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NEMATODES 313<br />

the epicuticle is the true limiting ‘membrane’<br />

in nematodes. Several observations refute that<br />

concept. For example, the epicuticle is not<br />

dissociated by treatments that dissolve most<br />

15m<br />

FIGURE 13.4 Light micrograph (500) highlighting<br />

fine structure of the cortical and medial (aqueous)<br />

layers of the cuticle of Ascaris suum. Micrograph was<br />

stained with trichrome. Scale bar 15 m. Abbreviations:<br />

c, cortical layer; ml, medial layer; bl, composite<br />

basal layer; h, hypodermis.<br />

membranes, and lipophilic markers do not<br />

exhibit the lateral mobility characteristic of<br />

membranes when inserted into the surface of<br />

nematodes. The epicuticle in most species<br />

examined is devoid of intramembranous particles<br />

that are normally associated with cell<br />

membranes. Finally, under in vitro conditions,<br />

charged solutes diffuse across the cuticle at<br />

relatively high rates, indicating the absence of<br />

a functional lipid barrier at the surface.<br />

However, the epicuticle contains at least<br />

some lipid components. In Brugia malayi,<br />

for example, a cuticular lipid fraction was isolated<br />

that contains high levels of lysophosphatidylethanolamine<br />

and a novel species<br />

that rapidly degrades to two or more products<br />

upon exposure to atmospheric oxygen. This<br />

lipid extract contains saturated and unsaturated<br />

fatty acids in the range 14–22 carbons, with<br />

the majority being mono- and di-unsaturated<br />

18-carbon species. Furthermore, this parasite<br />

secretes a glutathione peroxidase across its<br />

ep<br />

c<br />

ml<br />

bl<br />

h<br />

m<br />

FIGURE 13.5 Schematic of adult Ascaris suum cuticle and hypodermis. Abbreviations: ep, epicuticle; c, cortical<br />

layer; ml, medial (aqueous) layer; bl, composite basal layer; h, hypodermis; m, muscle (drawing courtesy of<br />

J.W. Bowman).<br />

BIOCHEMISTRY AND CELL BIOLOGY: HELMINTHS

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