Evolving complexity: how tinkering shapes cells, software and ecological networks.

A common trait of complex systems is that they can be represented by means of a network of interacting parts. It is, in fact, the network organization (more than the parts) that largely conditions most higher-level properties, which are not reducible to the properties of the individual parts. Can the topological organization of these webs provide some insight into their evolutionary origins? Both biological and artificial networks share some common architectural traits. They are often heterogeneous and sparse, and most exhibit different types of correlations, such as nestedness, modularity or hierarchical patterns. These properties have often been attributed to the selection of functionally meaningful traits. However, a proper formulation of generative network models suggests a rather different picture. Against the standard selection-optimization argument, some networks reveal the inevitable generation of complex patterns resulting from reuse and can be modelled using duplication-rewiring rules lacking functionality. These give rise to the observed heterogeneous, scale-free and modular architectures. Here, we examine the evidence for tinkering in cellular, technological and ecological webs and its impact in shaping their architecture. Our analysis suggests a serious consideration of the role played by selection as the origin of network topology. Instead, we suggest that the amplification processes associated with reuse might shape these graphs at the topological level. In biological systems, selection forces would take advantage of emergent patterns. This article is part of the theme issue 'Unifying the essential concepts of biological networks: biological insights and philosophical foundations'.