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Xylan

Xylan is a hemicellulosic polysaccharide that serves as a major structural component of plant cell walls, characterized by a linear backbone of β-1,4-linked D-xylopyranose residues often substituted with side chains such as 4-O-methylglucuronic acid, arabinose, and acetyl groups, with variations depending on plant species (e.g., glucuronoarabinoxylan in grasses and glucuronoxylan in hardwoods).^[1]^[2] As the second most abundant biopolymer on Earth after cellulose, it constitutes 20–30% of the dry weight in dicot secondary cell walls and up to 50% in grasses, contributing significantly to the global lignocellulosic biomass of approximately 180 billion tons annually (as of 2023).^[1]^[3]^[4] In plants, xylan biosynthesis occurs in the Golgi apparatus through a complex of glycosyltransferases, including IRX9, IRX10, and IRX14 from the GT43, GT47, and GT8 families, using UDP-xylose as the primary substrate and incorporating modifications like acetylation via TBL proteins to facilitate proper folding and secretion.^[2]^[1] Functionally, xylan cross-links cellulose microfibrils and interacts with lignin, enhancing cell wall rigidity, flexibility, and vascular tissue development; mutants with reduced xylan content exhibit stunted growth, collapsed xylem vessels, and altered stress tolerance due to compromised secondary wall integrity.^[2]^[1] Its structural domains—typically twofold helical in the major form for cellulose binding and threefold helical in the minor form—enable these interactions, while substitutions influence wall biomechanics and digestibility.^[2] Beyond biology, xylan's abundance in sources like hardwoods (yielding 31–67% upon extraction), softwoods, and agricultural residues positions it as a renewable resource for industrial applications, including biofuel production through enzymatic hydrolysis to xylooligosaccharides and ethanol, as well as biomaterials such as biodegradable films, hydrogels for drug delivery, and food packaging coatings leveraging its oxygen barrier properties and hydroxyl groups for chemical functionalization.^[3]^[1] Efforts in metabolic engineering, such as CRISPR-mediated modifications to substitution patterns, aim to reduce biomass recalcitrance and improve processing efficiency for pulping, bioenergy, and sustainable products.^[2]

Chemical Structure and Composition

Backbone and Linkages

Xylan is classified as a hemicellulose, a heterogeneous group of polysaccharides found in plant cell walls, where it serves as a major structural component second only to cellulose in abundance. The core architecture of xylan features a linear backbone composed primarily of β-1,4-linked D-xylopyranose residues, connected through glycosidic bonds that confer rigidity and enable interactions with other cell wall polymers.^[1] This β-1,4 linkage pattern mirrors that of cellulose but substitutes D-xylose for D-glucose, resulting in a polymer that is typically less crystalline and more amenable to substitution. The structural formula of the xylan backbone can be represented as a repeating unit of β-D-xylopyranosyl-(1→4)-β-D-xylopyranosyl, denoted as [Xylp-(β1→4)-Xylp]_n, where n typically ranges from about 100 to 300 units in vascular plants, varying by source and extraction method. This configuration arises from the polymerization of β-D-xylopyranose monomers, with the glycosidic bond forming between the anomeric carbon (C1) of one residue and the C4 hydroxyl of the adjacent residue in the pyranose ring form.^[1] The uniformity of these linkages provides xylan with a relatively straight-chain conformation, facilitating its role in cross-linking cellulose microfibrils.^[1] In comparison to cellulose, which consists of β-1,4-linked D-glucopyranose units, xylan's backbone lacks the C6 hydroxymethyl group present in glucose, leading to a flatter, ribbon-like structure that enhances its ability to bind parallel to cellulose surfaces via hydrogen bonding. This structural difference allows xylan to adopt a twofold helical screw conformation when associated with cellulose, contrasting with its threefold helical form in aqueous solution, and promotes efficient packing within the cell wall matrix.^[5] The absence of the bulky C6 substituent reduces steric hindrance, enabling stronger lateral interactions and contributing to the overall mechanical strength of secondary cell walls.^[5] The backbone structure of xylan was elucidated in the early 20th century through classical carbohydrate chemistry techniques, including methylation analysis, which confirmed the β-1,4 glycosidic linkages by identifying the positions of free hydroxyl groups after derivatization and hydrolysis. Seminal work in the 1930s, building on prior isolations from plant materials, established the linear polyxylose nature of the chain via periodate oxidation and linkage studies.^[6]

Substituents and Variations

Xylan, a major hemicellulosic polysaccharide, exhibits structural diversity primarily through substituents attached to its β-(1→4)-linked D-xylopyranosyl backbone, which modulate its physicochemical properties and biological roles.^[1] Common substituents include 4-O-methyl-α-D-glucuronopyranosyl (MeGlcA) residues, typically linked at the O-2 position of xylose units, and α-L-arabinofuranosyl (Araf) groups attached via α-(1→3) linkages at the O-3 position.^[7] Acetyl groups are frequently esterified to the O-2 and/or O-3 positions of xylose, with ferulic acid esters often bound to the O-5 position of Araf residues, particularly in grass xylans, facilitating cross-linking with lignin.^[1] These modifications vary by plant species and tissue, contributing to xylan’s adaptability in cell wall architecture.^[7] Xylan subtypes are classified based on predominant substituents and their sources. Arabinoxylan (AX), enriched with Araf branches, predominates in grasses such as cereals, where it forms glucuronoarabinoxylan (GAX) with additional MeGlcA groups.^[1] In contrast, glucuronoxylan (GX), featuring MeGlcA as the main side chain, is characteristic of hardwood dicots like poplar and Arabidopsis secondary cell walls.^[7] Xylans are further categorized as acidic (e.g., GX and GAX, due to uronic acid content) or neutral (e.g., arabinoxylan lacking uronic acids, as in cereal grains).^[7] These variations influence xylan’s interactions within the cell wall matrix.^[1] The degree of substitution, or branching, in xylan typically ranges from 5% to 40% of xylosyl residues bearing side chains, depending on the plant source and developmental stage; this metric is calculated as:

