  Robustness, the insensitivity of some of a biological system's
functionalities to a set of distinct conditions, is intimately linked to
fitness. Recent studies suggest that it may also play a vital role in enabling
the evolution of species. Increasing robustness, so is proposed, can lead to
the emergence of evolvability if evolution proceeds over a neutral network that
extends far throughout the fitness landscape. Here, we show that the design
principles used to achieve robustness dramatically influence whether robustness
leads to evolvability. In simulation experiments, we find that purely redundant
systems have remarkably low evolvability while degenerate, i.e. partially
redundant, systems tend to be orders of magnitude more evolvable. Surprisingly,
the magnitude of observed variation in evolvability can neither be explained by
differences in the size nor the topology of the neutral networks. This suggests
that degeneracy, a ubiquitous characteristic in biological systems, may be an
important enabler of natural evolution. More generally, our study provides
valuable new clues about the origin of innovations in complex adaptive systems.
