’ And Pangloss’s Theorem (from Voltaire) was ‘All is for the best in this best of all possible worlds.’
Evolutionary panglossians are vaguely aware that natural selection does a pretty effective job of making living things good at their business of living. Albatrosses seem beautifully designed for flying above the waves, penguins for flying beneath them (I happen to be writing this on a ship in Antarctic waters, marvelling through my binoculars at these prodigies of avian virtuosity). But panglossians forget, as it is so easy to forget, that this ‘good at’ applies to individuals, not to species. Good at flying, good at swimming, surviving, reproducing, yes, natural selection will tend to make individual animals good at those things. But there is no reason to expect natural selection to make species good at avoiding extinction, good at balancing their sex ratio, good at limiting their population in the interests of the common weal, good at husbanding their food supply and conserving their environment for the benefit of future generations. That would be panglossism. Group survival may emerge as a consequence of improved individual survival, but that is a fortunate by-product. Group survival is not what natural selection is about.
The panglossian error is tempting because we humans are blessed with foresight and can judge which actions are likely to benefit our species, or our town, or our nation, or the whole world, or any specified entity or interest group, in the future. We can foresee that to overfish the seas would be in the long run counterproductive for all fishermen. We can foresee a happier future if we limit our birth rate so that fewer individuals are born, to enjoy richer lives. We can decide that self-restraint now will pay dividends in the future. But natural selection has no foresight.
To be sure, a panglossian version of natural selection theory had been proposed, which, if only it worked, might achieve something like the ‘all is for the best’ utopia. But unfortunately it doesn’t work. At any rate, it was one of my aims in The Selfish Gene to persuade my readers that it doesn’t work. This was the theory called ‘group selection’. This maddeningly seductive error – the Great Group Selection Fallacy or GGSF – ran all through Konrad Lorenz’s popular 1964 book On Aggression. It also pervaded Robert Ardrey’s best-sellers, The Territorial Imperative and The Social Contract – where I was especially affronted by the mismatch between Ardrey’s erroneous message and the high quality of the English in which he expressed it.57 I aspired to publish a book on the same theme as Ardrey’s Social Contract (itself a sort of biological rewrite of Rousseau’s famous treatise); but mine would be based on rigorous natural selection theory, not the GGSF. My ambition was to undo the damage done by Ardrey and Lorenz – and by many television documentaries of the time, whose promulgation of the error was so ubiquitous that in The Selfish Gene I even dubbed it ‘the BBC Theorem’.
I was all too familiar with panglossism and the GGSF because I met them weekly in undergraduate essays. And indeed I, when an undergraduate myself, had laced many of my essays with the fallacious view that what really matters in natural selection is the survival of the species (my tutors never noticed). When I eventually came to write The Selfish Gene, my dream was that I would change all that. I was daunted by the knowledge that, in order to succeed, my book would need to be as well-written as Ardrey’s and sell as prolifically as Lorenz’s. I jokingly spoke of it as ‘my best-seller’, never really believing that it would become one but giving self-consciously ironic voice to my wilder ambition.
Natural selection is a purely mechanical, automatic process. The world is constantly tending to become full of entities that are good at surviving, denuded of things that are not. Natural selection has no foresight, but brains have, and that is why panglossism is so appealing to us. Brains may agonize about the long-term future and project forward this century’s self-indulgence into next century’s catastrophe. Natural selection cannot do that. Natural selection can’t agonize about anything. Natural selection can only blindly favour short-term gain, because every generation is automatically filled with the offspring of those individuals who did whatever it took, in the short term, to manufacture offspring more effectively than other individuals of their own generation.
And when you look carefully and hard at exactly what is going on as the generations flash by, your gaze is drawn irresistibly to the gene as the level at which natural selection really works. Natural selection automatically favours self-interest among entities that potentially can pass through the generational filter and survive into the distant future. As far as life on this planet is concerned, that means genes. Here’s how I put it in The Selfish Gene, where I introduced the phrase ‘survival machine’ to describe the role of (mortal) individual organisms vis-à-vis their (potentially immortal) genes:
The genes are the immortals . . . [they] have an expectation of life that must be measured not in decades but in thousands and millions of years.
In sexually reproducing species, the individual is too large and too temporary a genetic unit to qualify as a significant unit of natural selection. The group of individuals is an even larger unit. Genetically speaking, individuals and groups are like clouds in the sky or dust-storms in the desert. They are temporary aggregations or federations. They are not stable through evolutionary time. Populations may last a long while, but they are constantly blending with other populations and so losing their identity. They are also subject to evolutionary change from within. A population is not a discrete enough entity to be a unit of natural selection, not stable and unitary enough to be ‘selected’ in preference to another population.
