speculative essays on ‘the origin of the Chordates’ and other topics of remote phylogeny, one of my tutors rightly tried to shake my faith in the value of such speculations by suggesting that anything could, in principle, evolve into anything else. Even insects could evolve into mammals, if only the right sequence of selection pressures were provided in the right order. At the time, as most zoologists would, I dismissed the idea as obvious nonsense, and I still, of course, don’t believe that the right sequence of selection pressures ever would be provided. Nor did my tutor. But as far as the principle is concerned, a simple thought experiment shows it to be nearly incontrovertible. We need only prove that there exists a continuous series of small steps leading from an insect, say a stag beetle, to a mammal, say a stag. By this I mean that, starting with the beetle, we could lay out a sequence of hypothetical animals, each one as similar to the previous member of the series as a pair of brothers might be, and the sequence would culminate in a red deer stag.
The proof is easy, provided only that we accept, as everyone does, that beetle and deer have a common ancestor, however far back. Even if there is no other sequence of steps from beetle to deer, we know that at least one sequence must be obtained by simply tracing the beetle’s ancestors back to the common ancestor, then working forwards down the other line to the deer.
We have proved that there exists a trajectory of stepwise change connecting beetle to deer and, by implication, a similar trajectory from any modern animal to any other modern animal. In principle, therefore, we maypresume that a series of selection pressures could be artificially contrived to propel a lineage along one of these trajectories. It was a quick thought experiment along these lines that enabled me to say, when discussing D’Arcy Thompson’s transformations, that ‘In a sense it is obviously necessarily true that any animal form can be turned into a related form by a mathematical transformation, although it is not obvious that the transformation will be a simple one.’ In this book I shall make frequent use of the thought-experiment technique. I warn the reader of this in advance, since scientists are sometimes annoyed by the lack of realism in such forms of reasoning. Thought experiments are not supposed to be realistic. They are supposed to clarify our thinking about reality.
One feature of life in this world which, like sex, we have taken for granted and maybe should not, is that living matter comes in discrete packages called organisms. In particular, biologists interested in functional explanation usually assume that the appropriate unit for discussion is the individual organism. To us, ‘conflict’ usually means conflict between organisms, each one striving to maximize its own individual ‘fitness’. We recognize smaller units such as cells and genes, and larger units such as populations, societies and ecosystems, but there is no doubt that the individual body, as a discrete unit of action, exerts a powerful hold over the minds of zoologists, especially those interested in the adaptive significance of animal behaviour. One of my aims in this book is to break that hold. I want to switch emphasis from the individual body as focal unit of functional discussion. At the very least I want to make us aware of how much we take for granted when we look at life as a collection of discrete individual organisms.
The thesis that I shall support is this. It is legitimate to speak of adaptations as being ‘for the benefit of’ something, but that something is best not seen as the individual organism. It is a smaller unit which I call the active, germ-line replicator. The most important kind of replicator is the ‘gene’ or small genetic fragment. Replicators are not, of course, selected directly, but by proxy; they are judged by their phenotypic effects. Although for some purposes it