Xylan

Xylans are polysaccharides made up of β-1,4-linked xylose (a pentose sugar) residues with side branches of α-arabinofuranose and α-glucuronic acids and contribute to cross-linking of cellulose microfibrils and lignin through ferulic acid residues.[5] On the basis of substituted groups xylan can be categorized into three classes i) glucuronoxylan (GX) ii) neutral arabinoxylan (AX) and iii) glucuronoarabinoxylan (GAX).[6]

Studies on Arabidopsis mutants revealed that several Glycosyltransferases are involved in the biosynthesis of xylans.[7][8][9] Glycosyltransferases (GTs) catalyze the formation of glycosidic bonds between sugar molecules using nucleotide sugar as donor molecule.[8] In eukaryotes, GTs represent about 1% to 2% of gene products.[10] GTs are assembled into complexes existing in the Golgi apparatus. However, no xylan synthase complexes have been isolated from Arabidopsis tissues (dicot). The first gene involved in the biosynthesis of xylan was revealed on xylem mutants (irx) in Arabidopsis thaliana because of some mutation affecting xylan biosynthesis genes. As a result, abnormal plant growth due to thinning and weakening of secondary xylem cell walls was seen.[9] Arabidopsis mutant irx9 (At2g37090), irx14 (At4g36890), irx10/gut2 (At1g27440), irx10-L/gut1 (At5g61840) showed defect in xylan backbone biosynthesis.[9] Arabidopsis mutants irx7, irx8, and parvus are thought to be related to the reducing end oligosaccharide biosynthesis.[11] Thus, many genes have been associated with xylan biosynthesis but their biochemical mechanism is still unknown. Zeng et al. (2010) immuno-purified xylan synthase activity from etiolated wheat (Triticum aestivum) microsomes.[12] Jiang et al. (2016) reported a xylan synthase complex (XSC) from wheat that has a central core formed of two members of the GT43 and GT47 families (CAZy database). They purified xylan synthase activity from wheat seedlings through proteomics analysis and showed that two members of TaGT43 and TaGT47 are sufficient for the synthesis of a xylan-like polymer in vitro.[13]

Xylanase catalyzes the catabolism of xylan into xylose. Given that plants contain a lot of xylan, xylanase is thus important to the nutrient cycle.

Xylans play an important role in the integrity of the plant cell wall and increase cell wall recalcitrance to enzymatic digestion;[14] thus, they help plants to defend against herbivores and pathogens (biotic stress). Xylan also plays a significant role in plant growth and development. Typically, xylans content in hardwoods is 10-35%, whereas they are 10-15% in softwoods. The main xylan component in hardwoods is O-acetyl-4-O-methylglucuronoxylan, whereas arabino-4-O-methylglucuronoxylans are a major component in softwoods. In general, softwood xylans differ from hardwood xylans by the lack of acetyl groups and the presence of arabinose units linked by α-(1,3)-glycosidic bonds to the xylan backbone.[15]

The microanatomy, molecular physiology, and physical chemistry of the interactions between the three main structural biopolymers xylan, cellulose, and lignin in providing the rigidity of plant cell walls are topics of current research,[16][17] that may provide solutions in bioengineering, for example in biofuels manufacturing from maize, rice, and switchgrass.[17]

Xylan is used in different ways as part of our daily lives. For example, the quality of cereal flours and the hardness of dough are largely affected by the amount of xylan[6] thus, playing a significant role in bread industry. The main constituent of xylan can be converted into xylitol (a xylose derivative) which is used as a natural food sweetener, which helps to reduce dental cavities and acts as a sugar substitute for diabetic patients. It has many more applications in the livestock industry, because poultry feed has a high percentage of xylan.[6] Some macrophytic green algae contain xylan (specifically homoxylan[18]) especially those within the Codium and Bryopsis genera[19] where it replaces cellulose in the cell wall matrix. Similarly, it replaces the inner fibrillar cell-wall layer of cellulose in some red algae.

Xylan is one of the foremost anti-nutritional factors in common use feedstuff raw materials. Xylooligosaccharides produced from xylan are considered as "functional food" or dietary fibers[20] due their potential prebiotic properties.[21] Xylan can be converted in xylooligosaccharides by chemical hydrolysis using acids[22] or by enzymatic hydrolysis using endo-xylanases.[23] Some enzymes from yeast can exclusively converts xylan into only xylooligosaccharides-DP-3 to 7.[24]

Xylan is a major components of plant secondary cell walls which is a major source of renewable energy especially for second generation biofuels.[25] However, xylose (backbone of xylan) is a pentose sugar that is hard to ferment during biofuel conversion because microorganisms like yeast cannot ferment pentose naturally.[26]