An individual body seems discrete enough while it lasts, but alas, how long is that? Each individual is unique. You cannot get evolution by selecting between entities when there is only one copy of each entity! Sexual reproduction is not replication. Just as a population is contaminated by other populations, so an individual’s posterity is contaminated by that of his sexual partner. Your children are only half you, your grandchildren only a quarter you. In a few generations the most you can hope for is a large number of descendants, each of whom bears only a tiny portion of you – a few genes – even if a few do bear your surname as well.
Individuals are not stable things, they are fleeting. Chromosomes too are shuffled into oblivion, like hands of cards soon after they are dealt. But the cards themselves survive the shuffling. The cards are the genes. The genes are not destroyed by crossing-over, they merely change partners and march on. Of course they march on. That is their business. They are the replicators and we are their survival machines. When we have served our purpose we are cast aside. But genes are denizens of geological time: genes are forever.
I had already convinced myself of this truth a decade earlier, in almost exactly the same words, when I gave the 1966 undergraduate lectures in Oxford that I have already described. I recalled on page 199 the rhetorical flourishes with which I tried to persuade the undergraduates of the centrality of the immortal gene in the logic of natural selection. Here are my 1966 words, and you can see how similar they are to the equivalent, more rhetorical paragraphs of The Selfish Gene.
Genes are in a sense immortal. They pass through the generations, reshuffling themselves each time they pass from parent to offspring. The body of an animal is but a temporary resting place for the genes; the further survival of the genes depends on the survival of that body at least until it reproduces, and the genes pass into another body . . . the genes build themselves a temporary house, mortal, but efficient for as long as it needs to be . . . To use the terms ‘selfish’ and ‘altruistic’ then, our basic expectation on the basis of the orthodox neo-Darwinian theory of evolution is that Genes will be ‘selfish’.
When I recently found the text of that 1966 lecture (with its encouraging marginal note by Mike Cullen), I was surprised to realize that I had not then read George C. Williams’s book, Adaptation and Natural Selection, published in the same year:58
With Socrates’ death, not only did his phenotype disappear, but also his genotype . . . The loss of Socrates’ genotype is not assuaged by any consideration of how prolifically he may have reproduced. Socrates’ genes may be with us yet, but not his genotype, because meiosis and recombination destroy genotypes as surely as death.
It is only the meiotically dissociated fragments of the genotype that are transmitted in sexual reproduction, and these fragments are further fragmented by meoisis in the next generation. If there is an ultimately indivisible fragment it is, by definition, ‘the gene’ that is treated in the abstract discussions of population genetics.
When I eventually read Williams’s great book (some years later, I regret to say), his Socrates passage immediately resonated with me and I prominently acknowledged Williams’s importance, as well as Hamilton’s, for the theme of The Selfish Gene when I came to write it.
Williams and Hamilton were somewhat similar characters: quiet, withdrawn, self-effacing, deep-thinking. Williams had a dignity and mien which – perhaps enhanced by his high forehead and the cut of his beard – reminded many of Abraham Lincoln. Hamilton had more an air of A. A. Milne’s Eeyore. But when I wrote The Selfish Gene I didn’t know either man, just their published work and how central it was for our understanding of evolution.
Because genes are potentially immortal in the form of accurate copies, the difference between successful genes and unsuccessful genes really matters: it has long-term significance. The world becomes filled with genes that are good at being there, good at surviving through many generations. In practice that means good at cooperating with other genes in the business of building bodies that have what it takes to survive long enough to reproduce – for bodies are the temporary vehicles in which genes reside and which pass them on. Throughout The Selfish Gene I used that phrase ‘survival machine’ as my name for an organism. Organisms are the entities in life that actually do things – move, behave, search, hunt, swim, run, fly, feed their young. And the best way to explain everything that an organism does is to assume that it has been programmed, by the genes that ride inside it, to preserve those genes and pass them on before the organism itself dies.
I also used the word ‘vehicle’ as equivalent to ‘survival machine’. It reminds me of an amusing occasion when a Japanese television crew came to interview me about The Selfish Gene. They all travelled to Oxford from London packed into a black cab, with tripods and floodlights and, as it seemed, arms and legs sticking out of every window. The director informed me, in halting English (the official interpreter couldn’t make me understand him at all and had been sent off in disgrace), that he wanted to film me in the taxi as it drove around Oxford. This puzzled me, and I asked why. ‘Hoh!’ came the puzzled reply: ‘Are you not author of Taxicab Theory of Evolution?’ I afterwards guessed that the Japanese translators of my writings must have rendered ‘vehicle’ as ‘taxicab’.
