Proteoglycans are a diverse family of molecules that are characterized by having one or more glycosaminoglycan (GAG) chains attached to a protein core. These molecules are found inside cells, intercalated into membranes, and in the ECM, and appear to be ubiquitously associated with animal cells. Heterogeneity is found at both the protein level and particularly at the GAG level, where variations in both the type of GAG chain and the extent of modification are found.

Characteristically the GAG chain is composed of a repeat disaccharide sequence in which one sugar is an amino sugar (N-acetylglucosamine or N-acetylgalactosamine) and the other a uronic acid (galactose in one case). The main types of GAGs found in the matrix are shown in Table 2. As can be seen, the majority, chondroitin sulfate (CS), derma-tan sulfate (DS), heparan sulfate (HS) and keratan sulfate (KS), are sulfated and are charged on each sugar residue, while hyaluronan (HA) is non-sulfated and charged only on alternate residues. In addition, hyaluronan is the only GAG chain that is not coval-ently attached to a protein core in vivo. The other GAG chains are attached to the protein core via a

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