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Currency and commodity metabolites: Their identification and relation to the modularity of metabolic networks

The large-scale shape and function of metabolic networks are intriguing topics of systems biology. Such networks are on one hand commonly regarded as modular (i.e. built by a number of relatively independent subsystems), but on the other hand they are robust in a way not expected of a purely modular system. To address this question we carefully discuss the partition of metabolic networks into subnetworks. The practice of preprocessing such networks by removing the most abundant substrates, "currency metabolites," is formalized into a network-based algorithm. We study partitions for metabolic networks of many organisms and find cores of currency metabolites and modular peripheries of what we call "commodity metabolites." The networks are found to be more modular than random networks but far from perfectly divisible into modules. We argue that cross-modular edges are the key for the robustness of metabolism.