The interview itself was quite amusing. I rode in the taxi alone except for the cameraman and sound man. In the absence of the official interpreter, there was no interviewer, and I was bidden simply to talk ad lib about The Selfish Gene while we did a scenic tour of Oxford. The taxi driver doubtless had the streets of London intricately mapped in his enlarged hippocampus, but he didn’t know Oxford. It therefore fell to me to guide him, and my otherwise measured discourse on selfish genes was punctuated by frantic shouts of ‘Turn left here!’ or ‘Turn right at the traffic lights and then get in the right-hand lane!’ I hope they managed to find the unfortunate interpreter before returning to London.
In The Selfish Gene, I criticized the panglossian idea that animals have foresight and work out what would be good for the long-term future of their species or group. What is wrong with this is not the idea that animals ‘work out what would be good’. There is in any case no suggestion that the ‘working out’ is conscious. No: what is wrong is the idea of the species or group as the entity whose benefit is maximized. Biologists often legitimately use the language of ‘working out what would be good’ as a shorthand route to sound Darwinian reasoning. The trick is to identify the correct level in the hierarchy of life at which the shorthand metaphor of conscious reasoning is applied. It is quite OK to put yourself in the position of an individual animal and ask: ‘What would I do if I were trying to achieve the goal of propagating my genes?’
The Selfish Gene is filled with imagined soliloquies in which a hypothetical animal ‘reasons’ to itself: ‘Should I do X or Y?’ The meaning of ‘should’ is: ‘Would X or Y be better for my genes?’ This is legitimate, but only because it can be translated into the question: ‘Would a gene for making individuals do X (in this situation) become more frequent in the gene pool?’ The subjective soliloquy is justified by the fact that it can be translated into the language of gene survival.
One might be tempted to interpret ‘Should I do X or Y?’ as meaning ‘Would X or Y be more likely to prolong my own life?’ But if long life is bought at the expense of not reproducing – that is to say, if we pit individual longevity against gene survival – natural selection will not favour it. Reproduction can be a dangerous business. Male pheasants that are gorgeously coloured to attract females also attract predators. A drab, inconspicuous male would probably live longer than a brightly coloured, attractive male. But he’d be more likely to die unmated, and the genes for safety-first drabness are less likely to be passed on. Gene survival is what really matters in natural selection.
The following is a legitimate shorthand, put into the mouth of a male pheasant: ‘If I grow drab feathers I shall probably live a long time, but I won’t get a mate. If I grow bright feathers I’ll probably die young, but I’ll pass on lots of genes before I die, including the genes for making bright feathers. Therefore I should take the “decision” to grow bright feathers.’ Needless to say, ‘decision’ doesn’t mean what a human would ordinarily mean by it. Conscious thought is not involved. Organism-level shorthand can be confusing, but it works so long as you remember always to keep open the pathway to a translation back into gene language. No pheasant actually takes a ‘decision’ to grow bright feathers or drab. Instead, genes for growing bright feathers or drab have different probabilities of surviving through the generations.
It really can be helpful, when trying to understand from a Darwinian point of view what animals do, to see them as robot machines, ‘thinking’ about what steps to take in order to pass on their genes to future generations. Such steps may involve behaving in certain ways, or growing organs of a particular shape or character. It can also be helpful to think metaphorically of genes as ‘thinking’ about what steps to take in order to pass themselves on to future generations. Such steps will usually involve manipulating individual organisms via the processes of embryonic development.
But it is never even metaphorically legitimate to treat animals as thinking about what steps to take in order to preserve their species, or their group. Differential group or species survival is not what happens in natural selection. What happens is differential gene survival. Therefore, legitimate shorthands are of the form ‘If I were a gene, what would I do to preserve myself?’ Or – and ideally this should be exactly equivalent – ‘If I were an organism, what would I do to preserve my genes?’ But ‘If I were an organism, what would I do to preserve my species?’ is an illegitimate shorthand. So is – this time for a different reason – ‘If I were a species, what would I do to preserve myself?’ The latter metaphor is illegitimate because a species, unlike an individual organism, is not the kind of entity that even metaphorically behaves as an agent, doing things, acting upon decisions. Species don’t have brains and muscles, they are just collections of individual organisms that do. Species and groups are not ‘vehicles’. Individual organisms are.
I should point out that neither in my lectures of the 1960s nor in The Selfish Gene did I see as very novel the idea of the gene as the fundamental unit of natural selection. I thought of it – and clearly said so – as implicit in the orthodox neo-Darwinian theory of evolution: that is to say, the theory first clearly formalized in the 1930s by Fisher, Haldane, Wright and the other founding fathers of the so-called Modern Synthesis such as Ernst Mayr, Theodosius Dobzhansky, George Gaylord Simpson and Julian Huxley. It was only after The Selfish Gene was published that both critics and admirers came to see the idea as revolutionary. That was not how I thought of it at the time.
