Boolean nested canalizing functions: A comprehensive analysis

Yuan Li, John O. Adeyeye, David Murrugarra, Boris Aguilar, Reinhard Laubenbacher

Research output: Contribution to journalArticlepeer-review

36 Scopus citations


Boolean network models of molecular regulatory networks have been used successfully in computational systems biology. The Boolean functions that appear in published models tend to have special properties, in particular the property of being nested canalizing, a concept inspired by the concept of canalization in evolutionary biology. It has been shown that networks comprised of nested canalizing functions have dynamic properties that make them suitable for modeling molecular regulatory networks, namely a small number of (large) attractors, as well as relatively short limit cycles. This paper contains a detailed analysis of this class of functions, based on a novel normal form as polynomial functions over the Boolean field. The concept of layer is introduced that stratifies variables into different classes depending on their level of dominance. Using this layer concept a closed form formula is derived for the number of nested canalizing functions with a given number of variables. Additional metrics considered include Hamming weight, the activity number of any variable, and the average sensitivity of the function. It is also shown that the average sensitivity of any nested canalizing function is between 0 and 2. This provides a rationale for why nested canalizing functions are stable, since a random Boolean function in n variables has average sensitivity n2. The paper also contains experimental evidence that the layer number is an important factor in network stability.

Original languageEnglish
Pages (from-to)24-36
Number of pages13
JournalTheoretical Computer Science
StatePublished - Apr 15 2013

Bibliographical note

Funding Information:
The authors thank the referees for insightful comments that have improved the manuscript. The first and second authors were supported by Award # W911NF-11-10166 from the US DoD. The third and fifth authors were supported by NSF Grant CMMI-0908201.


  • Boolean function Nested canalizing function Layer number Extended monomial Multinomial coefficient Dynamical system Hamming weight Activity Average sensitivity

ASJC Scopus subject areas

  • Theoretical Computer Science
  • Computer Science (all)


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