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(a) Example HS tetrasaccharide with complete N-sulfation and illustrating uronate C5 epimers alongside points for O-sulfation; free acid form shown; (b) replacement of glycosidic oxygen with sulfur to create an S-glycoside analogue; and (c) interglycosidic torsion angles represented by Newman projections: Phi (Φ) is the O5′–C1′–O4–C4 angle and Psi (ψ) is the C1′–O4–C4–C3 angle, and O4 is the linkage oxygen.
