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. 2017 Dec 7;8:1973. doi: 10.1038/s41467-017-02076-0

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© The Author(s) 2017

Open Access This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons license, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commonslicense, unless indicated otherwise in a credit line to the material. If material is not included in the article’sCreative Commons license and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this license, visit http://creativecommons.org/licenses/by/4.0/.

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Fig. 1 — Models for heparan sulfate chain structures/organizations. a NDST (N-deacetylase/N-sulfotransferase). The first modification step of the heparan sulfate (HS) precursor chain is the conversion of N-acetyl moiety (NA) to N-sulfo moiety (NS), which is catalyzed by NDST. b The NA/NS domain model of a single HS chain. A single chain of HS generally contains two types of domains, N-acetyl (NA) and N-sulfo (NS). c Our model of distinct N-acetyl-rich HS and N-sulfo-rich HS clusters. See the main text for details.