Homeostasis is a critical property of living beings that involves the ability to self-regulate in response to changes in the environment in order to maintain a certain dynamic balance aecting form and/or function. Homeostasis is of particular importance in multicellular organisms, where it is intertwined with development [2,3]. Organisms have evolved intricate control mechanisms that ensure developmental processes achieve their end points and stabilize (e.g., dierentiate) as well as allow for a degree of adaptability to a range of conditions (e.g., stress or damage induced by wounding). is allows for the emergence of a more robust system that can tolerate both external and internal perturbations [4]. However, there are limitations to this tolerance, and oen it is the rare events that cause the most disruption [5]; think of the extinction of dinosaurs for an example. From an evolutionary point of view, this is a viable trade-o between the energetic cost of homeostasis versus the tness benet it would provide. In practical terms, homeostasis of living multicellular organisms is constrained in terms of the amount of disruption they can cope with and in terms of the amount of time they will remain homeostatic.

Many different single-cell-based models have been developed and applied to biological and medical problems. Computational approaches used are Monte-Carlo simulations, energy minimisation techniques, v