Having said that, however, I should add that not all the founding fathers of the Modern Synthesis were clear about this important implication of the theory that they collectively put together. To the end of his centenarian life, the authoritative German-American taxonomist Ernst Mayr expressed hostility to the idea of gene selectionism, in terms that suggested to me that he misunderstood it. And Julian Huxley, the founding father who actually coined the phrase ‘Modern Synthesis’, was an out-and-out group selectionist, without clearly realizing it. The first time I met the great Peter Medawar, he startled my student self with a deliciously sacrilegious remark, delivered with his characteristically patrician, yet impish style. ‘The trouble with Julian is that he really doesn’t understand evolution.’ Fancy saying that – of a Huxley! I could hardly believe my ears and, as you see, I have never forgotten it. I later heard another Nobel Prize-winner, the French molecular biologist Jacques Monod, say something a bit similar, though not about Huxley: ‘The trouble with natural selection is that everybody thinks he understands it.’
I mentioned that I began The Selfish Gene when power cuts interrupted my cricket research. I had completed only the first chapter of the book when I happened to meet an editor from the publishers Allen & Unwin. He was paying a routine visit to the Department of Zoology in search of possible books, and I told him about my embryonic project. He sat down and read that first chapter on the spot, liked it and encouraged me to continue. But then – unfortunately from one narrow point of view, fortunately from others – the industrial unrest came to an end and the lights came back on. I shoved my chapter in a drawer and forgot about it as I resumed my research on crickets.
From time to time during the next two years I contemplated returning to the book. The impetus was especially strong when I read and lectured about new publications that were beginning to appear in the early 1970s and proved beautifully compatible with the thesis of my gestating book. Most notable among these were papers by the young American biologist Robert Trivers, and others by the veteran British professor John Maynard Smith. Both these authors made use of the intuitive shortcut I mentioned (the philosopher Daniel Dennett would now call it an intuition pump):59 the shortcut of imagining that an individual organism behaves ‘as if’ consciously calculating the best policy for preserving and propagating its genes.
Trivers treated a parent animal as if it were a rational agent calculating what economists call the ‘opportunity cost’ of an action. A parent has to pay the costs of rearing each offspring. Among these costs might be food, including time and effort spent gathering it, time spent protecting the child from predators, and the risks incurred by the parent in doing so. Trivers wrapped them all up in one metric which he called Parental Investment or PI. Trivers’ key insight was that that PI must be an opportunity cost: the investment in any one child is measured as lost opportunities to invest in other children. Trivers used the notion to develop a penetrating theory of ‘parent–offspring conflict’. The decision on the best time to wean a child, for example, is subject to a ‘disagreement’ between the child and its mother, both behaving as rational economists whose ‘utility function’ is the long-term survival of their own genes. The mother ‘wants’ to terminate suckling earlier than the child does, because she places greater ‘value’ than he does on her future offspring, who will benefit from early weaning of the present child. The present child also ‘values’ his future siblings, but only half as highly as his mother does because of the way Hamilton’s Rule pans out. Therefore there is a period of ‘weaning conflict’, an uneasy phase of transition between the early time when both parties ‘agree’ that suckling should continue and the later time when both parties ‘agree’ that it should end. During this phase, when the mother ‘wants’ weaning but the child doesn’t, observers of animal behaviour should see the symptoms of a subtle battle between mother and child. In passing I should add that, long after The Selfish Gene was published, the Australian biologist David Haig cleverly showed how many of the ailments of pregnancy can be explained in terms of the same Triversian conflict going on inside the womb – not about weaning in this case, obviously, but about other aspects of the allocation of necessarily scarce resources.
Parent–offspring conflict was obviously a subject tailor-made for my book, and Trivers’ brilliant paper on the subject was one of the spurs that encouraged me to take my first chapter out of the drawer where it had languished since the end of the power strike. It became the inspiration for chapter 7 of The Selfish Gene, ‘Battle of the generations’. Chapter 8, ‘Battle of the sexes’, also made use of Trivers’ ideas, this time showing how males and females might compute their opportunity costs differently. When might a male, for example, desert his mate, leaving her ‘holding the baby’ and ‘in a cruel bind’ while he seeks a new one? Trivers also influenced chapter 10, ‘You scratch my back, I’ll ride on yours’. His paper in this case was an earlier one, on reciprocal altruism, which showed that kin selection is not the only evolutionary pressure towards altruism. Reciprocation – the repaying of favours – can also be very important, and it works across species, not just within them as kin selection does. So Trivers’ name was added to those of Hamilton and Williams among the four authors who had greatest influence on The Selfish Gene. I also asked him to write the foreword – which he graciously did, although at that point we had never met.
