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" "B$ZeMa,Ō 9ܰr$r%KP)sp:"Mi9ܰB$ : 1Minimal Defining Structure of a GlycosphingolipidGlycan-O-Ceramide  -Biosynthesis of Ceramide and Glucosylceramide$ Nomenclature Issues 1Isolation and purification of Glycosphingolipids 22Most glycosphingolipids obtained in good yield from cells and tissues by sequential organic extractions of increasing polarity Some separation by polarity achieved by two-phase extractions Subsequent fractionation away from other lipids in the extract using: DEAE ion exchange chromatography Silica gel thin layer and column separations HPLC adaptations of these methods are useful in obtaining complete separations. Principles of structural characterization of these molecules to be presented elsewhere$p d Op" d /$p d )%p d +$p d 0%p d OHBiosynthesis of different classes of glycans within the ER-Golgi PathwayII,$  GStepwise elongation of Glucosylceramidegenerates unique Core structures #Major Classes of Glycosphingolipids  EExamples of outer chains and modifications to Glycosphingolipid Cores-2!:Pathways for Ganglio-series Glycosphingolipid biosynthesis;;,   IMetabolic relationships in the biosynthesis and turnover of SphingolipidsJJ; /Turnover and Degradation of Glycosphingolipids ~Internalized from plasma membrane via endocytosis Pass through endosomes (some remodelling possible?) Terminal degradation in lysosomes - stepwise reactions by specific enzymes. Some final steps involve cleavages close to the cell membrane, and require facilitation by specific sphingolipid activator proteins (SAPs). Individual components, available for re-utilization in various pathways. At least some of glucosylceramide may remain intact and be recycled Human diseases in which specific enzymes or SAPs are genetically deficient ( storage disorders i$p d +%p d '$p d "%p d 2$p d 1%p d @&   # fR07Monoclonal antibodies (Mabs) against Glycosphingolipids$ Many  tumor-specific MAbs directed against glycans Majority react best with glycosphingolipids. Most MAbs are actually detecting  onco-fetal antigens Some used for diagnostic and prognostic applications in human diseases Few being exploited for attempts at monoclonal antibody therapy of tumors. Many used to demonstrate cell type-specific regulation of specific GSL structures in a temporal and spatial manner during development. Precise meaning of findings for cancer biology and development being explored..  $p d t   u&Biological Roles of Glycosphingolipids &Biological Roles of GlycosphingolipidszThought to be critical components of the epidermal (skin) permeability barrier Organizing role in cell membrane. Thought to associate with GPI anchors in the trans-Golgi, forming  rafts which target to apical domains of polarized epithelial cells May also be in glycosphingolipid enriched domains ( GEMs ) which are associated with cytosolic oncogenes and signalling molecules Physical protection against hostile environnments Binding sites for the adhesion of symbiont bacteria. Highly specific receptor targets for a variety of bacteria, toxins and viruses. 4$p d '%p d >O -! b%&Biological Roles of GlycosphingolipidsSpecific association of certain glycosphingolipids with certain membrane receptors. Can mediate low-affinity but high specificity carbohydrate-carbohydrate interactions between different cell types. Targets for autoimmune antibodies in Guillian-Barre and Miller-Fisher syndromes following Campylobacter infections and in some patients with human myeloma Shed in large amounts by certain cancers - these are found to have a strong immunosuppressive effects, via as yet unknown mechanisms$p d P ^ VExamples of interactions between glycosphingolipids and bacterial toxins or receptorsWW #4#/B19 Parvovirus and Uropathogenic E.coli Binding00 TExamples of proposed interactions between glycosphingolipids and mammalian receptors)GNatural and induced Genetic Disorders in Glycosphingolipid biosynthesisHH( One cultured cell line completely deficient in GlcCer synthase - thus, GSLs not essential for growth of single cells in a culture dish Many human genetic defects in GSL degradation result in  storage disorders with accumulation of specific intermediates. In contrast, human genetic defects in the biosynthesis of GSLs seem very rare. Targetted gene disruption of Ceramide Galactosyltransferase in mice: loss of GalCer and SulfatedGalCer (Sulfatide) in myelin. Mice form myelin with GlcCer, which replaces GalCer. Generalized tremors and mild ataxia, electrophysiological evidence for conduction deficits. With increasing age, progressive hindlimb paralysis and severe vacuolation of the ventral region of the spinal cord. Terminal differentiation and morphological maturation of oligodendrocytes is enhanced. 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