Life’s building blocks: the modular path to multiscale complexity

Modularity, the structuring of systems into discrete, interconnected units, is a fundamental organizing principle in biology across multiple scales. Recent progress in understanding the role of modularity as an evolutionary mechanism and a key driver of biological complexity has highlighted its importance in shaping the structure and function of living systems. Here, we propose a unifying framework that identifies the potential evolutionary advantages of modularity in systems ranging from molecular networks to ecologies, such as facilitating evolvability, enhancing robustness, improving information flows, and enabling the emergence of higher-level functions. Our analysis reveals the pervasiveness of modularity in living systems and highlights its crucial role in the evolution of multiscale hierarchies of increasing complexity.