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O-sugars bind to the flavan nucleus through a glycosidic bond, which is formed between the anomeric carbon of the sugar and a hydroxyl group on the flavan nucleus. The most common type of O-sugar linkage in flavonoids is the C-glycosidic bond, which involves the anomeric carbon of the sugar being linked to the carbon atom of the flavan nucleus. Other types of O-sugar linkages include the O-glycosidic bond, which involves the anomeric carbon of the sugar being linked to the oxygen atom of the flavan nucleus, and the S-glycosidic bond, which involves the anomeric carbon of the sugar being linked to the sulfur atom of the flavan nucleus.