The argument is moving towards the idea of a stereotyped, regularly repeating life cycle. Not only does each generation begin with a single-celled bottleneck. It also has a growth phase-'childhood'-of rather fixed duration. The fixed duration, the stereotypy, of the growth phase, makes it possible for particular things to happen at particular times during embryonic development, as if governed by a strictly observed calendar. To varying extents in different kinds of creature, cell divisions during development occur in rigid sequence, a sequence that recurs in each repetition of the life cycle. Each cell has its own location and time of appearance in the roster of cell divisions. In some cases, incidentally, this is so precise that embryologists can give a name to each cell, and a given cell in one individual organism can be said to have an exact counterpart in another organism.
So, the stereotyped growth cycle provides a clock, or calendar, by means of which embryological events may be triggered. Think of how readily we ourselves use the cycles of the earth's daily rotation, and its yearly circumnavigation of the sun, to structure and order our lives. In the same way, the endlessly repeated growth rhythms imposed by a bottlenecked life cycle will-it seems almost inevitable-be used to order and structure embryology. Particular genes can be switched on and off at particular times because the bottleneck/growth-cycle calendar ensures that there is such a thing as a particular time. Such well-tempered regulations of gene activity are a prerequisite for the evolution of embryologies capable of crafting complex tissues and organs. The precision and complexity of an eagle's eye or a swallow's wing couldn't emerge without clockwork rules for what is laid down when.
The third consequence of a bottlenecked life history is a genetic one. Here, the example of bottle-wrack and splurge-weed serves us again. Assuming, again for simplicity, that both species reproduce asexually, think about how they might evolve. Evolution requires genetic change, mutation. Mutation can happen during any cell division. In splurge-weed, cell lineages are broad-fronted, the opposite of bottlenecked. Each branch that breaks apart and drifts away is many-celled. It is therefore quite possible that two cells in a daughter will be more distant relatives of one another than either is to cells in the parent plant. (By 'relatives', I literally mean cousins, grandchildren and so on. Cells have definite lines of descent and these lines are branching, so words like second cousin can be used of cells in a body without apology.) Bottle-wrack differs sharply from splurge-weed here. All cells in a daughter plant are descended from a single spore cell, so all cells in a given plant are closer cousins (or whatever) of one another than of any cell in another plant.
This difference between the two species has important genetic consequences. Think of the fate of a newly mutated gene, first in splurge-weed, then in bottle-wrack. In splurge-weed, the new mutation can arise in any cell, in any branch of the plant. Since daughter plants are produced by broad-fronted budding, lineal descendants of the mutant cell can find themselves sharing daughter plants and grand-daughter plants with unmutated cells which are relatively distant cousins of themselves. In bottle-wrack, on the other hand, the most recent common ancestor of all the cells in a plant is no older than the spore that provided the plant's bottlenecked beginning. If that spore contained the mutant gene, all the cells of the new plant will contain the mutant gene. If the spore did not, they will not. Cells in bottle-wrack will be more genetically uniform within plants than cells in splurge-weed (give or take an occasional reverse-mutation). In bottle-wrack, the individual plant will be a unit with a genetic identity, will deserve the name individual. Plants of splurge-weed will have less genetic identity, will be less entitled to the name 'individual' than their opposite numbers in bottle-wrack.
This is not just a matter of terminology. With mutations around, the cells within a plant of splurge-weed will not have all the same genetic interests at heart. A gene in a splurge-weed cell stands to gain by promoting the reproduction of its cell. It does not necessarily stand to gain by promoting the reproduction of its 'individual' plant. Mutation will make it unlikely that the cells within a plant are genetically identical, so they won't collaborate wholeheartedly with one another in the manufacture of organs and new plants. Natural selection will choose among cells rather than 'plants'. In bottle-wrack, on the other hand, all the cells within a plant are likely to have the same genes, because only very recent mutations could divide them. Therefore they will happily collaborate in manufacturing efficient survival machines. Cells in different plants are more likely to have different genes. After all, cells that have passed through different bottlenecks may be distinguished by all but the most recent mutations-and this means the majority. Selection will therefore judge rival plants, not rival cells as in splurge-weed. So we can expect to see the evolution of organs and contrivances that serve the whole plant.
By the way, strictly for those with a professional interest, there is an analogy here with the argument over group selection. We can think of an individual organism as a 'group' of cells. A form of group selection can be made to work, provided some means can be found for increasing the ratio of between-group variation to within-group variation. Bottle-wrack's reproductive habit has exactly the effect of increasing this ratio; splurge-weed's habit has just the opposite effect. There are also similarities, which may be revealing but which I shall not explore, between 'bottlenecking' and two other ideas that have dominated this chapter. Firstly the idea that parasites will cooperate with hosts to the extent that their genes pass to the next generation in the same reproductive cells as the genes of the hosts- squeezing through the same bottleneck. And secondly the idea that the cells of a sexually reproducing body cooperate with each other only because meiosis is scrupulously fair.
