Modules

When people design a complex system, they use a modular approach. But why should biology?

For us, modularity is a way to limit complexity. Breaking a big problem into a series or hierarchy of smaller ones makes it more manageable and comprehensible, which is especially important if it is assembled by many people--or one person over an extended time.

The key to a successful module is that its "guts"--the way its parts work together--doesn't depend on how it connects to other stuff. The module can be thought of as a "black box," that just does its job. You don't have to think about it again. For this to work, the connections between modules must be weak, limited to well-defined inputs and outputs that don't directly affect its internal workings.

When it's done right, a module can be easily re-used in new situations. For example, the part of a computer operating system that offers a help menu is tapped by lots of programs without worrying about how it works.

But biology is not designed. Biological systems emerge from an evolutionary process that rewards only survival and reproduction, with no regard for elegance or comprehensibility. Re-usability sounds like a good thing in the long run, but doesn't help an individual survive in the here and now.

Nonetheless, modularity seems to be widespread feature of biological systems. Your gall bladder, for example, is a well-defined blob that receives and releases fluids like blood and bile, but otherwise keeps its own counsel. It can even be removed if necessary. And it does much the same thing in other people and animals.

At a smaller scale, many of the basic components of cells are the same for all eukaryotes. They have the discrete nuclei that define them, and they also have other organelles that perform essential functions, like mitochondria that generate energy. These modules work the same way, whether they happen to appear in a brain cell or a skin cell.

Even at the molecular level, re-usable modules abound. For example, although ribosomes don't have a membrane delimiting them, they consist of very similar bundles of proteins and RNA for all eukaryotic cells, and only modestly different bundles for bacteria. In addition to such complexes, many "pathways," or chains of molecular interactions, recur in many different species.

We have to be careful, of course: simply because we represent complex biological systems as modules doesn't mean they are there. The modules we think we see could simply reflect our limited capacity to understand messy reality. But when researchers have looked at this question carefully, they found that modules really exist in biology, much more than they would in a random system of similar complexity.

But why should evolution favor modular arrangements? And how does a modular structure change the way organisms evolve?