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The observed spatio-temporal ciliary beat patterns on multiciliated epitheliaare suspected to be the result of self-organizing processes on various levels.Here, we present an abstract epithelium model at the pluricellular level, whichintends to make the self-organization of ciliary beating patterns as well as ofthe associated fluid transport across the airway epithelium plausible. Ciliatedcells are modeled in terms of locally interacting oscillating two-stateactuators. The local interactions among these boolean actuators are triggeredby seeded mucus lumps. In the course of a simulation the actuators' state andthe associated mucus velocity field self-organize in tandem. We suggest toconsider the dynamics on multiciliated epithelia in the context of adaptive(boolean) networks. Within the framework of adaptive boolean networks ciliatedcells represent the nodes and as the mucus establishes the local interactionsamong nodes, its distribution determines the topology of the network.Furthermore, we present the results and insights from comprehensive parameterstudies. The results show evidence that so called deterministic update schemes,which are meant to represent intercellular signaling, lead to more realisticand robust dynamics and may therefore be favored by nature. Finally, we supposethat unciliated cells introduce a modular network topology on ciliatedepithelia causing the self-organization taking place simultaneously in each"ciliated module". This reasoning provides the first consistent explanation forthe meaning of the observed patchy expression patterns of the mucus modulationwave fields. Modularity may therefore be seen as a modular construction plan ofnature for ciliated epithelia whose number of cells range over several order ofmagnitudes.

Self-Organization of Self-Clearing Beating Patterns in an Array of Locally Interacting Ciliated Cells Formulated as an Adaptive Boolean Network