\text{Branching degree} = \left( \frac{\text{number of side chains}}{\text{total xylose units}} \right) \times 100

^[7] Higher branching, as in grass GAX, enhances solubility, while lower levels in woody GX promote rigidity.^[1] Nuclear magnetic resonance (NMR) spectroscopy serves as a primary analytical method for identifying and quantifying xylan substituents, providing detailed structural insights through characteristic chemical shifts.^[8] For instance, the anomeric proton of Araf residues appears as a peak around δ 5.2 ppm in ¹H NMR spectra, distinguishing it from backbone xylose signals. Two-dimensional NMR techniques, such as HSQC, further resolve substitution patterns by correlating proton and carbon shifts.^[9]

Natural Occurrence

In Plant Cell Walls

Xylan constitutes a major component of plant cell walls, particularly in secondary walls, where it accounts for 19–35% of the dry weight in angiosperms and 7–14% in gymnosperms.^[10]^[11] It is predominantly located in the secondary cell walls of vascular tissues such as xylem and sclerenchyma, where it integrates into the matrix alongside cellulose and lignin.^[12]^[13] Within these structures, xylan interacts with cellulose microfibrils through hydrogen bonding, forming helical ribbons that enhance wall cohesion, while also establishing surface contacts with lignin to contribute to overall wall hydrophobicity.^[10]^[14] These interactions underpin xylan's key functional roles in plant tissues, providing structural support by cross-linking cellulose and lignin to maintain wall rigidity and integrity during vascular development.^[15] Additionally, xylan facilitates water retention within the cell wall matrix, creating hydration gradients that support tissue flexibility and prevent desiccation.^[14] In primary walls, it aids cell expansion by contributing to an extensible network that accommodates growth under turgor pressure.^[16] In monocotyledonous plants like grasses, arabinoxylans—xylans substituted with arabinose—form feruloylated gel networks that further reinforce cell walls against mechanical stress.^[17] The accumulation of xylan in plant cell walls correlates with the evolutionary emergence of vascular plants approximately 400 million years ago, marking a pivotal adaptation for terrestrial growth and structural complexity.^[18] This development maintained consistent substitution patterns across vascular lineages, enabling diverse wall architectures tailored to environmental demands.^[19] In plants, substituent variations—such as glucuronic acid in eudicots and arabinose in monocots—further modulate these plant-specific forms.^[10]

In Microorganisms and Algae

Xylan occurs in the cell walls of various algae, serving as a structural component distinct from its role in vascular plants. In red algae (Rhodophyta), it is often present as sulfated xylans embedded in the matrix alongside microfibrils of cellulose and neutral β-1,3-xylans. These sulfated forms have been identified in species such as Polysiphonia, where they contribute to the cell wall's rigidity and ion-binding properties. Sulfated polysaccharides, which may include minor amounts of xylans, can comprise up to 38% of the dry cell wall weight in red algae.^[20]^[21] In green algae (Chlorophyta), xylan structures vary by order, with β-1,3-linked xylans forming triple-helical microfibrils in the cell walls of Bryopsidales members, replacing cellulose as the primary fibrillar polysaccharide. For example, Acetabularia acetabulum contains β-(1→3)-xylans as part of its structural fibrils, often co-occurring with cellulose and contributing to mechanical support in these siphonaceous forms. These algal xylans typically feature shorter chains compared to plant counterparts, with occasional O-acetylation enhancing solubility and flexibility. Ecologically, xylan in algae functions as a carbon storage reservoir, representing a significant yet underappreciated pool of organic carbon in marine environments, particularly through β-1,3-xylans in red and green species. This storage role supports algal growth and resilience in nutrient-variable oceanic conditions.^[22]

Physical and Structural Properties

Crystallinity and Morphology

Xylan exhibits predominantly amorphous characteristics, with low crystallinity, in stark contrast to the highly ordered structure of cellulose. This low degree of crystallinity arises primarily from the branching and irregular packing of xylan chains, which disrupt the formation of extensive crystalline domains. Unlike cellulose Iβ, which achieves crystallinities exceeding 70% through parallel alignment of glucan chains in hydrogen-bonded sheets, xylan lacks such well-defined ordered regions, contributing to its role as a matrix polysaccharide that fills spaces between cellulose microfibrils in plant cell walls.^[23]^[24]^[25] In the solid state, xylan adopts twisted ribbon-like or helical conformations, often manifesting as twofold or threefold helical screw structures when associated with cellulose or in isolated forms. X-ray diffraction (XRD) analysis of native xylan reveals broad diffraction peaks centered around 2θ ≈ 20°, indicative of its disordered, amorphous arrangement rather than sharp crystalline reflections.^[26] These morphological features underscore xylan's flexibility, with the backbone's β-1,4 linkages allowing conformational adaptability that further limits long-range order. While native xylan in plant cell walls is predominantly amorphous, isolated or modified forms can exhibit crystalline polymorphs, such as xylan I hydrate.^[27] Factors such as chemical modification influence xylan's supramolecular organization; for instance, deacetylation of native xylan slightly enhances crystallinity by promoting self-assembly into hydrate crystals, as the removal of acetyl groups reduces steric hindrance and facilitates interchain hydrogen bonding. Electron microscopy observations of native xylan in plant cell wall contexts depict it as intricate fibrillar networks, intertwining with cellulose to form composite structures that provide mechanical support without rigid crystallinity.^[28]^[29]

