'Slipped Sandwich' Model for Chitin and Chitosan Perception in Arabidopsis

Chitin, a linear polymer of N-acetyl-D-glucosamine, and chitosans, fully or partially deacetylated derivatives of chitin, are known to elicit defense reactions in higher plants. We compared the ability of chitin and chitosan oligomers and polymers (chitin oligomers with degree of polymerization [DP] 3 to 8; chitosan oligomers with degree of acetylation [DA] 0 to 35% and DP 3 to 15; chitosan polymers with DA 1 to 60% and DP approximately 1,300) to elicit an oxidative burst indicative of induced defense reactions in Arabidopsis thaliana seedlings. Fully deacetylated chitosans were not able to trigger a response; elicitor activity increased with increasing DA of chitosan polymers. Partially acetylated chitosan oligomers required a minimum DP of 6 and at least four N-acetyl groups to trigger a response. Invariably, elicitation of an oxidative burst required the presence of the chitin receptor AtCERK1. Our results as well as previously published studies on chitin and chitosan perception in plants are best explained by a new general model of LysMcontaining receptor complexes in which two partners form a long but off-set chitin-binding groove and are, thus, dimerized by one chitin or chitosan molecule, sharing a central GlcNAc unit with which both LysM domains interact. To verify this model and to distinguish it from earlier models, we assayed elicitor and inhibitor activities of selected partially acetylated chitosan oligomers with fully defined structures. In contrast to the initial 'continuous groove', the original 'sandwich', or the current 'sliding mode' models for the chitin/chitosan receptor, the here-proposed 'slipped sandwich' model-which builds on these earlier models and represents a consensus combination of these-is in agreement with all experimental observations.