There is evidence that, in living species too, ‘fossil genes’ occasionally come into their own again, and are re-used after lying dormant for a million years or so. To go into detail would carry us too far from the main pathway of this chapter, for you will remember that we are already out on a digression. The main point was that the total genetic capacity of a species may increase due to gene duplication. Reusing of old ‘fossil’ copies of existing genes is one way in which this can happen. There are other, more immediate, ways in which genes may be copied to widely distributed parts of the chromosomes, like files being duplicated to different parts of a disc, or different discs.
Humans have eight separate genes called globin genes (used for making haemoglobin, among other things), on various different chromosomes. It seems certain that all eight have been copied, ultimately from a single ancestral globin gene. About 1,100 million years ago, the ancestral globin gene duplicated, forming two genes. We can date this event because of independent evidence about how fast globins habitually evolve (see Chapters 5 and 11). Of the two genes produced by this original duplication, one became the ancestor of all the genes that make haemoglobin in vertebrates. The other became the ancestor of all the genes that make myoglobins, a related family of proteins that work in muscles. Various subsequent duplications have given rise to the so-called alpha, beta, gamma, delta, epsilon and zeta globins. The fascinating thing is that we can construct a complete family tree of all the globin genes, and even put dates on all the divergence points (delta and beta globin parted company, for example, about 40 million years ago; epsilon and gamma globins 100 million years ago). Yet the eight globins, descendants as they are of these remote branchings in distant ancestors, are still all present inside every one of us. They diverged to different parts of an ancestor’s chromosomes, and we have each inherited them on our different chromosomes. Molecules are sharing the same body with their remote molecular cousins. It is certain that a great deal of such duplication has gone on, all over the chromosomes, and throughout geological time. This is an important respect in which real life is more complicated than the biomorphs of Chapter 3. They all had only nine genes. They evolved by changes in those nine genes, never by increasing the number of genes to ten. Even in real animals, such duplications are rare enough not to invalidate my general statement that all members of a species share the same DNA ‘addressing’ system.
Duplication within the species isn’t the only means by which the number of cooperating genes has increased in evolution. An even rarer, but still possibly very important occurrence, is the occasional incorporation of a gene from another species, even an extremely remote species. There are, for example, haemoglobins in the roots of plants of the pea family. They don’t occur in any other plant families, and it seems almost certain that they somehow got into the pea family by cross-infection from animals, viruses perhaps acting as intermediaries.
An especially important event along these lines, according to the increasingly favoured theory of the American biologist Lynn Margulis, took place at the origin of the so-called eukaryotic cell. Eukaryotic cells include all cells except those of bacteria. The living world is divided, fundamentally, into bacteria versus the rest. We are part of the rest, and are collectively called the eukaryotes. We differ from bacteria mainly in that our cells have discrete little mini-cells inside them. These include the nucleus, which houses the chromosomes; the tiny bomb-shaped objects called mitochondria (which we briefly met in Figure 1), filled with intricately folded membranes; and, in the (eukaryotic) cells of plants, chloroplasts. Mitochondria and chloroplasts have their own DNA, which replicates and propagates itself entirely independently of the main DNA in the chromosomes of the nucleus. All the mitochondria in you are descended from the small population of mitochondria that travelled from your mother in her egg. Sperms are too small to contain mitochondria, so mitochondria travel exclusively down the female line, and male bodies are dead ends as far as mitochondria reproduction is concerned. Incidentally, this means that we can use mitochondria to trace our ancestry, strictly down the female line.
Margulis’s theory is that mitochondria and chloroplasts, and a few other structures inside cells, are each descended from bacteria. The eukaryotic cell was formed, perhaps 2 billion years ago, when several kinds of bacteria joined forces because of the benefits that each could obtain from the others. Over the aeons they have become so thoroughly integrated into the cooperative unit that became the eukaryotic cell, that it has become almost impossible to detect the fact, if indeed it is a fact, that they were once separate bacteria.
It seems that, once the eukaryotic cell had been invented, a whole new range of designs became possible. Most interestingly from our point of view, cells could manufacture large bodies comprising many billions of cells. All cells reproduce by splitting into two, both halves getting a full set of genes. As we saw in the case of the bacteria on a pin’s head, successive splittings into two can generate a very large number of cells in rather a short time. You start with one and it splits into two. Then each of the two splits, making four. Each of the four splits, making eight. The numbers go up by successive doublings, from 8 to 16, 32, 64, 128, 256, 512, 1,024, 2,048, 4,096, 8,192. After only 20 doublings, which doesn’t take very long, we are up in the millions. After only 40 doublings the number of cells is more than a trillion. In the case of bacteria, the enormous numbers of cells produced by successive doublings go their separate ways. The same is true of many eukaryotic cells, for instance protozoa such as amoebas. A major step in evolution was taken when cells that had been produced by successive splittings stuck together instead of going off independently. Higher-order structure could now emerge, just as it did, on an incomparably smaller scale, in the two-way branching computer biomorphs.
