Difference equation for tracking perturbations in systems of Boolean nested canalyzing functions

This paper studies the spread of perturbations through networks composed of Boolean functions with special canalyzing properties. Canalyzing functions have the property that at least for one value of one of the inputs the output is fixed, irrespective of the values of the other inputs. In this paper the focus is on partially nested canalyzing functions, in which multiple, but not all inputs have this property in a cascading fashion. They naturally describe many relationships in real networks. For example, in a gene regulatory network, the statement "if gene A is expressed, then gene B is not expressed regardless of the states of other genes" implies that A is canalyzing. On the other hand, the additional statement "if gene A is not expressed, and gene C is expressed, then gene B is automatically expressed; otherwise gene B's state is determined by some other type of rule" implies that gene B is expressed by a partially nested canalyzing function with more than two variables, but with two canalyzing variables. In this paper a difference equation model of the probability that a network node's value is affected by an initial perturbation over time is developed, analyzed, and validated numerically. It is shown that the effect of a perturbation decreases towards zero over time if the Boolean functions are canalyzing in sufficiently many variables. The maximum dynamical impact of a perturbation is shown to be comparable to the average impact for a wide range of values of the average sensitivity of the network. Percolation limits are also explored; these are parameter values which generate a transition of the expected perturbation effect to zero as other parameters are varied, so that the initial perturbation does not scale up with the parameters once the percolation limits are reached.