Solubility and Rheological Behavior

Xylan, a major hemicellulose, exhibits limited solubility in neutral solvents due to extensive intra- and intermolecular hydrogen bonding between its β-1,4-linked xylopyranose units, rendering native forms largely insoluble in water at ambient conditions.^[30] However, solubility increases in alkaline environments, where solutions such as 1% NaOH or 1 M KOH disrupt these bonds, allowing dissolution; for instance, xylan from various sources dissolves completely in 1% NaOH but remains insoluble in 1 N HCl.^[31] Arabinoxylans, substituted variants with arabinose side chains, display enhanced water solubility compared to unsubstituted xylan, attributed to reduced chain packing and increased hydrophilicity from branching.^[32] In solution, xylan forms viscous dispersions, particularly at concentrations exceeding 1% (w/v), where water-extractable arabinoxylans can generate highly viscous or even gel-like states, with intrinsic viscosities typically ranging from 100 to 500 mL/g measured via the Huggins equation from dilute solution viscometry.^[33] Rheological profiles reveal pseudoplastic (shear-thinning) behavior, wherein apparent viscosity decreases with increasing shear rate, following models such as η = η₀ (1 + kγⁿ) with n < 1, reflecting chain alignment and disentanglement under flow.^[34] This non-Newtonian response is evident in both native and modified xylan solutions, as demonstrated in studies of carboxymethyl xylan derivatives.^[35] Factors influencing these properties include molecular weight, which spans 10⁴ to 10⁶ Da for typical xylans, higher values correlating with elevated viscosity, and the degree of substitution, where lower branching in debranched forms promotes aggregation and gelation under acidic conditions (low pH).^[36]^[37] Such behaviors are critical for processing, as they govern solution stability and flow during extraction or formulation.

Biosynthesis

Enzymatic Mechanisms

Xylan biosynthesis occurs primarily in the Golgi apparatus of plant cells, where a multi-enzyme complex facilitates the assembly of the β-1,4-linked xylose backbone and its substituents. The core xylosyltransferases, including IRX9 (GT43 family), IRX10 (GT47 family), and IRX14 (GT8 family) in Arabidopsis thaliana, form the xylan synthase complex (XSC) responsible for elongating the polysaccharide chain using UDP-xylose as the activated donor substrate. These enzymes catalyze the sequential addition of β-D-xylopyranosyl (Xylp) units, with IRX9 and IRX14 initiating and extending shorter chains, while IRX10 promotes processive polymerization to achieve higher degrees of polymerization.^[38]^[39] The biosynthetic pathway initiates with the formation of a specific reducing end sequence (RES), consisting of β-D-Xylp-(1→3)-α-L-Rhap-(1→2)-α-D-GalpA-(1→4)-β-D-Xylp, which may serve as a primer for chain elongation; this step involves enzymes such as IRX7, IRX8, and PARVUS. Elongation proceeds iteratively via the XSC, transferring Xylp residues to the non-reducing end of the growing chain. The basic reaction for elongation is:

\text{Xylan-OH} + \text{[UDP](/page/UDP)-Xylp} \rightarrow \text{Xylan-Xylp} + \text{UDP}

Termination mechanisms remain poorly understood, potentially involving unknown regulatory signals that cap the chain length, typically resulting in polymers of 100-200 xylose units.^[40] Side chain modifications occur concurrently or subsequently, with glucuronyltransferases such as GUX1 and GUX2 (GT8 family) adding α-D-glucuronosyl (GlcAp) residues at the O-2 position of backbone xyloses every 7-10 units in dicots. These enzymes utilize UDP-glucuronic acid as the donor, enhancing xylan's interaction with cellulose microfibrils post-secretion. Acetylation of the xylan backbone occurs in the Golgi apparatus, mediated by TBL family acetyltransferases (e.g., TBL1, TBL11, TBL19 in Arabidopsis), which transfer acetyl groups from acetyl-CoA to specific O-2 and O-3 positions of xylose residues, typically resulting in 20-30% acetylation degree to promote proper folding and prevent aggregation during secretion.^[41] In Arabidopsis, the genome encodes 10-15 genes across GT families (e.g., GT8, GT43, GT47) dedicated to xylan backbone synthases and side-chain transferases, reflecting functional redundancy and tissue-specific expression in vascular tissues.^[42]^[43]

