Nutritional adaptations to parasitism within the platyhelminthes.

Some of the most significant alterations to the basic turbellarian plan are evident in the adaptations that relate to the acquisition of food by parasitic flatworms, reflecting the most potent of selection pressures in initiating and maintaining the host-parasite association. Nutritionally, ectoparasitic monogeneans show most correspondence with the predatory turbellarians, with certain monopisthocotylean members feeding by means of a protrusible pharynx and extracorporeal digestion, as skin-browsers of fish, with extensive intracellular digestion involving lysosomal enzymes in a well-differentiated gut. The more sheltered vascularised gill chamber of fish provides many polyopisthocotylean monogeneans with a totally renewable and more comprehensive diet in the form of blood, but haematophagy has necessitated a number of digestive adaptations, not least in resolving the problem of intracellular accumulations of haematin pigment. Haematophagy is the predominant feeding strategy of digeneans, but in contrast to monogeneans digestion of blood is largely extracellular; in schistosomes digestion is rapid, involving a battery of cathepsin-like cysteine proteinases and aminopeptidases. The external surfaces of all parasitic flatworms depart from turbellarian character and are composed of a multifunctional syncytial tegument, which is permeable to a variety of small organic solutes, some crossing by passive diffusion, others via facilitated or active mediated transport. The relative roles of the tegument and gut in trematode nutrition are difficult to assess, but can be related to the nature of the microhabitat within the host. Cestodes are highly adapted intestinal parasites bereft of any vestige of gut, and their tegument has become elaborated into a sophisticated and highly efficient digestive-absorptive layer, rivalling the vertebrate mucosa in its ability to gain kinetic advantage in the selective uptake of nutrient at the host-parasite interface. The patterns of energy metabolism in adult flatworm parasites are generally anaerobic and based on glycogen, with abbreviated metabolic pathways and the loss of biosynthetic capacities.