Authors: Richard Dawkins
* Not my favourite Yeats line, but apt in this case.
* For the sake of decorum / Pronounce it theorum.
DOGS, COWS AND CABBAGES
WHY did it take so long for a Darwin to arrive on the scene? What delayed humanity’s tumbling to that luminously simple idea which seems, on the face of it, so much easier to grasp than the mathematical ideas given us by Newton two centuries earlier – or, indeed, by Archimedes two millennia earlier? Many answers have been suggested. Perhaps minds were cowed by the sheer time it must take for great change to occur – by the mismatch between what we now call geological deep time and the lifespan and comprehension of the person trying to understand it. Perhaps it was religious indoctrination that held us back. Or perhaps it was the daunting complexity of a living organ such as an eye, freighted as it is with the beguiling illusion of design by a master engineer. Probably all those played a role. But Ernst Mayr, grand old man of the neo-Darwinian synthesis, who died in 2005 at the age of 100, repeatedly voiced a different suspicion. For Mayr, the culprit was the ancient philosophical doctrine of – to give it its modern name – essentialism. The discovery of evolution was held back by the dead hand of Plato.*
THE DEAD HAND OF PLATO
For Plato, the ‘reality’ that we think we see is just shadows cast on the wall of our cave by the flickering light of the camp fire. Like other classical Greek thinkers, Plato was at heart a geometer. Every triangle drawn in the sand is but an imperfect shadow of the true essence of triangle. The lines of the essential triangle are pure Euclidean lines with length but no breadth, lines defined as infinitely narrow and as never meeting when parallel. The angles of the essential triangle really do add up to exactly two right angles, not a picosecond of arc more or less. This is not true of a triangle drawn in the sand: but the triangle in the sand, for Plato, is but an unstable shadow of the ideal, essential triangle.
Biology, according to Mayr, is plagued by its own version of essentialism. Biological essentialism treats tapirs and rabbits, pangolins and dromedaries, as though they were triangles, rhombuses, parabolas or dodecahedrons. The rabbits that we see are wan shadows of the perfect ‘idea’ of rabbit, the ideal, essential, Platonic rabbit, hanging somewhere out in conceptual space along with all the perfect forms of geometry. Flesh-and-blood rabbits may vary, but their variations are always to be seen as flawed deviations from the ideal essence of rabbit.
How desperately unevolutionary that picture is! The Platonist regards any change in rabbits as a messy departure from the essential rabbit, and there will always be resistance to change – as if all real rabbits were tethered by an invisible elastic cord to the Essential Rabbit in the Sky. The evolutionary view of life is radically opposite. Descendants can depart indefinitely from the ancestral form, and each departure becomes a potential ancestor to future variants. Indeed, Alfred Russel Wallace, independent co-discoverer with Darwin of evolution by natural selection, actually called his paper ‘On the tendency of varieties to depart indefinitely from the original type’.
If there is a ‘standard rabbit’, the accolade denotes no more than the centre of a bell-shaped distribution of real, scurrying, leaping, variable bunnies. And the distribution shifts with time. As generations go by, there may gradually come a point, not clearly defined, when the norm of what we call rabbits will have departed so far as to deserve a different name. There is no permanent rabbitiness, no essence of rabbit hanging in the sky, just populations of furry, long-eared, coprophagous, whisker-twitching individuals, showing a statistical distribution of variation in size, shape, colour and proclivities. What used to be the longer-eared end of the old distribution may find itself the centre of a new distribution later in geological time. Given a sufficiently large number of generations, there may be no overlap between ancestral and descendant distributions: the longest ears among the ancestors may be shorter than the shortest ears among the descendants. All is fluid, as another Greek philosopher, Heraclitus, said; nothing fixed. After a hundred million years it may be hard to believe that the descendant animals ever had rabbits for ancestors. Yet in no generation during the evolutionary process was the predominant type in the population far from the modal type in the previous generation or the following generation. This way of thinking is what Mayr called population thinking. Population thinking, for him, was the antithesis of essentialism. According to Mayr, the reason Darwin was such an unconscionable time arriving on the scene was that we all – whether because of Greek influence or for some other reason – have essentialism burned into our mental DNA.