Genetic and Regulatory Pathways

The biosynthesis of xylan in plants is governed by a suite of genes, prominently exemplified by the IRX (irregular xylem) gene family in Arabidopsis thaliana. Genes such as IRX9 and IRX14 encode glycosyltransferases that catalyze the formation of the β-1,4-xylan backbone in secondary cell walls.^[44] Mutations in these genes, as seen in irx9 and irx14 mutants, result in severely reduced xylan levels and collapsed xylem vessels due to weakened cell wall integrity.^[45] These genetic disruptions highlight the critical role of the IRX family in vascular development and structural support.^[46] Transcriptional regulation of xylan biosynthesis is orchestrated by key transcription factors, including NAC and MYB families, which activate gene expression during secondary cell wall formation. NAC factors like NST1 and SND1 initiate the regulatory cascade, directly upregulating downstream MYB proteins such as MYB46, which in turn promote the expression of IRX genes and other wall-related loci.^[47] Hormone signaling further modulates this process; auxin promotes vascular differentiation and secondary wall thickening by enhancing NAC-MYB activity, while brassinosteroids synergize with auxin to fine-tune xylan deposition in xylem tissues.^[48] These regulatory networks ensure coordinated xylan synthesis in response to developmental cues.^[49] Evolutionary conservation of xylan biosynthetic genes underscores their ancient origins, with homologs present in algae, reflecting shared glycosyltransferase mechanisms between algal and plant kingdoms. In streptophyte algae, GT43 family members synthesize xylan backbones similar to those in land plants, indicating pre-land plant innovation.^[50] Plant-specific expansions, particularly in the IRX and related GT families, occurred post-Devonian, coinciding with the evolution of vascular systems and complex secondary walls in early tracheophytes. Environmental stresses, such as drought, influence xylan production through abscisic acid (ABA) signaling, which upregulates secondary wall biosynthesis to enhance cell wall rigidity and maintain turgor. ABA activates transcription factors that boost xylan deposition, as evidenced by increased hemicellulose content in drought-stressed tissues, thereby improving plant resilience.^[51] This stress-responsive pathway integrates with developmental regulation to adapt cell walls dynamically.^[52]

Degradation and Breakdown

Biological Hydrolysis

Biological hydrolysis of xylan refers to the enzymatic breakdown of this hemicellulosic polysaccharide by microorganisms and other organisms, facilitating nutrient recycling in natural ecosystems. This process primarily involves xylanolytic enzymes that target the β-1,4-xylosidic linkages in the xylan backbone, enabling the degradation of plant cell wall components without the need for harsh chemical conditions.^[53] The core enzymes in xylan hydrolysis are endoxylanases, classified mainly in glycoside hydrolase families GH10 and GH11, which randomly cleave internal β-1,4 linkages to produce shorter xylooligosaccharides. GH10 endoxylanases typically exhibit broader substrate specificity, accommodating substituted xylans with side chains like arabinose or acetyl groups, while GH11 enzymes are more specialized for unsubstituted regions of the backbone. Complementing these are exoxylanases, which remove xylobiose units from the non-reducing ends of xylooligosaccharides, and β-xylosidases, which hydrolyze terminal xylose residues to yield free xylose monomers. These enzymes act sequentially or synergistically to achieve complete degradation.^[54]^[55]^[56] In many organisms, xylan hydrolysis occurs through multi-enzyme complexes or xylanolytic systems that enhance efficiency via synergistic interactions. For instance, in filamentous fungi such as Trichoderma reesei, endoxylanases work in concert with accessory enzymes like β-xylosidases and arabinofuranosidases, where initial endo-cleavage exposes substrates for exo-acting enzymes, leading to higher overall yields of hydrolytic products. This cooperation is evident in the degradation of complex xylans, where the combined action can increase sugar release by over twofold compared to individual enzymes. Recent advances as of 2025 include the identification of xylan-degrading systems in halotolerant bacteria like Bacillus altitudinis, offering potential for saline industrial processes.^[57]^[58]^[59]^[60] The kinetics of endoxylanase action generally follow Michaelis-Menten kinetics, with typical Km values ranging from approximately 1–13 mg/mL for beechwood or oat spelt xylan substrates, indicating moderate substrate affinity. Optimal pH for most fungal and bacterial endoxylanases falls between 4.5 and 6.0, aligning with the acidic environments of plant cell walls or microbial habitats, though some variants show broader stability up to pH 7.0. These parameters underscore the enzymes' adaptation to natural degradation niches.^[61]^[62]^[63] Biologically, xylan hydrolysis plays crucial roles in carbon cycling and ecological interactions. In the human gut microbiome, bacteria like Bacteroides species utilize xylan via polysaccharide utilization loci (PULs) that encode coordinated xylanases and transporters, breaking down dietary xylan to fermentable xylose and short-chain fatty acids that support host health. Similarly, in plant-pathogen interactions, pathogens such as Xanthomonas campestris employ xylanolytic systems to degrade host cell walls, facilitating tissue invasion and nutrient acquisition during infection. These processes highlight xylan's importance in microbial nutrition and pathogenesis.^[64]^[65]^[66]

Chemical and Industrial Methods

Xylan extraction from lignocellulosic biomass is predominantly achieved through alkaline methods, such as treatment with sodium hydroxide (NaOH) solutions, which disrupt the lignocellulosic matrix and solubilize hemicelluloses like xylan.^[67] Hot water extraction serves as an alternative, employing elevated temperatures (typically 160–200°C) to partially hydrolyze and release xylan without added chemicals.^[68] The efficiency of these processes is quantified by the yield equation:

