Pontamine fast scarlet 4B: a new fluorescent dye for visualising cell wall organisation in radiata pine tracheids

Hemicelluloses are polysaccharides in plant cell walls that have beta-(1-->4)-linked backbones with an equatorial configuration. Hemicelluloses include xyloglucans, xylans, mannans and glucomannans, and beta-(1-->3,1-->4)-glucans. These types of hemicelluloses are present in the cell walls of all terrestrial plants, except for beta-(1-->3,1-->4)-glucans, which are restricted to Poales and a few other groups. The detailed structure of the hemicelluloses and their abundance vary widely between different species and cell types. The most important biological role of hemicelluloses is their contribution to strengthening the cell wall by interaction with cellulose and, in some walls, with lignin. These features are discussed in relation to widely accepted models of the primary wall. Hemicelluloses are synthesized by glycosyltransferases located in the Golgi membranes. Many glycosyltransferases needed for biosynthesis of xyloglucans and mannans are known. In contrast, the biosynthesis of xylans and beta-(1-->3,1-->4)-glucans remains very elusive, and recent studies have led to more questions than answers.

This review discuses the ultrastructural aspects of cell wall lignification and lignin topochemistry. Lignification results from the enzyme mediated polymerization of monolignols initiated by unknown factors (initiation sites) located at the corners of cells and in the middle lamella. Lignification results in the filling of pores within the carbohydrate matrix following a sequence from the outer regions of the wall towards the lumen. The amount and chemical characteristics of lignin vary across the cell wall, with the presence of reaction wood, and among cell types.