Now, for the first time, large body size became a possibility. A human body is a truly colossal population of cells, all descended from one ancestor, the fertilized egg; and all therefore cousins, children, grandchildren, uncles, etc. of other cells in the body. The 10 trillion cells that make up each one of us are the product of a few dozens of generations of cell doublings. These cells are classified into about 210 (according to taste) different kinds, all built by the same set of genes but with different members of the set of genes turned on in different kinds of cells. This, as we have seen, is why liver cells are different from brain cells, and bone cells are different from muscle cells.
Genes working through the organs and behaviour patterns of many-celled bodies can achieve methods of ensuring their own propagation that are not available to single cells working on their own. Many-celled bodies make it possible for genes to manipulate the world, using tools built on a scale that is orders of magnitude larger than the scale of single cells. They achieve these large-scale indirect manipulations via their more direct effects on the miniature scale of cells. For instance, they change the shape of the cell membrane. The cells then interact with one another in huge populations to produce large-scale group effects such as an arm or a leg or (more indirectly) a beaver’s dam. Most of the properties of an organism that we are equipped to see with our naked eyes are so-called ‘emergent properties’. Even the computer biomorphs, with their nine genes, had emergent properties. In real animals they are produced at the whole-body level by interactions between cells. An organism works as an entire unit, and its genes can be said to have effects on the whole organism, even though each copy of any one gene exerts its immediate effects only within its own cell.
We have seen that a very important part of a gene’s environment is the other genes that it is likely to meet in successive bodies as the generations go by. These are the genes that are permuted and combined within the species. Indeed, a sexually reproducing species can be thought of as a device that permutes a discrete set of mutually accustomed genes in different combinations. Species, according to this view, are continually shuffling collections of genes that meet each other within the species, but never meet genes of other species. But there is a sense in
which the genes of different species, even if they don’t meet at close quarters inside cells, nevertheless constitute an important part of each other’s environment. The relationship is often hostile rather than cooperative, but this can be treated as just a reversal of sign. This is where we come to the second major theme of this chapter, ‘arms races’. There are arms races between predators and prey, parasites and hosts, even — though the point is a more subtle one and I shan’t discuss it further — between males and females within one species.
Arms races are run in evolutionary time, rather than on the timescale of individual lifetimes. They consist of the improvement in one lineage’s (say prey animals’) equipment to survive, as a direct consequence of improvement in another (say predators’) lineage’s evolving equipment. There are arms races wherever individuals have enemies with their own capacity for evolutionary improvement. I regard arms races as of the utmost importance because it is largely arms races that have injected such ‘progressiveness’ as there is in evolution. For, contrary to earlier prejudices, there is nothing inherently progressive about evolution. We can see this if we consider what would have happened if the only problems animals had had to face had been those posed by the weather and other aspects of the nonliving environment.
After many generations of cumulative selection in a particular place, the local animals and plants become well fitted to the conditions, for instance the weather conditions, in that place. If it is cold the animals come to have thick coats of hair, or feathers. If it is dry they evolve leathery or waxy waterproof skins to conserve what little water there is. The adaptations to local conditions affect every part of the body, its shape and colour, its internal organs, its behaviour, and the chemistry in its cells.
If the conditions in which a lineage of animals lives remain constant; say it is dry and hot and has been so without a break for 100 generations, evolution in that lineage is likely to come to a halt, at least as far as adaptations to temperature and humidity are concerned. The animals will become as well fitted as they can be to the local conditions. This doesn’t mean that they couldn’t be completely redesigned to be even better. It does mean that they can’t improve themselves by any small (and therefore likely) evolutionary step: none of their immediate neighbours in the local equivalent of ‘biomorph space’ would do any better.
Evolution will come to a standstill until something in the conditions changes: the onset of an ice age, a change in the average rainfall of the area, a shift in the prevailing wind. Such changes do happen when we are dealing with a timescale as long as the evolutionary one. As a consequence, evolution normally does not come to a halt, but constantly ‘tracks’ the changing environment. If there is a steady downward drift in the average temperature in the area, a drift that persists over centuries, successive generations of animals will be propelled by a steady selection ‘pressure’ in the direction, say, of growing longer coats of hair. If, after a few thousand years of reduced temperature the trend reverses and average temperatures creep up again, the animals will come under the influence of a new selection pressure, and will be pushed towards growing shorter coats again.
But so far we have considered only a limited part of the environment, namely the weather. The weather is very important to animals and plants. Its patterns change as the centuries go by, so this keeps evolution constantly in motion as it ‘tracks’ the changes. But weather patterns change in a haphazard, inconsistent way. There are other parts of an animal’s environment that change in more consistently malevolent directions, and that also need to be ‘tracked’. These parts of the environment are living things themselves. For a predator such as a hyena, a part of its environment that is at least as important as the weather is its prey, the changing populations of gnus, zebras and antelopes. For the antelopes and other grazers that wander the plains in search of grass, the weather may be important, but the lions, hyenas and other carnivores are important too. Cumulative selection will see to it that animals are well fitted to outrun their predators or outwit their prey, no less than it sees to it that they are well fitted to the prevailing weather conditions. And, just as long-term fluctuations in the weather are ‘tracked’ by evolution, so long-term changes in the habits or weaponry of predators will be tracked by evolutionary changes in their prey. And vice versa, of course.