For the mind encased in Platonic blinkers, a rabbit is a rabbit is a rabbit. To suggest that rabbitkind constitutes a kind of shifting cloud of statistical averages, or that today’s typical rabbit might be different from the typical rabbit of a million years ago or the typical rabbit of a million years hence, seems to violate an internal taboo. Indeed, psychologists studying the development of language tell us that children are natural essentialists. Maybe they have to be if they are to remain sane while their developing minds divide things into discrete categories each entitled to a unique noun. It is no wonder that Adam’s first task, in the Genesis myth, was to give all the animals names.
And it is no wonder, in Mayr’s view, that we humans had to wait for our Darwin until well into the nineteenth century. To dramatize how very anti-essentialist evolution is, consider the following. On the ‘population-thinking’ evolutionary view, every animal is linked to every other animal, say rabbit to leopard, by a chain of intermediates, each so similar to the next that every link could in principle mate with its neighbours in the chain and produce fertile offspring. You can’t violate the essentialist taboo more comprehensively than that. And it is not some vague thought-experiment confined to the imagination. On the evolutionary view, there really is a series of intermediate animals connecting a rabbit to a leopard, every one of whom lived and breathed, every one of whom would have been placed in exactly the same species as its immediate neighbours on either side in the long, sliding continuum. Indeed, every one of the series was the child of its neighbour on one side and the parent of its neighbour on the other. Yet the whole series constitutes a continuous bridge from rabbit to leopard – although, as we shall see later, there never was a ‘rabbipard’. There are similar bridges from rabbit to wombat, from leopard to lobster, from every animal or plant to every other. Maybe you have reasoned for yourself why this startling result follows necessarily from the evolutionary world-view, but let me spell it out anyway. I’ll call it the hairpin thought experiment.
Take a rabbit, any female rabbit (arbitrarily stick to females, for convenience: it makes no difference to the argument). Place her mother next to her. Now place the grandmother next to the mother and so on back in time, back, back, back through the megayears, a seemingly endless line of female rabbits, each one sandwiched between her daughter and her mother. We walk along the line of rabbits, backwards in time, examining them carefully like an inspecting general. As we pace the line, we’ll eventually notice that the ancient rabbits we are passing are just a little bit different from the modern rabbits we are used to. But the rate of change will be so slow that we shan’t notice the trend from generation to generation, just as we can’t see the motion of the hour hand on our watches – and just as we can’t see a child growing, we can only see later that she has become a teenager, and later still an adult. An additional reason why we don’t notice the change in rabbits from one generation to another is that, in any one century, the variation within the current population will normally be greater than the variation between mothers and daughters. So if we try to discern the movement of the ‘hour hand’ by comparing mothers with daughters, or indeed grandmothers with granddaughters, such slight differences as we may see will be swamped by the differences among the rabbits’ friends and relations gambolling in the meadows round about.
Nevertheless, steadily and imperceptibly, as we retreat through time, we shall reach ancestors that look less and less like a rabbit and more and more like a shrew (and not very like either). One of these creatures I’ll call the hairpin bend, for reasons that will become apparent. This animal is the most recent common ancestor (in the female line, but that is not important) that rabbits share with leopards. We don’t know exactly what it looked like, but it follows from the evolutionary view that it definitely had to exist. Like all animals, it was a member of the same species as its daughters and its mother. We now continue our walk, except that we have turned the bend in the hairpin and are walking forwards in time, aiming towards the leopards (among the hairpin’s many and diverse descendants, for we shall continually meet forks in the line, where we consistently choose the fork that will eventually lead to leopards). Each shrew-like animal along our forward walk is now followed by her daughter. Slowly, by imperceptible degrees, the shrew-like animals will change, through intermediates that might not resemble any modern animal much but strongly resemble each other, perhaps passing through vaguely stoat-like intermediates, until eventually, without ever noticing an abrupt change of any kind, we arrive at a leopard.