\text{Yield} = \frac{\text{xylan extracted}}{\text{total xylan}} \times 100

with reported yields commonly ranging from 70% to 90%, influenced by factors like biomass type, pretreatment severity, and pH adjustment.^[67]^[69] Chemical hydrolysis of xylan involves acid-catalyzed cleavage of β-1,4-glycosidic bonds, often using 1–2% sulfuric acid (H₂SO₄) under hydrothermal conditions of 120–180°C to produce xylose or xylooligosaccharides.^[70]^[71] Alkaline hydrogen peroxide (H₂O₂) pretreatments provide an alternative for xylan solubilization, where the oxidant aids delignification under alkaline pH and temperatures around 80–120°C to enhance accessibility and minimize degradation products like furfural.^[72]^[73] To improve xylan’s processability, chemical modifications such as esterification are applied, substituting hydroxyl groups with acyl chains to enhance solubility in non-polar solvents like chloroform.^[74] On an industrial scale, xylan recovery occurs during kraft pulping in pulp mills, where delignification with hot alkaline white liquor (containing NaOH and Na₂S) at 160–170°C extracts hemicelluloses as soluble byproducts from hardwood or softwood chips.^[75]^[76] Autohydrolysis offers a chemical-free approach for xylan decomposition, conducted at approximately 180°C under pressure to promote internal acid catalysis via acetyl group release, yielding xylooligosaccharides (XOS) with a degree of polymerization (DP) of 2–10 while preserving the biomass for downstream uses.^[77]^[78] This method enhances safety by eliminating corrosive reagents and improves efficiency through reduced wastewater generation compared to acid-based processes.^[79]

Applications and Research

Industrial and Commercial Uses

In the pulp and paper industry, xylanases are employed to hydrolyze xylan, facilitating lignin removal during the bleaching process and thereby reducing chlorine consumption by up to 25-50% while improving pulp brightness.^[80] This enzymatic treatment enhances overall bleaching efficiency without significantly compromising fiber integrity. Additionally, retaining approximately 5-10% xylan in the pulp serves as a strength enhancer, improving tensile properties and fiber bonding in the final paper product through better carbohydrate-lignin associations.^[81] In the food industry, arabinoxylan, a substituted form of xylan, functions as a thickener in gluten-free products, where it helps mimic the viscoelastic properties of gluten to improve texture and stability.^[82] In baking applications, particularly for rye and wheat-based doughs, arabinoxylan enhances dough stability by increasing water absorption and gas retention, leading to improved loaf volume and crumb structure.^[83]^[84] For biofuel production, xylan from lignocellulosic biomass is hydrolyzed to xylose, which is then fermented into ethanol using engineered yeasts or bacteria, enabling second-generation biofuel pathways from non-food feedstocks.^[85] This process unlocks hemicellulosic fractions in agricultural waste, with global production potential estimated at 20-50 billion liters of ethanol annually from sources like crop residues and forestry byproducts.^[86]^[87] Beyond these sectors, xylan derivatives serve as fiber additives in animal feed, promoting gut health and nutrient utilization in poultry and swine diets by modulating digesta viscosity.^[88] In textiles, modified xylan acts as a dye thickener and binder, offering eco-friendly alternatives to synthetic polymers in printing pastes due to its biocompatibility and film-forming properties.^[89] The market for hemicellulose derivatives, including those from xylan, was valued at approximately USD 1.6 billion in 2023, driven by demand in these industrial applications.^[90]

Emerging Biomedical and Environmental Research

In recent years, xylooligosaccharides (XOS), short-chain carbohydrates derived from xylan, have emerged as promising prebiotics in biomedical research, particularly for modulating gut microbiota. Studies demonstrate that XOS supplementation selectively promotes the growth of beneficial bacteria such as Bifidobacterium species, enhancing microbial diversity and short-chain fatty acid production in both animal models and human trials. For instance, in a 2023 clinical study, XOS supplementation improved constipation symptoms by enriching Bifidobacterium abundance and modulating gut microbiota.^[91] Similarly, a 2021 mouse model showed XOS increasing Bifidobacterium populations throughout the intestine, correlating with anti-inflammatory metabolite profiles.^[92] These effects position XOS as a supportive therapy for gut health disorders, with human fecal microbiota analyses confirming bifidogenic activity at doses of 1-5 g/day without adverse effects. XOS also exhibit anti-inflammatory potential in inflammatory bowel disease (IBD) models, mitigating intestinal damage through microbiota modulation. Research from 2021 using a lipopolysaccharide-induced inflammation model in piglets found that XOS at 0.02% dietary levels reduced pro-inflammatory cytokines like TNF-α and IL-6, while preserving gut barrier integrity via increased mucin production.^[93] These findings, supported by reviews, highlight XOS's role in attenuating IBD symptoms by fostering anti-inflammatory microbial shifts, though clinical translation requires larger randomized trials. In materials science, xylan-based hydrogels have gained attention for controlled drug delivery due to their biocompatibility and tunable porosity. A 2018 study developed carboxymethyl xylan-acrylamide hydrogels using sodium bicarbonate as a pore-forming agent, achieving pore sizes of 100-130 µm and a 5-fluorouracil release rate of 71% over four hours at pH 7.4, ideal for sustained colonic delivery.^[94] These hydrogels exhibit swelling ratios up to 500% and biodegradation within weeks, offering advantages over synthetic polymers in reducing systemic toxicity. Xylan-chitosan nanocomposites further advance biomedical materials by combining natural polysaccharides for enhanced mechanical performance. Incorporating 12 wt% cellulose nanowhiskers into chitosan-xylan matrices yielded films with tensile strengths up to 57.2 MPa, a 70% improvement over unreinforced versions, alongside improved elongation at break for flexible wound dressings. These properties stem from strong hydrogen bonding interactions, enabling applications in tissue engineering scaffolds that support cell adhesion without cytotoxicity.^[95] Environmentally, xylan-derived biodegradable films address plastic pollution by providing renewable alternatives for packaging. A 2023 development of xylan plastic (XP) from industrial hemicellulose waste achieved tensile strengths of 55 MPa and full biodegradability in soil within 60 days, outperforming polyethylene in toughness (2.2 MJ/m³) while being non-toxic to mammalian cells.^[96] Such films reduce reliance on fossil-based plastics, with life-cycle analyses showing up to 80% lower carbon footprints when sourced from agricultural residues. In wastewater treatment, lignocellulolytic enzymes including xylanases facilitate the breakdown of lignocellulosic pollutants, enhancing bioremediation efficiency. A 2021 review demonstrated that fungal enzymes can degrade lignin derivatives in industrial effluents, reducing chemical oxygen demand through multi-enzyme action.^[97] Recent reviews emphasize immobilized enzymes on lignocellulosic supports for continuous treatment, achieving high pollutant removal in pulp mill wastewater without deactivation over multiple cycles. Key milestones include gene-editing advances for biofuel applications, such as CRISPR/Cas9 optimizations in microbial strains expressing xylanases, which boosted lignocellulosic hydrolysis yields for ethanol production. Additionally, research on marine algal β-1,3-xylan pathways has informed carbon capture strategies, where engineered algae utilize xylan-rich biomass to sequester CO₂ while yielding biofuels.

