Mucins produced by goblet cells of the respiratory mucosa are condensed while stored in secretory granules. Mucin condensation and its decondensation upon exocytosis can be explained by the theory of polymer gel phase transition. After the opening of a secretory pore, Ca2+ inside the granule is exchanged for extracellular Na+. Na/Ca exchange triggers a polymer gel phase transition whereby the mucin polymer matrix undergoes massive swelling and thereby changes from a condensed to a hydrated phase. Swelling of the granular content is driven by a Donnan potential and results in the release of secretory product and the formation of small mucin gels, which later anneal to each other to form the respiratory mucus. Because of the tangled rather than cross-linked topology of the mucin network, the rheologic properties of the respiratory mucus depend primarily on hydration. As mucins are polyionic, the hydration of mucus is controlled by a Donnan equilibrium. Hence, mucus hydration and rheology are determined by two factors: the quantity, chain length, and charge density of the secreted mucins, and the amount and the ionic and polyionic composition of the water transported across the respiratory mucosa.