Various things must be said about this thought experiment. First, we happen to have chosen to walk from rabbit to leopard, but I repeat that we could have chosen porcupine to dolphin, wallaby to giraffe or human to haddock. The point is that for any two animals there has to be a hairpin path linking them, for the simple reason that every species shares an ancestor with every other species: all we have to do is walk backwards from one species to the shared ancestor, then turn through a hairpin bend and walk forwards to the other species.
Second, notice that we are talking only about locating a chain of animals that links a modern animal to another modern animal. We are most emphatically not evolving a rabbit into a leopard. I suppose you could say we are de- evolving back to the hairpin, then evolving forwards to the leopard from there. As we’ll see in a later chapter, it is unfortunately necessary to explain, again and again, that modern species don’t evolve into other modern species, they just share ancestors: they are cousins. This, as we shall see, is also the answer to that disquietingly common plaint: ‘If humans have evolved from chimpanzees, how come there are still chimpanzees around?’
Third, on our forward march from the hairpin animal, we arbitrarily choose the path leading to the leopard. This is a real path of evolutionary history, but, to repeat this important point, we choose to ignore numerous branch points where we could have followed evolution to countless other end points – for the hairpin animal is the grand ancestor not only of rabbits and leopards but of a large fraction of modern mammals.
The fourth point, which I have already emphasized, is that, however radical and extensive the differences between the ends of the hairpin – rabbit and leopard, say – each step along the chain that links them is very, very small. Every individual along the chain is as similar to its neighbours in the chain as mothers and daughters are expected to be. And more similar to its neighbours in the chain, as I have also mentioned, than to typical members of the surrounding population.
You can see how this thought experiment drives a coach and horses through the elegant Greek temple of Platonic ideal forms. And you can see how, if Mayr is right that humans are deeply imbued with essentialist preconceptions, he might well also be right about why we historically found evolution so hard to stomach.
The word ‘essentialism’ itself wasn’t invented till 1945 and so was not available to Darwin. But he was only too familiar with the biological version of it in the form of the ‘immutability of species’, and much of his effort was directed towards combating it under that name. Indeed, in several of Darwin’s books – more so in others than On the Origin of Species itself – you’ll understand fully what he’s on about only if you shed modern presuppositions about evolution, and remember that a large part of his audience would have been essentialists who never doubted the immutability of species. One of Darwin’s most telling weapons in arguing against this supposed immutability was the evidence from domestication, and it is domestication that will occupy the rest of this chapter.
Sculpting the gene pool
Darwin knew plenty about animal and plant breeding. He communed with pigeon fanciers and horticulturalists, and he loved dogs.* Not only is the first chapter of On the
Origin of Species all about domestic varieties of animals and plants; Darwin also wrote a whole book on the subject. The Variation of Animals and Plants under Domestication has chapters on dogs and cats, horses and asses, pigs, cattle, sheep and goats, rabbits, pigeons (two chapters; pigeons were a particular love of Darwin), chickens and various other birds, and plants, including the amazing cabbages. Cabbages are a vegetable affront to essentialism and the immutability of species. The wild cabbage, Brassica oleracea, is an undistinguished plant, vaguely like a weedy version of a domestic cabbage. In just a few centuries, wielding the fine and coarse chisels furnished by the toolbox of selective breeding techniques, horticulturalists have sculpted this rather nondescript plant into vegetables as strikingly different from each other and from the wild ancestor as broccoli, cauliflower, kohlrabi, kale, Brussels sprouts, spring greens, romanescu and, of course, the various kinds of vegetables that are still commonly called cabbage.
Another familiar example is the sculpting of the wolf, Canis lupus, into the two hundred or so breeds of dog, Canis familiaris, that are recognized as separate by the UK Kennel Club, and the larger number of breeds that are genetically isolated from one another by the apartheid-like rules of pedigree breeding.