We can use the general term ‘enemies’ of a species, to mean other living things that work to make life difficult. Lions are enemies of zebras. It may seem a little callous to reverse the statement to ‘Zebras are enemies of lions’. The role of the zebra in the relationship seems too innocent and wronged to warrant the pejorative ‘enemy’. But individual zebras do everything in their power to resist being eaten by lions, and from the lions’ point of view this is making life harder for them. If zebras and other grazers all succeeded in their aim, the lions would die of starvation. So by our definition zebras are enemies of lions. Parasites such as tapeworms are enemies of their hosts, and hosts are enemies of parasites since they tend to evolve measures to resist them. Herbivores are enemies of plants, and plants are enemies of herbivores, to the extent that they manufacture thorns, and poisonous or nasty-tasting chemicals.
Lineages of animals and plants will, in evolutionary time, ‘track’ changes in their enemies no less assiduously than they track changes in average weather conditions. Evolutionary improvements in cheetah weaponry and tactics are, from the gazelles’ point of view, like a steady worsening of the climate, and they are tracked in the same kind of way. But there is one enormously important difference between the two. The weather changes over the centuries, but it does not change in a specifically malevolent way. It is not out to ‘get’ gazelles. The average cheetah will change over the centuries, just like the mean annual rainfall changes. But whereas mean annual rainfall will drift up and down, with no particular rhyme or reason, the average cheetah, as the centuries go by, will tend to become better equipped to catch gazelles than his ancestors were. This is because the succession of cheetahs, unlike the succession of annual weather conditions, is itself subject to cumulative selection. Cheetahs will tend to become fleeter of foot, keener of eye, sharper of tooth. However ‘hostile’ the weather and other inanimate conditions may seem to be, they have no necessary tendency to get steadily more hostile. Living enemies, seen over the evolutionary timescale, have exactly that tendency.
The tendency for carnivores to get progressively ‘better’ would soon run out of steam, as do human arms races (for reasons of economic cost which we shall come to), were it not for the parallel tendency in the prey. And vice versa. Gazelles, no less than cheetahs, are subject to cumulative selection, and they too will tend, as the generations go by, to improve their ability to run fast, to react swiftly, to become invisible by blending into the long grass. They too are capable of evolving in the direction of becoming better enemies, in this case enemies of cheetahs. From the cheetahs’ point of view the mean annual temperature does not get systematically better or worse as the years go by, except in so far as any change for a well-adapted animal is a change for the worse. But the mean annual gazelle does tend to get systematically worse — more difficult to catch because better adapted to evade cheetahs. Again, the tendency towards progressive improvement in gazelles would slow to a halt, were it not for the parallel tendency to improvement shown by their predators. One side gets a little better because the other side has. And vice versa. The process goes into a vicious spiral, on a timescale of hundreds of thousands of years.
In the world of nations on their shorter timescale, when two enemies each progressively improve their weaponry in response to the other side’s improvements, we speak of an ‘arms race’. The evolutionary analogy is close enough to justify borrowing the term, and I make no apology to my pompous colleagues who would purge our language of such illuminating images. I have introduced the idea here in terms of a simple example, gazelles and cheetahs. This was to get across the important difference between a living enemy, which itself is subject to evolu
tionary change, and an inanimate non-malevolent condition such as the weather, which is subject to change, but not systematic, evolutionary change. But the time has come to admit that in my efforts to explain this one valid point I may have misled the reader in other ways. It is obvious, when you come to think about it, that my picture of an ever-advancing arms race was too simple in at least one respect. Take running speed. As it stands so far, the arms-race idea seems to suggest that cheetahs and gazelles should have gone on, generation after generation, getting ever faster until both travelled faster than sound. This has not happened and it never will. Before resuming the discussion of arms races, it is my duty to forestall misunderstandings.
The first qualification is this. I gave an impression of a steady upward climb in the prey-catching abilities of cheetahs, and the predator-avoiding abilities of gazelles. The reader might have come away with a Victorian idea of the inexorability of progress, each generation better, finer and braver than its parents. The reality in nature is nothing like that. The timescale over which significant improvement might be detected is, in any case, likely to be far longer than could be detected by comparing one typical generation with its predecessor. The ‘improvement’, moreover, is far from continuous. It is a fitful affair, stagnating or even sometimes going ‘backwards’, rather than moving solidly ‘forwards’ in the direction suggested by the arms-race idea. Changes in conditions, changes in the inanimate forces I have lumped under the general heading of ‘the weather’, are likely to swamp the slow and erratic trends of the arms race, as far as any observer on the ground could be aware. There may well be long stretches of time in which no ‘progress’ in the arms race, and perhaps no evolutionary change at all, takes place. Arms races sometimes culminate in extinction, and then a new arms race may begin back at square one. Nevertheless, when all this is said, the arms-race idea remains by far the most satisfactory explanation for the existence of the advanced and complex machinery that animals and plants possess. Progressive ‘improvement’ of the kind suggested by the arms-race image does go on, even if it goes on spasmodically and interruptedly; even if its net rate of progress is too slow to be detected within the lifetime of a man, or even within the timespan of recorded history.