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Oligosaccharide chain distortion is induced by HPSE interaction. (a) Linear helical structure of a free heparin tetrasaccharide, adapted from an NMR derived model of heparin with all IdoUAs restrained in 2SO42. (b) Configuration of dp4 in the HPSE active site, showing a bend around the −2, −1, +1 sugars caused by HPSE interaction with −2 N-sulfate and +1 6O-sulfate. (c) Configuration of M09 S05a in complex with HPSE, showing the same distortion around −2→−1 as observed for dp4.