Incidentally, the wild ancestor of all domestic dogs really does seem to be the wolf and only the wolf (although its domestication may have happened independently in different places around the world). Evolutionists haven’t always thought so. Darwin, along with many of his contemporaries, suspected that several species of wild canid, including wolves and jackals, had contributed ancestry to our domestic dogs. The Nobel Prize-winning Austrian ethologist Konrad Lorenz was of the same view. His Man Meets Dog, published in 1949, pushes the notion that domestic dog breeds fall into two main groups: those derived from jackals (the majority) and those derived from wolves (Lorenz’s own favourites, including Chows). Lorenz seems to have had no evidence at all for his dichotomy, other than the differences that he thought he saw in the personalities and characters of the breeds. The matter remained open until molecular genetic evidence came along to clinch it. There is now no doubt. Domestic dogs have no jackal ancestry at all. All breeds of dogs are modified wolves: not jackals, not coyotes and not foxes.
The main point I want to draw out of domestication is its astonishing power to change the shape and behaviour of wild animals, and the speed with which it does so. Breeders are almost like modellers with endlessly malleable clay, or like sculptors wielding chisels, carving dogs or horses, or cows or cabbages, to their whim. I shall return to this image shortly. The relevance to natural evolution is that, although the selecting agent is man and not nature, the process is otherwise exactly the same. This is why Darwin gave so much prominence to domestication at the beginning of On the Origin of Species. Anybody can understand the principle of evolution by artificial selection. Natural selection is the same, with one minor detail changed.
Strictly speaking, it is not the body of the dog or the cabbage that is carved by the breeder/sculptor but the gene pool of the breed or species. The idea of a gene pool is central to the body of knowledge and theory that goes under the name of the ‘Neo-Darwinian Synthesis’. Darwin himself knew nothing of it. It was not a part of his intellectual world, nor indeed were genes. He was aware, of course, that characteristics run in families; aware that offspring tend to resemble their parents and siblings; aware that particular characteristics of dogs and pigeons breed true. Heredity was a central plank of his theory of natural selection. But a gene pool is something else. The concept of a gene pool has meaning only in the light of Mendel’s law of the independent assortment of hereditary particles. Darwin never knew Mendel’s laws, for although Gregor Mendel, the Austrian monk who was the father of genetics, was Darwin’s contemporary, he published his findings in a German journal which Darwin never saw.
A Mendelian gene is an all-or-nothing entity. When you were conceived, what you received from your father was not a substance, to be mixed with what you received from your mother as if mixing blue paint and red paint to make purple. If this were really how heredity worked (as people vaguely thought in Darwin’s time) we’d all be a middling average, halfway between our two parents. In that case, all variation would rapidly disappear from the population (no matter how assiduously you mix purple paint with purple paint, you’ll never reconstitute the original red and blue). In fact, of course, anybody can plainly see that there is no such intrinsic tendency for variation to decrease in a population. Mendel showed that this is because when paternal genes and maternal genes are combined in a child (he didn’t use the word ‘gene’, which wasn’t coined until 1909), it is not like blending paints, it is more like shuffling and reshuffling cards in a pack. Nowadays, we know that genes are lengths of DNA code, not physically separate like cards, but the principle remains valid. Genes don’t blend; they shuffle. You could say they are shuffled badly, with groups of cards sticking together for several generations of shuffling before chance happens to split them.
Any one of your eggs (or sperms if you are male) contains either your father’s version of a particular gene or your mother’s version, not a blend of the two. And that particular gene came from one and only one of your four grandparents; and from one and only one of your eight great-grandparents.*
Hindsight says this should have been obvious all along. When you cross a male with a female, you expect to get a son or a daughter, not a hermaphrodite.† Hindsight says anybody in an armchair could have generalized the same all-or-none principle to the inheritance of each and every characteristic. Fascinatingly, Darwin himself was glimmeringly close to this, but he stopped just short of making the full connection. In 1866 he wrote, in a letter to Alfred Wallace:My dear Wallace
I do not think you understand what I mean by the non-blending of certain varieties. It does not refer to fertility. An instance will explain. I crossed the Painted Lady and Purple sweet peas, which are very differently coloured varieties, and got, even out of the same pod, both varieties perfect but none intermediate. Something of this kind, I should think, must occur at first with your butterflies . . . Though these cases are in appearance so wonderful, I do not know that they are really more so than every female in the world producing distinct male and female offspring.
