Materials, and descriptions of primary antibodies and chemicals, immunohistochemical DS-epi1 staining, characterization of metabolically labeled CS/DS chains from the cell layer and cell culture medium, immunoblot analysis and phospho-kinase array, qRT-PCR, digital holographic imaging, O-sulfotransferase assay, micro array RNA analysis and characterization of the position of the incorporated sulfates are listed in SI Materials and Methods.