A preference for link operator functions can drive Boolean biological networks towards critical dynamics

Boolean modelling is a powerful framework to understand the operating principles of biological networks. The regulatory logic between biological entities in these networks is expressed as Boolean functions (BFs). There exist various types of BFs (and thus regulatory logic rules) which are meaningful in the biological context. In this contribution, we explore one such type, known as link operator functions (LOFs). We theoretically enumerate these functions and show that, among all BFs and even within the biologically consistent effective and unate functions (EUFs), the LOFs form a tiny subset. We then find that the AND-NOT LOFs are particularly abundant in reconstructed biological Boolean networks. By leveraging these facts, namely, the tiny representation of LOFs in the space of EUFs and their presence in the biological dataset, we show that the space of acceptable models can be shrunk by applying steady-state constraints to BFs, followed by the choice of LOFs which satisfy those constraints. Finally, we demonstrate that among a wide range of BFs, the LOFs drive biological network dynamics towards criticality.