Darwin came that close to discovering Mendel’s law of the non-blending of (what we would now call) genes.* The case is analogous to the claim, by various aggrieved apologists, that other Victorian scientists, for example Patrick Matthew and Edward Blyth, had discovered natural selection before Darwin did. In a sense that is true, as Darwin acknowledged, but I think the evidence shows that they didn’t understand how important it is. Unlike Darwin and Wallace, they didn’t see it as a general phenomenon with universal significance – with the power to drive the evolution of all living things in the direction of positive improvement. In the same way, this letter to Wallace shows that Darwin got tantalizingly close to grasping the point about the non-blending nature of heredity. But he didn’t see its generality, and in particular he failed to see it as the answer to the riddle of why variation didn’t automatically disappear from populations. That was left to twentieth-century scientists, building on Mendel’s before-his-time discovery.†
So now the concept of the gene pool starts to make sense. A sexually reproducing population, such as, say, all the rats on Ascension Island, remotely isolated in the South Atlantic, is continually shuffling all the genes on the island. There is no intrinsic tendency for each generation to become less variable than the previous generation, no tendency towards ever more boringly grey, middling intermediates. The genes remain intact, shuffled about from individual body to individual body as the generations go by, but not blending with one another, never contaminating each other. At any one time, the genes are all sitting in the bodies of individual rats, or they are moving into new rat bodies via sperms. But if we take a long view across many generations, we see all the rat genes on the island being mixed up as though they were cards in a single well-shuffled pack: one single pool of genes.
I’m guessing that the rat gene pool on a small and isolated island such as Ascension is a self-contained and rather well-stirred pool, in the sense that the recent ancestors of any one rat could have lived anywhere on the island, but probably not anywhere other than on the island, give or take the occasional stowaway on a ship. But the gene pool of the rats on a large land mass such as Eurasia would be much more complicated. A rat living in Madrid would derive most of its genes from ancestors living in the western end of the Eurasian continent rather than, say, Mongolia or Siberia, not because of specific barriers to gene flow (though those exist too) but because of the sheer distances involved. It takes time for sexual shuffling to work a gene from one side of a continent to the other. Even if there are no physical barriers such as rivers or mountain ranges, gene flow across such a large land mass will still be slow enough for the gene pool to deserve the name ‘viscous’. A rat living in Vladivostok would trace most of its genes back to ancestors in the east. The Eurasian gene pool would be shuffled, as on Ascension Island, but not homogeneously shuffled because of the distances involved. Moreover, partial barriers such as mountain ranges, large rivers or deserts would further get in the way of homogeneous shuffling, thereby structuring and complicating the gene pool. These complications don’t devalue the idea of the gene pool. The perfectly stirred gene pool is a useful abstraction, like a mathematician’s abstraction of a perfect straight line. Real gene pools, even on small islands like Ascension, are imperfect approximations, only partially shuffled. The smaller and less broken-up the island, the better the approximation to the abstract ideal of the perfectly stirred gene pool.
Just to round off the thought about gene pools, each individual animal that we see in a population is a sampling of the gene pool of its time (or rather its parents’ time). There is no intrinsic tendency in gene pools for particular genes to increase or decrease in frequency. But when there is a systematic increase or decrease in the frequency with which we see a particular gene in a gene pool, that is precisely and exactly what is meant by evolution. The question, therefore, becomes: why should there be a systematic increase or decrease in a gene’s frequency? That, of course, is where things start to get interesting, and we shall come to it in due course.
