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Proteoglycans are proteins that carry sulfated glycosaminoglycans (GAGs). They help form and maintain morphogen gradients, guiding cell migration and differentiation during animal development. While no sulfated GAGs have been found in marine sponges, chondroitin sulfate (CS) and heparan sulfate (HS) have been identified in Cnidarians, Lophotrocozoans and Ecdysozoans. The general view that nematodes such as  Caenorhabditis elegans , which belong to Ecdysozoa, produce HS but only chondroitin without sulfation has therefore been puzzling. We have analyzed GAGs in  C. elegans  using reversed-phase ion-pairing HPLC, mass spectrometry and immunohistochemistry. Our analyses included wild type  C. elegans  but also a mutant lacking two HS sulfotransferases ( hst-6 hst-2 ), as we suspected that the altered HS structure could boost CS sulfation. We could indeed detect sulfated CS in both wild type and mutant nematodes. While 4- O -sulfation of galactosamine dominated, we also detected 6- O -sulfated galactosamine residues. Finally, we identified the product of the gene  C41C4.1  as a  C. elegans  CS-sulfotransferase and renamed it  chst-1  (CarboHydrate SulfoTransferase) based on loss of CS-4- O -sulfation in a  C41C4.1  mutant and  in vitro  sulfotransferase activity of recombinant C41C4.1 protein. We conclude that  C. elegans  indeed manufactures CS, making this widely used nematode an interesting model for developmental studies involving CS.

scientific reports

Nematodes join the family of chondroitin sulfate-synthesizing organisms: Identification of an active chondroitin sulfotransferase in Caenorhabditis elegans