Microanatomy of secretory granule release from guinea pig tracheal goblet cells.

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Abstract
The microanatomy of mucin granule release from epithelial goblet cells has been investigated in guinea pig tracheae. Using a tannic acid arrest procedure, granule release under basal conditions and after high K+ or acetylcholine (ACh) application was arrested and a variety of granule fusion sites were identified in ultrathin sections and freeze-fracture replicas. Rather than there being subclasses of secretory cells containing either electron-lucent granules (indicative of mucin) or smaller electron-dense (serous) granules, the majority of secretory cells in both control and treated groups contained granules with an electrondense core surrounded by an electron-lucent region. Granule release sites were of three principal types: (1) simple exocytosis, where the membranes of single granules fused directly with the plasma membrane to give an "omega" profile; (2) compound exocytosis, where granule membranes, fused together intracellularly, were found in continuity with the plasma membrane; and (3) apocrine-like secretion, which involved the loss of the central apical mass of granules together with elements of the cell cytoplasm. In treated preparations, there was an increase in the number of cells exhibiting fusion sites; the percentage showing simple fusions fell from 82% to 59% (with ACh) and 57% (with KCl), whereas the percentage of cells exhibiting compound and apocrine-like secretion increased. Dense cores were frequently retained at the sites of fusion and, despite the expansive decondensation of mucin known to occur, there was also evidence of some retention of the electron-lucent material.

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