Something funny happens to the gene pools of domestic dogs. Breeders of pedigree Pekineses or Dalmatians go to elaborate lengths to stop genes crossing from one gene pool to another. Stud books are kept, going back many generations, and miscegenation is the worst thing that can happen in the book of a pedigree breeder. It is as though each breed of dog were incarcerated on its own little Ascension Island, kept apart from every other breed. But the barrier to interbreeding is not blue water but human rules. Geographically the breeds all overlap, but they might as well be on separate islands because of the way their owners police their mating opportunities. Of course, from time to time the rules are broken. Like a rat stowing away on a ship to Ascension Island, a whippet bitch, say, escapes the leash and mates with a spaniel. But the mongrel puppies that result, however loved they may be as individuals, are cast off the island labelled Pedigree Whippet. The island itself remains a pure whippet island. Other pure-bred whippets ensure that the gene pool of the virtual island labelled Whippet continues uncontaminated. There are hundreds of man-made ‘islands’, one for each breed of pedigree dog. Each one is a virtual island, in the sense that it is not geographically localized. Pedigree whippets or Pomeranians are to be found in many different places around the world, and cars, ships and planes are used to ferry the genes from one geographical place to another. The virtual genetic island that is the Pekinese gene pool overlaps geographically, but not genetically (except when a bitch breaks cover), with the virtual genetic island that is the boxer gene pool and the virtual island that is the St Bernard gene pool.
Now let’s return to the remark that opened my discussion of gene pools. I said that if human breeders are to be seen as sculptors, what they are carving with their chisels is not dog flesh but gene pools. It appears to be dog flesh because the breeder might announce an intention to, say, shorten the snouts of future generations of boxers. And the end product of such an intention would indeed be a shorter snout, as though a chisel had been taken to the ancestor’s face. But, as we have seen, a typical boxer in any one generation is a sampling of the contemporary gene pool. It is the gene pool that has been carved and whittled over the years. Genes for long snouts have been chiselled out of the gene pool and replaced by genes for short snouts. Every breed of dog, from dachshund to Dalmatian, from boxer to borzoi, from poodle to Pekinese, from Great Dane to chihuahua, has been carved, chiselled, kneaded, moulded, not literally as flesh and bone but in its gene pool.
It isn’t all done by carving. Many of our familiar breeds of dog were originally derived as hybrids of other breeds, often quite recently, for example in the nineteenth century. Hybridization, of course, represents a deliberate violation of the isolation of the gene pools on virtual islands. Some hybridization schemes are designed with such care that the breeders would resent their products being described as mongrels or mutts (as President Obama delightfully described himself). The ‘Labradoodle’ is a hybrid between a standard poodle and a Labrador retriever, the result of a carefully crafted quest for the best virtues of both breeds. Labradoodle owners have established societies and associations just like those of pure-bred pedigree dogs. There are two schools of thought in the Labradoodle Fancy, and those of other such designer hybrids. There are those who are happy to go on making Labradoodles by mating poodles and Labradors together. And there are those who are trying to initiate a new Labradoodle gene pool that will breed true, when Labradoodles are mated together. At present, second-generation Labradoodle genes recombine to produce more variety than pure-bred pedigree dogs are supposed to show. This is how many ‘pure’ breeds got their start: they went through an intermediate stage of high variation, subsequently trimmed down through generations of careful breeding.
Sometimes, new breeds of dog get their start with the adoption of a single major mutation. Mutations are the random changes in genes that constitute the raw material for evolution by non-random selection. In nature, large mutations seldom survive, but geneticists like them in the laboratory because they are easy to study. Breeds of dog with very short legs, like basset hounds and dachshunds, acquired them in a single step with the genetic mutation called achondroplasia, a classic example of a large mutation that would be unlikely to survive in nature. A similar mutation is responsible for the commonest kind of human dwarfism: the trunk is of nearly normal size, but the legs and arms are short. Other genetic routes produce miniature breeds that retain the proportions of the original. Dog breeders can achieve changes in size and shape by selecting combinations of a few major mutations such as achondroplasia and lots of minor genes. Nor do they need to understand the genetics in order to achieve change effectively. Without any understanding at all, just by choosing who mates with whom, you can breed for all kinds of desired characteristics. This is what dog breeders, and animal and plant breeders generally, achieved for centuries before anybody understood anything about genetics. And there’s a lesson in that about natural selection, for nature, of course, has no understanding or awareness of anything at all.