The fourth was John Maynard Smith, who later became a beloved mentor. As a boy I had met the book that he would refer to as ‘my little Penguin’, and I was much taken by the smiling author photograph: shock of nutty professor hair askew like the pipe in his mouth, thick round glasses in need of a clean – the sort of man I was immediately drawn to. I also liked the biographical note, which explained that he had been an engineer designing aircraft, but gave it up and went back to university to study biology because he noticed that ‘aircraft were noisy and old fashioned’. Many years later, a new edition of that book, The Theory of Evolution, was published by Cambridge University Press and I was honoured to be invited to write the foreword.60 I included the following tribute to this genial hero:
Readers of ‘campus novels’ know that a conference is where you can catch academics at their worst. The conference bar, in particular, is the academy in microcosm. Professors huddle together in exclusive, conspiratorial corners, talking not about science or scholarship but about ‘tenure-track hiring’ (their word for jobs) and ‘funding’ (their word for money). If they do talk shop, too often it will be to make an impression rather than to enlighten. John Maynard Smith is a splendid, triumphant, lovable exception. He values creative ideas above money, plain language above jargon. He is always the centre of a lively, laughing crowd of students and young research workers of both sexes. Never mind the lectures or the ‘workshops’; be blowed to the motor coach excursions to local beauty spots; forget your fancy visual aids and radio microphones; the only thing that really matters at a conference is that John Maynard Smith must be in residence and there must be a spacious, convivial bar. If he can’t manage the dates you have in mind, you must just reschedule the conference. He doesn’t have to give a formal talk (although he is a riveting speaker) and he doesn’t have to chair a formal session (although he is a wise, sympathetic and witty chairman). He has only to turn up and your conference will succeed. He will charm and amuse the young research workers, listen to their stories, inspire them, rekindle enthusiasms that might be flagging, and send them back to their laboratories or their muddy fields, enlivened and invigorated, eager to try out the new ideas he has generously shared with them.
My relationship with John didn’t get off to an outstandingly good start, however. I first met him in 1966 when, as Dean of Biological Sciences, he interviewed me for a job at the University of Sussex. I was already pretty much committed to going to Berkeley. However, there was a job going at Sussex and Richard Andrew, their resident expert on our shared subject of animal behaviour, pressed me with flattering urgency to apply. I told Richard about my near-commitment to go to Berkeley, but he said there was no harm in doing the Sussex interview anyway, so I thought: What the hell, why not? I’m afraid my ‘what the hell’ attitude didn’t endear me to Maynard Smith in the interview. I said I wouldn’t lecture about animal taxonomy. He said it was part of the job. I rather arrogantly said: Well, I’ve got a job offer at Berkeley and I’m not quite sure why I am doing this interview anyway. He was nice about it when he and Dr Andrew took me to lunch, but, as I said, it was not a good start to what was later to prove a delightful friendship.
In the early 1970s, Maynard Smith began the long series of papers in which, together with colleagues such as Geoffrey Parker and the late George Price, he deployed a version of the mathematical theory of games to solve a number of problems in evolution. These ideas were immensely congenial to the idea of the selfish gene, and Maynard Smith’s papers constituted the other major stimulus that led me to dust off my old chapter 1 and write the whole book.
Maynard Smith’s particular contribution was the notion of the evolutionarily stable strategy or ESS. ‘Strategy’ in this sense can be construed as ‘preprogrammed rule’. Maynard Smith set up mathematical models in which preprogrammed rules with names like (for the particular case of animal combat) Hawk, Dove, Retaliator, Bully, are let loose in an imagined (or simulated) world to interact with each other. Once again, it is important to understand that the animals implementing the rules are not assumed to be consciously aware of what they are doing, or why. Each preprogrammed rule has a frequency in the population (like genes in a gene pool, although the link with DNA doesn’t have to be made explicit in the models). The frequencies change in accordance with ‘payoffs’. In the social and economic sciences where game theory originated, payoffs can be thought of as equivalent to money. In evolutionary game theory, payoffs have the special meaning of reproductive success: high payoffs for a strategy lead to increased representation in the population.
The key point is that a successful strategy is not necessarily one that wins its particular contests against other strategies. A successful strategy is one that numerically dominates the population. And since a numerically dominant strategy is by definition likely to encounter copies of itself, it will stay numerically dominant only if it flourishes in the presence of copies of itself. This is the meaning of ‘evolutionarily stable’ in Maynard Smith’s ‘ESS’. We expect to see ESSs in nature, because if a strategy is evolutionarily unstable, it will tend to disappear from the population as rival strategies outbreed it.
I won’t expound evolutionary game theory any further here because I did that in The Selfish Gene, and the same applies to Trivers’ ideas on parental investment. Here it is sufficient to say that the publications of Trivers and Maynard Smith in the early 1970s rekindled my interest in the ideas of Hamilton that had inspired me in the 1960s, and moved me to return to the book whose first chapter had slumbered in a drawer since the end of the power strikes. Maynard Smith’s game-theoretic ideas dominated the chapter on aggression, and inspired my treatment of many topics in later chapters.
