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An investigation of the mechanism of infection by digenetic trematodes: the penetration of the miracidium of Fasciola hepatica into its snail host Lymnaea truncatula

The penetration barrier presented to the miracidium by the snail epithelium can be divided into three layers. The chemical composition and physical configuration of the outermost of these plays an important part in the initial attachment response of the miracidium. Attachment can be stimulated in the absence of the snail by pure chemicals in solution. However, the surface to which the miracidium attaches must have the correct physical configuration otherwise the miracidium is unable to form a stable attachment. In vivo, the miracidial body begins to contract and relax following attachment to the snail. This coincides with the start of secretion by the apical gland and accessory gland cells. The snail's columnar epithelium is rapidly cytolysed so that 10 min after attachment the anterior of the miracidium has reached the underlying connective tissues. As the miracidium penetrates the snail, its ciliated epithelial cells are shed in sequence from anterior to posterior. This shedding removes a protective barrier against osmosis which is probably the acid mucopolysaccharide present in the epithelial cells. The mechanism of shedding is not understood but involves the reversal of binding by the desmosomal mucosubstance which attaches the epithelial cells to surrounding intercellular ridges. The miracidium metamorphoses into the sporocyst as it penetrates the snail, by forming a new body surface. The material for this is extruded from the vesiculated cells which lie beneath the musculature of the body wall. The process of surface formation coincides with cell shedding and moves backwards as cells are shed. At not more than 2·5 h after attachment the extruded cytoplasm forms a thin continuous layer over the surface of the organism. Contacts with underlying cells appear to have been broken and the cytoplasm is underlain by a thin fibrous basal lamella. In the first 24 h after penetration the surface of this syncytium becomes thrown into folds and metamorphosis into the sporocyst can be considered complete.