Add time:09/03/2019         Source:sciencedirect.com

Most fungi have multiple Chitin (cas 1398-61-4) synthases (CSs) that may make chitin at different sites on the cell surface, at different times during growth, and in response to cell wall stress. The structure-based model for CS function is for transfer of GlcNAc from UDP-GlcNAc at the cytoplasmic face of the plasma membrane to the non-reducing end of a growing chitin chain, which is concomitantly translocated through a transmembrane channel formed by the synthase. Two aspects of CS mechanism are investigated: how chains might be initiated, and what governs how long they can get. First, it is shown that CSs incorporate free GlcNAc into di-N-acetylchitobiose and into insoluble chitin in a UDP-GlcNAc-dependent manner, and therefore that GlcNAc primes chitin synthesis. Second, average lengths of insoluble chitin chains were measured by determining the molar ratio of priming GlcNAc to chain-extending, UDP-GlcNAc-derived GlcNAc, and showed dependence on UDP-GlcNAc concentration, approaching a maximum at higher concentrations of substrate. These results, together with previous findings that 2-acylamido GlcN analogues prime formation of chitin oligosaccharides and stimulate chitin synthesis are discussed in the context of the structure-based model, and lead to the following proposals. 1) CSs may “self-prime” by hydrolyzing UDP-GlcNAc to yield GlcNAc. 2) A CS’s active site is not continuously occupied by a nascent chitin chain, rather, CSs can release chitin chains, then re-initiate, and therefore synthesize chitin chains in bursts. 3) The length of chitin chains made by a given CS will impact that CS’s contribution to construction of the fungal cell wall.

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