SummaryMixed-linkage (1 fi 3),(1 fi 4)-b-D-glucan (MLG) is widely considered to be a defining feature of the cell walls of plants in the Poales order. However, we conducted an extensive survey of cell-wall composition in diverse land plants and discovered that MLG is also abundant in the walls of the horsetail Equisetum arvense. MALDI-TOF MS and monosaccharide linkage analysis revealed that MLG in E. arvense is an unbranched homopolymer that consists of short blocks of contiguous 1,4-b-linked glucose residues joined by 1,3-b linkages. However, in contrast to Poaceae species, MLG in E. arvense consists mostly of cellotetraose rather than cellotetriose, and lacks long 1,4-b-linked glucan blocks. Monosaccharide linkage analyses and immunochemical profiling indicated that, in E. arvense, MLG is a component of cell walls that have a novel architecture that differs significantly from that of the generally recognized type I and II cell walls. Unlike in type II walls, MLG in E. arvense does not appear to be co-extensive with glucuroarabinoxylans but occurs in walls that are rich in pectin. Immunofluorescence and immunogold localization showed that MLG occurs in both young and old regions of E. arvense stems, and is present in most cell types apart from cells in the vascular tissues. These findings have important implications for our understanding of cell-wall evolution, and also demonstrate that plant cell walls can be constructed in a way not previously envisaged.Keywords: mixed-linkage (1 fi 3),(1 fi 4)-b-D-glucan, Equisetum arvense, cell wall, cell-wall evolution.
IntroductionAlmost all plant cells are surrounded by a complex, multifunctional and glycan-rich cell wall (Carpita and Gibeaut, 1993;Fry, 2004). As well as providing mechanical support, cell walls act as defensive barriers against pathogens and the environment, and play important roles in determining cell fate and in modulating growth and development (Bacic et al., 1988;Carpita and Gibeaut, 1993;O'Neill et al., 1990;Ridley et al., 2001). Materials from cell walls are also commercially important, and are a source of functional food ingredients, industrial fibres, nutraceuticals and feedstocks for second-generation bio-fuels (Himmel et al., 2007;Kim and Triplett, 2001;Reyna-Villasmil et al., 2007;Willats et al., 2006). In the non-lignified primary walls of growing cells, the main load-bearing components are cellulose microfibrils that are crossed-linked by non-pectic, non-cellulosic polysaccharides (Carpita and Gibeaut, 1993;Cosgrove, 2005). This network is embedded in a complex hydrated matrix of pectins and glycoproteins. The structure of cellulose is conserved across the plant kingdom, but the structures and relative amounts of other cell-wall components are highly variable, not just between plant species but also between organs, within tissues and even within cell-wall microdomains (McCann et al., 2007;Willats et al., 2001).
ª 2008 The Authors Journal compilation ª 2008 Blackwell Publishing LtdThe Plant Journal (2008Journal ( ) 54, 510-521 doi: 10.1111Jo...