So, finally, in 1975, having finished my ‘hierarchical organization’ paper, I took the sabbatical leave to which I was entitled, stayed at home every morning, and devoted myself to my typewriter and The Selfish Gene. Indeed, so devoted was I to the task that I didn’t attend the crucial meeting at which New College was electing a new Warden. A colleague slipped out of the meeting and telephoned urgently to tell me the vote was extremely close and to beg me to come quickly. I now think that, although my sabbatical leave entitled me to do so, my absenting myself from such a crucial vote was an act of self-indulgent irresponsibility. The meeting would have taken only a few hours of my time, and the repercussions of my lost vote potentially might have been felt for many years. Fortunately, the man for whom I would have voted got in anyway (and became an excellent Warden), so I don’t have to endure a burden of guilt for changing the course of college history. Actually, his rival would have been very good too, and college meetings would certainly have been amusing as he was justly reputed to be the wittiest man in Oxford.
I wrote The Selfish Gene in a frenzy of creative energy. I’d completed three or four chapters when I spoke to my friend Desmond Morris about publication. A legendarily successful author himself, Desmond arranged a meeting with Tom Maschler, doyen of London publishers. I met Mr Maschler in his high-ceilinged, book-lined room at Jonathan Cape in London. He’d read my chapters and liked them, but urged me to change the title. ‘Selfish’, he explained to me, is a ‘down word’. Why not The Immortal Gene? With hindsight, he was very probably right. I can’t now remember why I didn’t follow his advice. I think I should have done.
I didn’t, in any case, pursue him as a publisher because matters were rather forcefully taken out of my hands. At lunch one day in New College Roger (now Sir Roger) Elliott, Oxford’s Professor of Theoretical Physics, said he had heard I was writing a book, and asked me about it. I told him a little of what I was trying to do, and he seemed interested. As it happened, he was a member of the Board of Delegates of Oxford University Press, and he passed the word to Michael Rodgers, the appropriate editor at that ancient publishing house. Michael wrote and asked to see my chapters. I sent them to him.
And then the whirlwind hit – beginning with his characteristically loud voice, over the telephone: ‘I’ve read your chapters. Haven’t been able to sleep since. I MUST HAVE THAT BOOK!’ Well, some people might resist that kind of persuasion, but not me. Michael was clearly my kind of publisher. I signed the contract and set to work with redoubled urgency to complete the book.
I now find it quite hard to comprehend how we all used to tolerate the burden of writing in the age before computer word processors. Pretty much every sentence I write is revised, fiddled with, re-ordered, crossed out and reworked. I reread my work obsessively, subjecting the text to a kind of Darwinian sieving which, I hope and believe, improves it with every pass. Even as I type a sentence for the first time, at least half the words are deleted and changed before the sentence ends. I have always worked like this. But while a computer is naturally congenial to this way of working, and the text itself remains clean with every revision, on a typewriter the result was a mess. Scissors and sticky tape were tools of the trade as important as the typewriter itself. The growing typescript of The Selfish Gene was covered with xxxxxxx deletions, handwritten insertions, words ringed and moved with arrows to other places, strips of paper inelegantly taped to the margin or the bottom of the page. One would think it a necessary part of composition that one should be able to read one’s text fluently. This would seem to be impossible when working on paper. Yet, mysteriously, writing style does not seem to have shown any general improvement since the introduction of computer word processors. Why not?
The Selfish Gene went through two fair copies typed up by Pat Searle, the motherly secretary of the Animal Behaviour Research Group. Each one went to Michael Rodgers and came back with his helpful, handwritten annotations. In particular, he excised some purple passages that my romantically youthful enthusiasm had pushed way over the top. In Peter Medawar’s metaphor of the writer as organist, ‘a scientist’s fingers, unlike a historian’s, must never stray toward the diapason’. The end of chapter 2 of The Selfish Gene is about as purple as science prose should get, and I blush to recall (and am glad I haven’t preserved) the paragraph that followed it. Here’s the paragraph of paler purple that survived Michael’s moderating pen. It’s the end of the chapter on the origin of life and the spontaneous arising in the primeval soup of ‘replicators’, which later moved into the world of ‘vehicles’ – living organisms.
