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Biochemical covalent modification networks exhibit a remarkable suite of steady state and dynamical properties such as multistationarity, oscillations, ultrasensitivity and absolute concentration robustness. This paper focuses on conditions required for a network of this type to have a species with absolute concentration robustness. We find that the robustness in a substrate is endowed by its interaction with a bifunctional enzyme, which is an enzyme that has different roles when isolated versus when bound as a substrate-enzyme complex. When isolated, the bifunctional enzyme promotes production of more molecules of the robust species while when bound, the same enzyme facilitates degradation of the robust species. These dual actions produce robustness in the large class of covalent modification networks. For each network of this type, we find the network conditions for the presence of robustness, the species that has robustness, and its robustness value. The unified approach of simultaneously analyzing a large class of networks for a single property, i.e. absolute concentration robustness, reveals the underlying mechanism of the action of bifunctional enzyme while simultaneously providing a precise mathematical description of bifunctionality. 

Abstract Reaction networks are commonly used within the mathematical biology and mathematical chemistry communities to model the dynamics of interacting species. These models differ from the typical graphs found in random graph theory since their vertices are constructed from elementary building blocks, i.e. the species. We consider these networks in an Erdös–Rényi framework and, under suitable assumptions, derive a threshold function for the network to have a deficiency of zero, which is a property of great interest in the reaction network community. Specifically, if the number of species is denoted by n and the edge probability by $$p_n$$ , then we prove that the probability of a random binary network being deficiency zero converges to 1 if $$p_n\ll r(n)$$ as $$n \to \infty$$ , and converges to 0 if $$p_n \gg r(n)$$ as $$n \to \infty$$ , where $$r(n)=\frac{1}{n^3}$$ .