To sum up, we have seen three reasons why a bottlenecked life history tends to foster the evolution of the organism as a discrete and unitary vehicle. The three may be labelled, respectively, 'back to the drawing board', 'orderly timing-cycle', and 'cellular uniformity'. Which came first, the bottlenecking of the life cycle, or the discrete organism? I should like to think that they evolved together. Indeed I suspect that the essential, defining feature of an individual organism is that it is a unit that begins and ends with a single-celled bottleneck. If life cycles become bottlenecked, living material seems bound to become boxed into discrete, unitary organisms. And the more that living material is boxed into discrete survival machines, the more will the cells of those survival machines concentrate their efforts on that special class of cells that are destined to ferry their shared genes through the bottleneck into the next generation. The two phenomena, bottlenecked life cycles and discrete organisms, go hand in hand. As each evolves, it reinforces the other. The two are mutually enhancing, like the spiralling feelings of a woman and a man during the progress of a love affair.
The Extended Phenotype is a long book and its argument cannot easily be crammed into one chapter. I have been obliged to adopt here a condensed, rather intuitive, even impressionistic style. I hope, nevertheless, that I have succeeded in conveying the flavour of the argument.
Let me end with a brief manifesto, a summary of the entire selfish gene/extended phenotype view of life. It is a view, I maintain, that applies to living things everywhere in the universe. The fundamental unit, the prime mover of all life, is the replicator. A replicator is anything in the universe of which copies are made. Replicators come into existence, in the first place, by chance, by the random jostling of smaller particles. Once a replicator has come into existence it is capable of generating an indefinitely large set of copies of itself. No copying process is perfect, however, and the population of replicators comes to include varieties that differ from one another. Some of these varieties turn out to have lost the power of self-replication, and their kind ceases to exist when they themselves cease to exist. Others can still replicate, but less effectively. Yet other varieties happen to find themselves in possession of new tricks: they turn out to be even better self-replicators than their predecessors and contemporaries. It is their
descendants that come to dominate the population. As time goes by, the world becomes filled with the most powerful and ingenious replicators.
Gradually, more and more elaborate ways of being a good replicator are discovered. Replicators survive, not only by virtue of their own intrinsic properties, but by virtue of their consequences on the world. These consequences can be quite indirect. All that is necessary is that eventually the consequences, however tortuous and indirect, feed back and affect the success of the replicator at getting itself copied.
The success that a replicator has in the world will depend on what kind of a world it is-the pre-existing conditions. Among the most important of these conditions will be other replicators and their consequences. Like the English and German rowers, replicators that are mutually beneficial will come to predominate in each other's presence. At some point in the evolution of life on our earth, this ganging up of mutually compatible replicators began to be formalized in the creation of discrete vehicles-cells and, later, many-celled bodies. Vehicles that evolved a bottlenecked life cycle prospered, and became more discrete and vehicle-like.
This packaging of living material into discrete vehicles became such a salient and dominant feature that, when biologists arrived on the scene and started asking questions about life, their questions were mostly about vehicles-individual organisms. The individual organism came first in the biologist's consciousness, while the replicators-now known as genes-were seen as part of the machinery used by individual organisms. It requires a deliberate mental effort to turn biology the right way up again, and remind ourselves that the replicators come first, in importance as well as in history.
One way to remind ourselves is to reflect that, even today, not all the phenotypic effects of a gene are bound up in the individual body in which it sits. Certainly in principle, and also in fact, the gene reaches out through the individual body wall and manipulates objects in the world outside, some of them inanimate, some of them other living beings, some of them a long way away. With only a little imagination we can see the gene as sitting at the centre of a radiating web of extended phenotypic power. And an object in the world is the centre of a converging web of influences from many genes sitting in many organisms. The long reach of the gene knows no obvious boundaries. The whole world is criss-crossed with causal arrows joining genes to phenotypic effects, far and near.
It is an additional fact, too important in practice to be called incidental but not necessary enough in theory to be called inevitable, that these causal arrows have become bundled up. Replicators are no longer peppered freely through the sea; they are packaged in huge colonies-individual bodies. And phenotypic consequences, instead of being evenly distributed throughout the world, have in many cases congealed into those same bodies. But the individual body, so familiar to us on our planet, did not have to exist. The only kind of entity that has to exist in order for life to arise, anywhere in the universe, is the immortal replicator.
The End.
The Selfish Gene Page 35