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May 24, 2010 · Various materials have been used as an amorphous standard, such as ball-milled cellulose, regenerated cellulose, and xylan or lignin powder.
[25]
Folding of xylan onto cellulose fibrils in plant cell walls revealed by ...
Dec 21, 2016 · The twofold screw xylan is spatially close to cellulose, and has similar rigidity to the cellulose microfibrils, but reverts to the threefold ...
[26]
Structure of native cellulose microfibrils, the starting point for ...
Dec 25, 2017 · It has long been known that the thin microfibrils of cellulose found in higher plants contain both crystalline and disordered chains, or chain ...
[27]
Bottom-up Construction of Xylan Nanocrystals in Dimethyl Sulfoxide
Feb 8, 2021 · The 2-fold helical structure is unique for xylan for which only a 3-fold helical form has been reported. The DMSO molecules participated in the ...
[28]
Synergistic Epichlorohydrin-Crosslinked Carboxymethyl Xylan ... - NIH
Aug 20, 2025 · ... broad peaks at 2θ = 20–21° and 26–27° are attributed to ... Comparison of X-ray diffraction (XRD) patterns between native xylan and ECX.
[29]
Crystalline nanoxylan from hot water extracted wood xylan at multi ...
Jul 1, 2024 · As discussed above, xylans readily self-assemble into crystal hydrates in case the biopolymer is less branched and deacetylated. However, the ...
[30]
Structural Imaging of Native Cryo-Preserved Secondary Cell Walls ...
We have adapted low temperature scanning electron microscopy (cryo-SEM) for imaging the nanoscale architecture of angiosperm and gymnosperm cell walls in their ...
[31]
Xylan - an overview | ScienceDirect Topics
This is generally explained by the fact that only one hydrogen bond is present between adjacent xylosyl residues,99 whereas two hydrogen bonds between adjacent ...
[32]
Solubility of xylan in some solvents. - ResearchGate
The xylan dissolves completely in alkali (1% NaOH), dissolves in hot water and is slightly soluble in cold water, and insoluble in acid (1N HCl).
[33]
Arabinoxylans as Functional Food Ingredients: A Review - PMC - NIH
Apr 1, 2022 · The viscosity of AX is positively correlated with solubility because of the high water-holding capacity of soluble AX [118,119]. Like solubility ...
[34]
Characterization of Water Extractable Arabinoxylans from a Spring ...
The intrinsic viscosity and viscosimetric molecular weight values for WEAX were 3.5 dL/g and 504 kDa, respectively. WEAX solution at 2% (w/v) formed gels ...
[35]
Shear rate dependence of viscosity for (a) xylan, (b) CX, and (c)...
4, the viscosities of xylan, CX, and CX-g-PHGH-1 all decreased with the increase of shear rate. In other words, they exhibited pseudoplastic or shearthinning ...
[36]
Shear rate dependence of viscosity for (a) xylan, (b) CX and (c) CX-g ...
With regard to SLX viscosity, the xylan was exhibited as a pseudoplastic whereby increasing the spindle speed or shear rate force damages the network ...Missing: equation | Show results with:equation
[37]
Reassigning the role of a mesophilic xylan hydrolysing family GH43 ...
Xylan, being a large polymer with an estimated molecular weight of ∼ 106 Da, requires a systematic set of xylan degrading enzymes to achieve its complete ...
[38]
Assembly of Debranched Xylan from Solution and on Nanocellulosic ...
This study focused on the assembly characteristics of debranched xylan onto cellulose surfaces. A rye arabinoxylan polymer with an initial arabinose/xylose ...Missing: gel | Show results with:gel
[39]
https://doi.org/10.1111/tpj.12641
[40]
https://doi.org/10.3389/fpls.2019.00176
[41]
https://doi.org/10.1016/j.molp.2015.01.006
[42]
https://doi.org/10.1111/tpj.12135
[43]
https://doi.org/10.1105/tpc.105.035501
[44]
Highly Branched Xylan Made by IRREGULAR XYLEM14 and ...
Xylans play important roles in strengthening secondary walls because most xylan-deficient mutants have collapsed xylem vessels, dwarfed stature, and reduced ...
[45]
Arabidopsis genes IRREGULAR XYLEM (IRX15) and IRX15L ...
Jan 19, 2011 · Direct measurement of xylan using PACE analysis demonstrates a 50% decrease in xylan that is likely to account for the decrease in cell wall ...
[46]
The Arabidopsis irregular xylem8 Mutant Is Deficient in ...
The majority of the irx mutants have been identified based on genetic screens for collapsed xylem cells (Turner et al., 2001). However, recent reports using ...
[47]
NAC-MYB-based transcriptional regulation of secondary cell wall ...
In this review, we summarize our current knowledge of the transcriptional regulation of SCW formation in plant cells.
[48]
Hormonal regulation of secondary cell wall formation
May 22, 2015 · During shoot and root tip growth, the combinatorial effect of hormones such as auxin, cytokinins, and brassinosteroids regulates xylem ...
[49]
Current Models for Transcriptional Regulation of Secondary Cell ...