Was there to be any end to the gradual improvement in the techniques and artifices used by the replicators to ensure their own continuation in the world? There would be plenty of time for improvement. What weird engines of self-preservation would the millennia bring forth? Four thousand million years on, what was to be the fate of the ancient replicators? They did not die out, for they are past masters of the survival arts. But do not look for them floating loose in the sea; they gave up that cavalier freedom long ago. Now they swarm in huge colonies, safe inside gigantic lumbering robots, sealed off from the outside world, communicating with it by tortuous indirect routes, manipulating it by remote control. They are in you and in me; they created us, body and mind; and their preservation is the ultimate rationale for our existence. They have come a long way, those replicators. Now they go by the name of genes, and we are their survival machines.
That paragraph encapsulates the central metaphor of the book, and also its science-fictiony feel. Indeed, I began my preface with the words:
This book should be read almost as though it were science fiction. It is designed to appeal to the imagination. But it is not science fiction: it is science. Cliché or not, ‘stranger than fiction’ expresses exactly how I feel about the truth. We are survival machines – robot vehicles blindly programmed to preserve the selfish molecules known as genes. This is a truth which still fills me with astonishment. Though I have known about it for years, I never seem to get fully used to it. One of my hopes is that I may have some success in astonishing others.
And the opening lines of chapter 1 continued the science fiction mood:
Intelligent life on a planet comes of age when it first works out the reason for its own existence. If superior creatures from space ever visit earth, the first question they will ask, in order to assess the level of our civilization, is: ‘Have they discovered evolution yet?’ Living organisms had existed on earth, without ever knowing why, for over three thousand million years before the truth finally dawned on one of them. His name was Charles Darwin.
Niko Tinbergen hated that opening, when the book was published and he read it. He didn’t like anything that suggested that humanity is an intelligent species, and he felt deeply wounded by the terrible effects we have had on the world. But that really wasn’t the kind of point I was making.
I should say something about the last chapter: ‘Memes: the new replicators’. Given that the rest of the book thrust the gene to centre stage as the starring replicator in the evolution of life, it was important to dispel the impression that the replicator has to be DNA. In keeping with the science fiction mood of the opening, I pointed out that on other planets the evolution of life could be fostered by a completely different system of self-replication – but that, whatever it was, it would have to have certain qualities, such as high fidelity of copying.
Casting around for an example, I could have used computer viruses if they had been invented in 1975. Instead, I lit upon human culture as a new ‘primeval soup’:
But do we have to go to distant worlds to find other kinds of replicator and other, consequent kinds of evolution? I think that a new kind of replicator has recently emerged on this very planet. It is staring us in the face. It is still in its infancy, still drifting clumsily about in its primeval soup, but already it is achieving evolutionary change at a rate which leaves the old gene panting far behind.
The new soup is the soup of human culture. We need a name for the new replicator, a noun which conveys the idea of a unit of cultural transmission, or a unit of imitation. ‘Mimeme’ comes from a suitable Greek root, but I want a monosyllable that sounds a bit like ‘gene’. I hope my classicist friends will forgive me if I abbreviate mimeme to meme. If it is any consolation, it could alternatively be thought of as being related to ‘memory’, or to the French word même. It should be pronounced to rhyme with ‘cream’.
Examples of memes are tunes, ideas, catch-phrases, clothes fashions, ways of making pots or of building arches. Just as genes propagate themselves in the gene pool by leaping from body to body via sperms or eggs, so memes propagate themselves in the meme pool by leaping from brain to brain, via a process which, in the broad sense, can be called imitation.
I went on to discuss various ways in which the idea of memes might be applied, for example to the spread and inheritance of religion. My primary intention, however, was not to make a contribution to the theory of human culture, but to downplay the gene as the only conceivable replicator that might lie at the root of a Darwinian process. I was trying to push ‘Universal Darwinism’ (the title of a later paper, based on my lecture to the 1982 conference commemorating Darwin’s death). Nevertheless I am delighted that the philosopher Daniel Dennett, the psychologist Susan Blackmore and others have run, so productively, with the meme ball. More than thirty books have now been published with the word ‘meme’ in their title, and the word has made it into the Oxford English Dictionary (whose criterion is that it must be used, without attribution or definition, in a significant number of published places).
Publication of one’s first book is a heady time for a young author. I made frequent trips to the stately neo-classical OUP building in Walton Street, and sometimes the London office in Ely House, to meet the various people involved in the complex business of production, design, marketing and so on. When it came to jacket design, the science fiction mood of the book led me again to the elegantly porticoed North Oxford door of Desmond Morris. As well as being a biologist, television personality, anthropological collector, (implausible) raconteur61 and best-selling author, Desmond is an accomplished surrealist painter. His paintings have an unmistakably biological feel. He has created a dreamscape in which other-worldly creatures live and move and have their evolution – for they do evolve, from canvas to canvas: just what was needed for The Selfish Gene. He was delighted by the idea of providing a jacket design, and Michael Rodgers and I went to look at the paintings on his walls and in his studio. The Expectant Valley stood out, not just for its bold colours and air of brooding fecundity, but also more mundanely in that it provided a convenient space to accommodate the title. We chose it with pleasure and I believe it enhanced the sales of the book.
