Mix and Match

It's easier to reconfigure a complex system to do new things if it is built from simpler, independent modules. But in biology, modules may be useful for more immediate reasons.

Biological systems, ranging from communities to molecular networks, often feature a modular organization, which for one thing makes it easier for a species to evolve in response to changes in the environment. In some cases, this flexibility might have been selected during prior changes. But modularity can also make life easier for a single organism during its lifetime, and be selected for this reason.

In their book The Plausibility of Life, Marc Kirschner and John Gerhart include modularity as one aspect of "facilitated variation." In particular, they say, genetic changes that affect the "weak linkages" between modules can cause major changes in the resulting phenotype. As long as the modules, the "conserved core processes," are not disrupted, the resulting organism is likely to be viable, and possibly an improvement on its predecessors.

In describing facilitated variation, Kirschner and Gerhart defer the question of whether facilitating rapid future evolution alone causes these features to be selected. Perhaps it does, in some circumstances. But in any case, we can regard the presence of features that enable rapid reconfiguration as an observational fact.

Moreover, the practical challenges of development demand the same sort of robust flexibility that encourages rapid evolutionary change. Over the development of a complex organism, various cells are exposed to drastically different local environments. In addition, genetic or other changes pose unpredictable challenges to the molecular and other systems of the cells. Throughout these changes, critical processes, like metabolism and DNA replication, need to keep working reliably.

To survive and reproduce in the face of these variations organisms need a robust and flexible organization. Features that allow such flexibility should be selected, if only because they improve individual fitness. These same features may then increase evolutionary adaptability, whether or not that adaptability is, by itself, evolutionarily favored.

Whether adaptability is selected for its evolutionary potential or only for helping organisms thrive in a chaotic environment, it has a profound effect on subsequent evolution. A flexible organization including modularity and other features allows small genetic alterations to be leveraged into large but nonfatal changes in the developing creature, so the population can rapidly explore possible innovations.