Apr 3, 2018 · Transcriptional regulation in secondary wall formation has been extensively elucidated in the dicot model species Arabidopsis thaliana (Zhong ...
[50]
Identification of an algal xylan synthase indicates that there is ...
Feb 20, 2018 · Our analysis points to the evolutionary appearance of the various xylan synthesis‐specific genes identified to date, describing innovations of ...
[51]
Integrity of xylan backbone affects plant responses to drought - PMC
Jun 25, 2024 · Drought induces ABA signaling, resulting ... Drought signaling pathways partially overlap with other signaling events including cell wall ...
[52]
Abscisic acid regulates secondary cell-wall formation and lignin ...
ABA is the major hormone that controls a plant's ability to survive water-deficit stress, and the content of lignin is increased and lignin biosynthesis-related ...
[53]
Xylan degradation, a metabolic property shared by rumen and ... - NIH
Xylan is the second most abundant plant cell wall polysaccharide next to cellulose in monocotyledonous grasses, which comprise the primary feedstuff of grazing ...Missing: EPS | Show results with:EPS
[54]
Functional Characterization of the GH10 and GH11 Xylanases ... - NIH
We found that the GH11 xylanase may be more efficient at catalyzing biomass degradation than the GH10 xylanase.
[55]
Synergistic mechanism of GH11 xylanases with different action ...
May 10, 2021 · Aspergillus niger, possessing numerous xylan degrading isozyme-encoding genes, are highly effective xylan degraders in xylan-rich habitats.Missing: biofilms | Show results with:biofilms
[56]
Exo-Xylanase - an overview | ScienceDirect Topics
Primarily, endoxylanase breaks the main chain of xylan for the formation of xylo-oligomers and β-xylosidase converts xylo-oligomers into xylose monomers.
[57]
Synergistic effect of Aspergillus niger and Trichoderma reesei ...
Aug 13, 2014 · This was reflected in an overall strong synergistic effect in releasing sugars during saccharification using A. niger and T. reesei enzyme sets.2.3 Trichoderma Reesei And... · 2.5 Enzymatic Assays · 3 Results
[58]
Biochemical characterization of a novel exo-oligoxylanase from ...
Jul 29, 2019 · The complete degradation of xylans requires synergistic reaction of several xylanolytic enzymes, in which xylanases (EC 3.2.1.8) and β- ...
[59]
Co-production of Xylooligosaccharides and Xylose From Poplar ...
Feb 1, 2021 · This study focuses on enzymatic hydrolysis of poplar sawdust xylan for production of XOS and xylose by a GH11 endo-1,4-β-xylanase MxynB-8 and a GH39 β- ...
[60]
Purification, characterization, and mode of action of endoxylanases ...
Jan 1, 1992 · The Km and Vmax values with water-soluble oat spelts xylan as substrate were 2.6 mg ml-1 and 33.6 mumol min-1 mg-1 for endoxylanase 1 and ...
[61]
Application of an endo-xylanase from Aspergillus japonicus in the ...
... Vmax of 467.4 ± 30.38 μmol/min/mg, with a km of 2.59 ± 0.17 mg/mL, a kcat of 253.95 ± 16.51 s−1 and a kcat/km value of 98.05 ± 4.41 mL s−1 mg−1. The endo- ...
[62]
Characterization and structural analysis of the endo-1,4-β-xylanase ...
Oct 13, 2023 · When the effect of pH on the enzymatic activity of TsaGH11 was investigated, the optimum pH was observed to be slightly acidic, ranging from pH ...
[63]
Xylan utilization in human gut commensal bacteria is ... - NIH
Here we demonstrate that two human gut commensal Bacteroides are equipped with unique enzymes that allow degradation of xylan, a common hemicellulose in human ...
[64]
Xylan degradation by the human gut Bacteroides xylanisolvens ...
May 4, 2016 · Xylans are not degraded by human digestive enzymes in the upper digestive tract and therefore reach the colon where they are subjected to ...
[65]
The xylan utilization system of the plant pathogen Xanthomonas ...
Feb 27, 2013 · Xylan is a major structural component of plant cell wall and the second most abundant plant polysaccharide in nature.
[66]
Delignification outperforms alkaline extraction for xylan ...
Nov 20, 2016 · For other grass-like feedstocks, such as for wheat straw, alkaline extraction showed also good xylan yields (70–90%) (Escarnot et al., 2011; ...
[67]
Xylan and xylose decomposition during hot water pre-extraction
Mar 1, 2021 · By increasing the initial pH of the xylan solution, the furfural concentration can be reduced effectively to 2% and the formation of formic acid ...
[68]
Preparation and characterization of xylan by an efficient approach ...
Han and Xie (2018) extracted xylan from corn stalks with microwave pretreatment (650 W, 21 min) assisted alkali method. The yield of xylan can reach 76.57 %.
[69]
Kinetic Model of Xylose Dehydration for a Wide Range of Sulfuric ...
Jun 4, 2020 · This paper presents a robust kinetic model for the dehydration of xylose in concentrated sulfuric acid (ie, 0.1– 2 M) at 120–160 °C
[70]
