Chapter 27 : Plant Cell Wall and Chitin Degradation

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A large group of fungi has specialized in the degradation of the complex plant cell walls. The natural resistance of plant cell walls to microbial and enzymatic decomposition is largely responsible for the high cost of lignocellulose conversion. A polymer that is structurally related to the plant cell wall polysaccharide cellulose but does not occur in plants is chitin. Due to practical applications, most strategies to use plant cell walls in biotechnological processes exploit the cellulose and hemicellulose sugars following depolymerization. Most of the plant cell wall polysaccharides occur in the form of lignocelluloses. Xyloglucan is quantitatively the predominant hemicellulosic polysaccharide of dicotyledons and nongraminaceous monocotyledons, comprising up to 20% of the plant cell wall. Degradation and catabolism of the individual carbon sources present in complex mixtures follow a mainly energy-driven hierarchy, but adaptation of saprobic and plant pathogenic fungi to their habitats has resulted in species-specific carbon source priorities. A list of fungal glycoside hydrolases (GH) and carbohydrate esterases (CE) that are involved in the degradation of the side chains of plant cell wall polysaccharides is provided. Fungi depolymerize pectin by using not only hydrolytic enzymes (PGAs) but also enzymes that cleave polysaccharide chains via a β-elimination mechanism, resulting in the formation of a Δ-4,5-unsaturated bond at the newly formed, nonreducing end. Many aspects of chitin degradation resemble that of cellulose and have potential impacts on the development of second-generation (“lignocellulosic”) bioethanol. N-acetylglucosamine, the monomer of chitin, is an excellent carbon source for but only induces N-acetylglucosaminidases.

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27
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Image of FIGURE 1

Schematic structure of major hemicelluloses in plant cell walls. Hemicelluloses consist of branched polysaccharides that have a backbone composed of 1,4-linked β-D-pentosyl/ hexosyl residues. The predominant hemicellulose in many primary walls is xyloglucan, while the other hemicelluloses, including glucuronoxylan, arabinoxylan, arabinoglucuronoxylan, and galactoglucomannan, occur in both primary and secondary cell walls.

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27
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Image of FIGURE 2

Cellulose degradation and regulation. Cellulose is extracellularly degraded by the enzymatic attack of three types of enzymes including cellbiohydrolase (CBH), endoglucanase (EG), and β-glucosidase (BGL). Swollenin (SWO) disrupts the crystalline structure of the cellulose and supports the enzymatic cellulose breakdown. Negative-acting (CRE1, HAP complex, and ACE1) and positive-acting (ACE2 and XYR1) regulators control cellulase expression on the level of transcription.

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27
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Image of FIGURE 3

Schematic structure of different pectins. Rhamnogalacturonan I and homogalacturonan are the two main structures of the plant cell wall pectin. The main chain of rhamnogalacturonan I (shown on top) is decorated with different arabinan, galactan, and arabinogalactan side chains (hairy region), whereas on the main chain of homogalacturonan, only methyl and acetyl esters are found (smooth region).

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27
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Image of FIGURE 4

Fungal D-xylose and L-arabinose catabolism. The fungal catabolic pathway for D-xylose and L-arabinose is an interconnected pathway that channels both sugars in the pentose phosphate pathway. NADPH-dependent reductions alternate with NAD-dependent oxidations before D-xylulose is finally phosphorylated by xylulokinase to D-xylulose 5-phosphate. In the main enzyme for the first step in both pathways is the D-xylose reductase XYL1.

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27
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Image of FIGURE 5

Fungal D-galactose catabolism. D-Galactose occurs in nature in the two anomeric forms α- and β-D-galactose. The galactokinase of the classical Leloir pathway (left) is specific for α-D-galactose, and therefore, β-D-galactose has to be epimerized to the α-anomer before it can enter this pathway. A second pathway (right) was found recently in and . It starts with the reduction of both anomeric forms of D-galactose to galactitol. Two hypothetical drafts for the further degradation of galactitol are summarized.

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27
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Fungal glycoside hydrolases (GH) and carbohydrate esterases (CE) involved in the degradation of the side chains of plant cell wall polysaccharides

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27
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Properties of fungal chitinases from phylogenetic subgroups A, B, and C

Citation: Kubicek C, Seidl V, Seiboth B. 2010. Plant Cell Wall and Chitin Degradation, p 396-413. In Borkovich K, Ebbole D (ed), Cellular and Molecular Biology of Filamentous Fungi. ASM Press, Washington, DC. doi: 10.1128/9781555816636.ch27

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