The American zoologist Raymond Coppinger makes the point that puppies of different breeds are much more similar to each other than adult dogs are. Puppies can’t afford to be different, because the main thing they have to do is suck,* and sucking presents pretty much the same challenges for all breeds. In particular, in order to be good at sucking, a puppy can’t have a long snout like a borzoi or a retriever. That’s why all puppies look like pugs. You could say that an adult pug is a puppy whose face didn’t properly grow up. Most dogs, after they are weaned, develop a relatively longer snout. Pugs, bulldogs and Pekineses don’t; they grow in other departments, while the snout retains its infantile proportions. The technical term for this is neoteny, and we’ll meet it again when we come on to human evolution in Chapter 7.
If an animal grows at the same rate in all its parts, so that the adult is just a uniformly inflated replica of the infant, it is said to grow isometrically. Isometric growth is quite rare. In allometric growth, by contrast, different parts grow at different rates. Often, the rates of growth of different parts of an animal bear some simple mathematical relation to each other, a phenomenon that was investigated especially by Sir Julian Huxley in the 1930s. Different breeds of dog achieve their different shapes by means of genes that change the allometric growth relationships between the parts of the body. For example, bulldogs get their Churchillian scowl from a genetic tendency towards slower growth of the nasal bones. This has knock-on effects on the relative growth of the surrounding bones, and indeed all the surrounding tissues. One of these knock-on effects is that the palate is pulled up into an awkward position, so the bulldog’s teeth stick out and it has a tendency to dribble. Bulldogs also have breathing difficulties, which are shared by Pekineses. Bulldogs even have difficulty being born because the head is disproportionately big. Most if not all the bulldogs you see today were born by caesarian section.
Borzois are the opposite. They have extra long snouts. Indeed, they are unusual in that the elongation of the snout begins before they are born, which probably makes borzoi puppies less proficient suckers than other breeds. Coppinger speculates that the human desire to breed borzois for long snouts has reached a limit imposed by the survival capacity of puppies trying to suck.
What lessons do we learn from the domestication of the dog? First, the great variety among breeds of dogs, from Great Danes to Yorkies, from Scotties to Airedales, from ridgebacks to dachshunds, from whippets to St Bernards, demonstrates how easy it is for the non-random selection of genes – the ‘carving and whittling’ of gene pools – to produce truly dramatic changes in anatomy and behaviour, and so fast. Surprisingly few genes may be involved. Yet the changes are so large – the differences between breeds so dramatic – that you might expect their evolution to take millions of years instead of just a matter of centuries. If so much evolutionary change can be achieved in just a few centuries or even decades, just think what might be achieved in ten or a hundred million years.
Viewing the process over centuries, it is no empty fancy that human dog breeders have seized dog flesh like modelling clay and pushed it, pulled it, kneaded it into shape, more or less at will. Of course, as I pointed out earlier, we have really been kneading not dog flesh but dog gene pools. And ‘carved’ is a better metaphor than ‘kneaded’. Some sculptors work by taking a lump of clay and kneading it into shape. Others take a lump of stone or wood, and carve it by subtracting bits with a chisel. Obviously dog fanciers don’t carve dogs into shape by subtracting bits of dog flesh. But they do something close to carving dog gene pools by subtraction. It is more complicated than pure subtraction, however. Michelangelo took a single chunk of marble, and then subtracted marble from it to reveal David lurking inside. Nothing was added. Gene pools, on the other hand, are continually added to, for example by mutation, while at the same time non-random death subtracts. The analogy to sculpture breaks down here, and should not be pushed too tenaciously, as we’ll see again in Chapter 8.
