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      Acetylation of woody lignocellulose: significance and regulation

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          Abstract

          Non-cellulosic cell wall polysaccharides constitute approximately one quarter of usable biomass for human exploitation. In contrast to cellulose, these components are usually substituted by O-acetyl groups, which affect their properties and interactions with other polymers, thus affecting their solubility and extractability. However, details of these interactions are still largely obscure. Moreover, polysaccharide hydrolysis to constituent monosaccharides is hampered by the presence of O-acetyl groups, necessitating either enzymatic (esterase) or chemical de-acetylation, increasing the costs and chemical consumption. Reduction of polysaccharide acetyl content in planta is a way to modify lignocellulose toward improved saccharification. In this review we: (1) summarize literature on lignocellulose acetylation in different tree species, (2) present data and current hypotheses concerning the role of O-acetylation in determining woody lignocellulose properties, (3) describe plant proteins involved in lignocellulose O-acetylation, (4) give examples of microbial enzymes capable to de-acetylate lignocellulose, and (5) discuss prospects for exploiting these enzymes in planta to modify xylan acetylation.

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          Rhamnogalacturonan II: structure and function of a borate cross-linked cell wall pectic polysaccharide.

          Rhamnogalacturonan II (RG-II) is a structurally complex pectic polysaccharide that was first identified in 1978 as a quantitatively minor component of suspension-cultured sycamore cell walls. Subsequent studies have shown that RG-II is present in the primary walls of angiosperms, gymnosperms, lycophytes, and pteridophytes and that its glycosyl sequence is conserved in all vascular plants examined to date. This is remarkable because RG-II is composed of at least 12 different glycosyl residues linked together by more than 20 different glycosidic linkages. However, only a few of the genes and proteins required for RG-II biosynthesis have been identified. The demonstration that RG-II exists in primary walls as a dimer that is covalently cross-linked by a borate diester was a major advance in our understanding of the structure and function of this pectic polysaccharide. Dimer formation results in the cross-linking of the two homogalacturonan chains upon which the RG-II molecules are constructed and is required for the formation of a three-dimensional pectic network in muro. This network contributes to the mechanical properties of the primary wall and is required for normal plant growth and development. Indeed, changes in wall properties that result from decreased borate cross-linking of pectin may lead to many of the symptoms associated with boron deficiency in plants.
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            Poplar carbohydrate-active enzymes. Gene identification and expression analyses.

            Over 1,600 genes encoding carbohydrate-active enzymes (CAZymes) in the Populus trichocarpa (Torr. & Gray) genome were identified based on sequence homology, annotated, and grouped into families of glycosyltransferases, glycoside hydrolases, carbohydrate esterases, polysaccharide lyases, and expansins. Poplar (Populus spp.) had approximately 1.6 times more CAZyme genes than Arabidopsis (Arabidopsis thaliana). Whereas most families were proportionally increased, xylan and pectin-related families were underrepresented and the GT1 family of secondary metabolite-glycosylating enzymes was overrepresented in poplar. CAZyme gene expression in poplar was analyzed using a collection of 100,000 expressed sequence tags from 17 different tissues and compared to microarray data for poplar and Arabidopsis. Expression of genes involved in pectin and hemicellulose metabolism was detected in all tissues, indicating a constant maintenance of transcripts encoding enzymes remodeling the cell wall matrix. The most abundant transcripts encoded sucrose synthases that were specifically expressed in wood-forming tissues along with cellulose synthase and homologs of KORRIGAN and ELP1. Woody tissues were the richest source of various other CAZyme transcripts, demonstrating the importance of this group of enzymes for xylogenesis. In contrast, there was little expression of genes related to starch metabolism during wood formation, consistent with the preferential flux of carbon to cell wall biosynthesis. Seasonally dormant meristems of poplar showed a high prevalence of transcripts related to starch metabolism and surprisingly retained transcripts of some cell wall synthesis enzymes. The data showed profound changes in CAZyme transcriptomes in different poplar tissues and pointed to some key differences in CAZyme genes and their regulation between herbaceous and woody plants.
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              O-Acetylation of Plant Cell Wall Polysaccharides

              Plant cell walls are composed of structurally diverse polymers, many of which are O-acetylated. How plants O-acetylate wall polymers and what its function is remained elusive until recently, when two protein families were identified in the model plant Arabidopsis that are involved in the O-acetylation of wall polysaccharides – the reduced wall acetylation (RWA) and the trichome birefringence-like (TBL) proteins. This review discusses the role of these two protein families in polysaccharide O-acetylation and outlines the differences and similarities of polymer acetylation mechanisms in plants, fungi, bacteria, and mammals. Members of the TBL protein family had been shown to impact pathogen resistance, freezing tolerance, and cellulose biosynthesis. The connection of TBLs to polysaccharide O-acetylation thus gives crucial leads into the biological function of wall polymer O-acetylation. From a biotechnological point understanding the O-acetylation mechanism is important as acetyl-substituents inhibit the enzymatic degradation of wall polymers and released acetate can be a potent inhibitor in microbial fermentations, thus impacting the economic viability of, e.g., lignocellulosic based biofuel production.
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                Author and article information

                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                21 May 2013
                2013
                : 4
                : 118
                Affiliations
                [1] 1Department of Forest Genetics and Plant Physiology, Swedish University of Agricultural Sciences Umea, Sweden
                [2] 2Department of Food and Environmental Sciences, University of Helsinki Helsinki, Finland
                Author notes

                Edited by: Maurice Bosch, Aberystwyth University, UK

                Reviewed by: Peter Ulvskov, Copenhagen University, Denmark; Paul Gatenholm, Chalmers University of Technology, Sweden

                *Correspondence: Ewa J. Mellerowicz, Department of Forest Genetics and Plant Physiology, Swedish University of Agricultural Sciences, S901-83 Umeå, Sweden. e-mail: ewa.mellerowicz@ 123456slu.se

                This article was submitted to Frontiers in Plant Biotechnology, a specialty of Frontiers in Plant Science.

                Article
                10.3389/fpls.2013.00118
                3659327
                23734153
                5ee6de84-5620-430b-9da6-652d22096551
                Copyright © Pawar, Koutaniemi, Tenkanen and Mellerowicz.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits use, distribution and reproduction in other forums, provided the original authors and source are credited and subject to any copyright notices concerning any third-party graphics etc.

                History
                : 23 February 2013
                : 15 April 2013
                Page count
                Figures: 1, Tables: 2, Equations: 0, References: 100, Pages: 8, Words: 0
                Categories
                Plant Science
                Mini Review Article

                Plant science & Botany
                cell wall,wood,biofuel,saccharification, o-acetylation,hemicellulose,acetyl esterase

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