The Kinetics Studies on Hydrolysis of Hemicellulose - PMC - NIH
Nov 18, 2021 · It obtained the highest xylooligosaccharides yield of 45.18% from 1% sulfuric acid at 120°C for 60 min. Meanwhile, it achieved the highest ...
[71]
Effect of sulfuric acid on production of xylooligosaccharides and ...
Hydrogen peroxide reacts with acetic acid with sulfuric acid as a catalyst to generate peracetic acid, which has strong oxidizing properties and is used to ...Missing: H2SO4 | Show results with:H2SO4
[72]
Xylan solubilisation from oil palm frond and sago palm bark via in ...
Sep 15, 2025 · The optimal conditions were identified using mixture of 2 % NH4OH: 1 % H2O2: 1 % Na2S2O4 for OPF and 2 % NH4OH: 1 % H2O2: 1 % NaBH4 for SPB, ...
[73]
Syntheses and characterization of xylan esters - ScienceDirect.com
Aug 17, 2012 · Chemical modification of xylan brought about by esterification resulted to an improvement in its solubility in CHCl3 and increase in its ...
[74]
Hemicellulose extraction from aspen chips prior to kraft pulping ...
In this study kraft white liquor with 18% active alkali and 25% sulfidity was used to partially extract xylan from commercial aspen chips.
[75]
On the recovery of hemicellulose before kraft pulping - BioResources
Both parameters were close to the values (721 mL/g intrinsic viscosity and 88.4% brightness) determined for a commercial pulp used as a reference.
[76]
An integrated process to produce prebiotic xylooligosaccharides by ...
In this study, raw alkali-extracted xylan was subjected to autohydrolysis at 180 °C for 40 min, following preliminary experiments to establish the optimal ...
[77]
A sustainable process for procuring biologically active fractions of ...
Jul 29, 2019 · Although ultrafiltration can separate the XOS with DP of 2–10 from prehydrolyzate containing higher molar mass XOS and lignin fragments, the ...
[78]
Production of Xylooligosaccharides from Jiuzao by Autohydrolysis ...
The autohydrolysis treatment condition with the highest XOS yield changed from 190 °C to 180 °C compared to that without enzymatic hydrolysis, indicating the ...
[79]
Applicability of microbial xylanases in paper pulp bleaching: A review
May 1, 2014 · This review summarizes the application of xylanases in the bleaching of pulp, with emphasis on the mechanism and effects of xylanase treatment on pulp and paper
[80]
[PDF] IMPROVEMENT OF PAPER STRENGTH BY INCREASING THE ...
Extracted xylan from beech dissolving pulp and eucalyptus kraft pulp was precipitated on unrefined, bleached, once- dried softwood kraft.Missing: enhancer | Show results with:enhancer
[81]
Technological properties of arabinoxylans as baking additives
Dec 23, 2021 · Arabinoxylans can be used as a baking additive to improve dough consistency, increase loaf volumes, improve crumb structure and decrease ...
[82]
Pentosan extraction from rye bran on pilot scale for application in ...
Arabinoxylan, a pentosan, is the most important thickening agent and key component in rye for bread making. The idea of the study was to use pentosans for ...
[83]
[PDF] Impact of isolated and chemically modified dietary fiber on bakery ...
Oct 2, 2023 · This review focuses on the impact of isolated fiber ingredients, including arabinoxylan, β‐glucan, cellulose, resistant starch, and pectin, on ...
[84]
Enzymatic deconstruction of xylan for biofuel production - PMC - NIH
This review focuses on current understanding of the molecular basis for substrate specificity and catalysis by enzymes involved in xylan deconstruction.
[85]
[PDF] A review of production of bioethanol from agricultural wastes
Oct 11, 2023 · Among the four agricultural wastes stated, rice straw has the highest potential production of bioethanol, with 205 billion liters produced.
[86]
Bioethanol Market Size, Share & Value | Analysis Report [2032]
For instance, The U.S. Department of Energy (DOE) reports that cellulosic ethanol has the potential to produce up to 100 billion gallons of fuel annually in the ...Bioethanol Market Trends · Bioethanol Market Growth... · Bioethanol Market...<|separator|>
[87]
Friend or Foe? Impacts of Dietary Xylans, Xylooligosaccharides, and ...
Xylan is naturally present in typical feedstuffs fed to animals and has been shown to cause increased digesta viscosity reducing nutrient digestibility and ...
[88]
Chemical Modification of Xylan - IntechOpen
It is used in particular in the food, cosmetic and pharmaceutical sectors, as a thickener, emulsifier or gelling agent. In addition, its binding properties ...2. Industrial Upgrading Of... · 2.1 A New Way Of Chemical... · 4. The Results
[89]
Hemicellulose Market Size, Share, Scope, Trends & Forecast
Rating 4.5 (45) Hemicellulose Market size was valued at USD 1.6 Billion in 2023 and is projected to reach USD 2.7 Billion by 2031, growing at a CAGR of 4.6%Missing: derivatives | Show results with:derivatives