Darwin’s theory of evolution is possibly the most important and influential theory in biology. I am not going to argue against that claim, as I do believe that it is a fine piece of theoretical work, and a great conceptual advance in biology. However, I also find that the explanatory power of Darwin’s theory is often overstated. I recently visited a public exhibition in a museum about Darwin. Nice exhibition overall, but I was a bit bothered by the claim that Darwin’s theory explains the origin, diversity and adaptedness of species, case solved. I have the same feeling when I read in many articles or when I hear in conversations with many scientists that such and such observed feature of living organisms is “explained by evolution”. The reasoning generally goes like this: such biological structure is apparently beneficial to the organism, and therefore the existence of that structure is explained by evolution. As if the emergence of that structure directly followed from Darwin’s account of evolution.

To me, the Darwinian argument is often used as magic, and is mostly void of any content. Replace “evolution” by “God” and you will notice no difference in the logical structure or arguments. Indeed, what the argument actually contains is 1) the empirical observation that the biological organism is apparently adapted to its environment, thanks to the biological feature under scrutiny; 2) the theoretical claim that organisms are adapted to their environment. Note that there is nothing in the argument that actually involves evolution, i.e., the change of biological organisms through some particular process. Darwin is only invoked to back up the theoretical claim that organisms are adapted, but there is nothing specifically about Darwinian evolution that is involved in the argument. It could well be replaced by God, Lamarck or aliens.

What makes me uneasy is that many people seem to think that Darwin’s theory fully explains how biological organisms get to be adapted to their environment. But even in its modern DNA form, it doesn’t. It describes some of the important mechanisms of adaptation, but there is an obvious gap. I am not saying that Darwin’s theory is wrong, but simply that it only addresses part of the problem.

What is Darwin’s theory of evolution? It is based on three simple steps: variation, heredity and selection. 1) Individuals of a given species vary in different respects. 2) Those differences are inherited. In the modern version, new variations occur randomly at this step, and so variations are introduced gradually over generations. 3) Individuals with adapted features survive and reproduce more than others (by definition of “adapted feature”), and therefore spread those features in the population. There is ample empirical evidence for these three claims, and that was the great achievement of Darwin.

The gap in the theory is the nature and distribution of variations. In the space of all possible small variations in structure that one might imagine, do we actually see them in a biological population? Well for one, there are a substantial number of individuals that actually survive for a certain time, so a large number of those variations are not destructive. Since the metaphor of the day is to see the genome as a code for a program, let us consider computer programs. Take a functional program and randomly change 1% of all the bits. What is the probability that 1) the program doesn’t crash, 2) it produces something remotely useful? I would guess that the probability is vanishingly small. You will note that this is not a very popular technique in software engineering. Another way to put it: consider the species of programs that calculate combinatorial functions (say, factorials, binomial coefficients and the like). Surely one might argue that individuals vary by small changes, but conversely, would a small random change in the code typically produce a new combinatorial function?

So it doesn’t follow logically from the three steps of Darwin’s theory that biological organisms should be adapted and survive to changing environments. There is a critical ingredient that is missing: to explain how, in sharp contrast with programs, a substantial fraction of new variations are constructive rather than destructive. In more modern terms, how is it that completely random genetic mutations result in variations in phenotypes that are not arbitrary?

Again I am not saying that Darwin is wrong, but simply that his theory only addresses part of the problem, and that it is not correct to claim that Darwin’s theory fully explains how biological organisms are adapted to their environment (ie, perpetuate themselves). A key point, and a very important research question, is to understand how new variations can be constructive. This can be addressed within the Darwinian framework, as I outlined in a previous post. It leads to a view that departs quite substantially from the program metaphor. A simple remark: the physical elements that are subject to random variation cannot be mapped to the physical elements of structure (e.g. molecules) that define the phenotype, for otherwise those random variations would lead to random (ie mostly destructive) phenotypes. Rather, the structure of the organism must be the result of a self-regulatory process that can be steered by the elements subject to random variation. This is consistent with the modern view of the genome as a self-regulated network of genes, and with Darwin’s theory. But it departs quite substantially from the magic view of evolution theory that is widespread in the biological literature (at least in neuroscience), and instead points to self-regulation and optimization processes operating at the scale of the individual (not of generations).