Mucus is an ubiquitous polymer hydrogel that functions as a protective coat on the surface of integument and mucosa of species ranging from simple animals (such as coelenterates) to mammals. The polymer matrix of mucus is made out of long-chain glycoproteins called mucins that are tangled together, forming a randomly woven, highly polyionic network (Lee et al., 1977; Verdugo et al., 1983). Mucin-containing granules, produced by mammalian goblet cells in vitro, undergo massive post-exocytotic swelling. Their swelling kinetics is similar to the swelling of condensed artificial polymer gels (Verdugo, 1984; Tanaka and Fillmore, 1979). We had proposed that mucins must be condensed in the secretory granule and expand by hydration during or after exocytosis (Verdugo, 1984; Tam and Verdugo, 1981). However, the polyionic charges of mucins prevents condensation unless they (the mucins) are appropriately shielded. The present experiments were designed to assert the presence of an intragranular shielding cation and its role in secretion. Giant mucin granules of the slug (Ariolimax columbianus) are released intact from mucus-secreting cells of the slug's skin. They burst spontaneously outside the cell, forming, upon hydration, the typical slug mucus (Deyrup-Olsen et al., 1983). We report here that these granules contain from 2.5 to 3.6 moles calcium/kg dry material, and that calcium is released from the granules immediately before the burst that discharges their secretory product. Therefore, we propose that calcium functions as a shielding cation of polyionic mucins, and that the bursting discharge of mucins from secretory granules must result from the release of calcium from the intragranular compartment.(ABSTRACT TRUNCATED AT 250 WORDS)

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Mucous secretion by the body wall of the terrestrial slugs (Ariolimax columbianus, Arionidae; and other species) was found to involve at least three distinct stages--release of vesicles, formation of granules, and organization of strands. Mucus is stored intracellularly in membrane-bound vesicles, and these are shed intact from the mucous cells. Disruption of the vesicle membrane, with release of contents, can be effected by endogenous lytic agent(s), as well as by exogenous surfactants, lipid solvents, or hypotonic media. Thereupon 1-micron granules are released. These may be stable, or they may change to material that is finely granular or in the form of strands; the transition to strands is facilitated by shear stress exerted through the fluid containing the mucous components. Lectins organize, or are organized with, the strands, as evidenced by agglutination of erythrocytes on them. Mucous formation, as seen in the living slug, differs markedly from the one-step process of exocytosis of fluid mucus inferred from studies of mucous membranes fixed for ultrastructural investigation.