The idea of sculpture calls to mind the over-muscled physiques of human body-builders, and non-human equivalents such as the Belgian Blue breed of cattle. This walking beef factory has been contrived via a particular genetic alteration called ‘double muscling’. There is a substance called myostatin, which limits muscle growth. If the gene that makes myostatin is disabled, muscles grow larger than usual. It is quite often the case that a given gene can mutate in more than one way to produce the same outcome, and indeed there are various ways in which the myostatin-producing gene can be disabled, with the same effect. Another example is the breed of pig called the Black Exotic, and there are individual dogs of various breeds that show the same exaggerated musculature for the same reason. Human body-builders achieve a similar physique by an extreme regime of exercise, and often by the use of anabolic steroids: both environmental manipulations that mimic the genes of the Belgian Blue and the Black Exotic. The end result is the same, and that is a lesson in itself. Genetic and environmental changes can produce identical outcomes. If you wanted to rear a human child to win a body-building contest and you had a few centuries to spare, you could start by genetic manipulation, engineering exactly the same freak gene as characterizes Belgian Blue cattle and Black Exotic pigs. Indeed, there are some humans known to have deletions of the myostatin gene, and they tend to be abnormally well muscled. If you started with a mutant child and made it pump iron as well (presumably the cattle and pigs could not be cajoled into this), you could probably end up with something more grotesque than Mr Universe.
Political opposition to eugenic breeding of humans sometimes spills over into the almost certainly false assertion that it is impossible. Not only is it immoral, you may hear it said, it wouldn’t work. Unfortunately, to say that something is morally wrong, or politically undesirable, is not to say that it wouldn’t work. I have no doubt that, if you set your mind to it and had enough time and enough political power, you could breed a race of superior body-builders, or high-jumpers, or shot-putters; pearl fishers, sumo wrestlers, or sprinters; or (I suspect, although now with less confidence because there are no animal precedents) superior musicians, poets, mathematicians or wine-tasters. The reason I am confident about selective breeding for athletic prowess is that the qualities needed are so similar to those that demonstrably work in the breeding of racehorses and carthorses, of greyhounds and sledge dogs. The reason I am still pretty confident about the practical feasibility (though not the moral or political desirability) of selective breeding for mental or otherwise uniquely human traits is that there are so few examples where an attempt at selective breeding in animals has ever failed, even for traits that might have been thought surprising. Who would have thought, for example, that dogs could be bred for sheep-herding skills, or ‘pointing’, or bull-baiting?
You want high milk yield in cows, orders of magnitude more gallons than could ever be needed by a mother to rear her babies? Selective breeding can give it to you. Cows can be modified to grow vast and ungainly udders, and these continue to yield copious quantities of milk indefinitely, long after the normal weaning period of a calf. As it happens, dairy horses have not been bred in this way, but will anyone contest my bet that we could do it if we tried? And of course, the same would be true of dairy humans, if anyone wanted to try. All too many women, bamboozled by the myth that breasts like melons are attractive, pay surgeons large sums of money to implant silicone, with (for my money) unappealing results. Does anyone doubt that, given enough generations, the same deformity could be achieved by selective breeding, after the manner of Friesian cows?
About twenty-five years ago I developed a computer simulation to illustrate the power of artificial selection: a kind of computer game equivalent to breeding prize roses or dogs or cattle. The player is faced with an array of nine shapes on the screen – ‘computer biomorphs’ – the middle one of which is the ‘parent’ of the surrounding eight. All the shapes are constructed under the influence of a dozen or so ‘genes’, which are simply numbers handed down from ‘parent’ to ‘offspring’, with the possibility of small ‘mutations’ intervening on the way. A mutation is just a slight increment or decrement in the numerical value of the parent’s gene. Each shape is constructed under the influence of a particular set of numbers, which are its own particular values of the dozen genes. The player looks over the array of nine shapes and sees no genes but chooses the preferred ‘body’ shape she wants to breed from. The other eight biomorphs disappear from the screen, the chosen one glides to the centre, and ‘spawns’ eight new mutant ‘children’. The process repeats for as many ‘generations’ as the player has time for, and the average shape of the ‘organisms’ on the screen gradually ‘evolves’ as the generations go by. Only genes are passed from generation to generation, so, by directly choosing biomorphs by eye, the player is inadvertently choosing genes. That is just what happens when breeders choose dogs or roses to breed from.