Abstract
Biological networks are often modular. Explanations for this peculiarity either assume an adaptive advantage of a modular design such as higher robustness, or attribute it to neutral factors such as constraints underlying network assembly. Interestingly, most insights on the origin of modularity stem from models in which interactions are either determined by highly simplistic mechanisms, or have no mechanistic basis at all. Yet, empirical knowledge suggests that biological interactions are often mediated by complex structural or behavioural traits. Here, we investigate the origins of modularity using a model in which interactions are determined by potentially complex traits. Specifically, we model system elements-such as the species in an ecosystem-as finite-state machines (FSMs), and determine their interactions by means of communication between the corresponding FSMs. Using this model, we show that modularity probably emerges for free. We further find that the more modular an interaction network is, the less complex are the traits that mediate the interactions. Altogether, our results suggest that the conditions for modularity to evolve may be much broader than previously thought.