As it happened, Desmond had an exhibition around this time in a small gallery in Walton Street near the OUP building, and The Expectant Valley was one of the paintings on sale. Its price, £750, happened to be exactly equal to the advance the publishers had paid me for my book. The coincidence was too much to resist and, after repeated visits to the gallery during which I became fond of many of the paintings, I bought The Expectant Valley. I think Desmond was a bit embarrassed, and he kindly threw in another, slightly similar painting, The Titillator. The two go rather well together.
The Selfish Gene was published in the autumn of 1976; I was thirty-five. It was reviewed widely, surprisingly so for a first work by an unknown author, and I still don’t really know why it received the attention it did. There was no launch party and no obvious fanfare organized by the publishers. Some months after publication it came to the notice of Peter Jones, one of the producers on the BBC’s ‘flagship’ science series Horizon. Peter asked me if I would like to present a documentary on the subject, but I was much too shy at that time to dare appear on television, and I recommended John Maynard Smith instead. He did a good job – he had a wonderfully warm and engaging manner – and the documentary, which had the same name, The Selfish Gene, must have given a good boost to the book’s sales, at least in Britain. But the broadcast came too late to account for the wide review coverage the book received.
I don’t do it any more, but for that first book I kept a scrap-book of reviews, and I have just been glancing at them again. There were more than 100, and a rereading doesn’t generally bear out the common perception of the book as controversial. Almost all the reviews were favourable. Among early reviewers were the psychiatrist Anthony Storr, the anthropologists Lionel Tiger and Francis Huxley (son of Julian), the naturalist Bruce Campbell and the philosopher Bernard Williams, whom I came to know much later as one of those entertaining conversationalists whose wit had the capacity to ‘raise the game’ of any companion. There were hostile reviews from two biologists identified with the political left, Steven Rose and Richard Lewontin, and – more subtly barbed – from Cyril Darlington on the opposite side of the political spectrum. But these were rare. Most of the reviewers got the message, expounded it fairly and were very nice about the book. Especially warming for me were the highly favourable reviews by Peter Medawar and W. D. Hamilton. Hamilton even hit the particular nail I had originally targeted in my quest to answer Lorenz, Ardrey and the panglossians of the 1960s and the ‘BBC Theorem’:
This book should be read, can be read, by almost everybody. It describes with great skill a new face of the theory of evolution. With much of the light, unencumbered style that has lately sold new and sometimes erroneous biology to the public, it is, in my opinion, a more serious achievement. It succeeds in the seemingly impossible task of using simple, untechnical English to present some rather recondite and quasi-mathematical themes of recent evolutionary thought. Seen through this book in their broad perspective at last, these will surprise and refresh even many research biologists who might have supposed themselves already in the know. At least, so they surprised this reviewer. Yet, to repeat, the book remains easily readable by anyone with the least grounding in science.
There was nobody in the world whom I would rather have surprised in such a way than ‘this reviewer’. I was also touched by the way Bill Hamilton ended his beautifully written review with poems, one by Wordsworth, and the other by Housman, whose Shropshire Lad I often found myself identifying with Bill’s complex personality:
From far, from eve and morning
And yon twelve-winded sky,
The stuff of life to knit me
Blew hither: here am I
. . .
Speak now, and I will answer;
How shall I help you, say;
Ere to the wind’s twelve quarters
I take my endless way.
Not a bad epitaph for an evolutionary scientist, and Bill Hamilton was probably the greatest evolutionary scientist of the second half of the twentieth century. While this volume of autobiography was in its final stages, I found a treasure among a bundle of old papers, with Bill’s handwriting at the top: it was a copy of the last page of his lecture notes, containing a rewriting of another Housman poem, ‘The Immortal Part’, to incorporate the idea of the ‘immortal’ gene. I have no memory of the lecture he is referring to, or when he gave it, and the paper is undated. I have reproduced it in the web appendix.
Long after The Selfish Gene was published, Bill became my close colleague at Oxford and I saw him almost daily at lunch in New College. I am humbly proud of the role played by my book in bringing his brilliant ideas to a wider audience. But I like to hope, too, that there are other ways in which the book changed the way my professional colleagues think about their subject. I like to think it is no accident that, if you visit a biological field station in the Serengeti, or Antarctica, or the Amazon or the Kalahari, and listen to the active researchers talking shop over their beers in the evenings, what you hear will be laced with talk of genes. It’s not that they’re talking about the molecular antics of DNA – although that is interesting too – but the underlying assumption of these conversations is that the behaviour of the animals and plants under study is aimed at preserving genes and propagating them through succeeding generations.