In Silico Comparison of the Hemicelluloses Xyloglucan and Glucuronoarabinoxylan in Protecting Cellulose from Degradation

Total Page:16

File Type:pdf, Size:1020Kb

In Silico Comparison of the Hemicelluloses Xyloglucan and Glucuronoarabinoxylan in Protecting Cellulose from Degradation Computation 2015, 3, 336-353; doi:10.3390/computation3030336 OPEN ACCESS computation ISSN 2079-3197 www.mdpi.com/journal/computation Article In Silico Comparison of the Hemicelluloses Xyloglucan and Glucuronoarabinoxylan in Protecting Cellulose from Degradation Indrakumar Vetharaniam 1,*, Martin Upsdell 1, William J. Kelly 2, Graeme T. Attwood 2, Christina D. Moon 2 and Philip J. Harris 3 1 AgResearch Limited, Ruakura Research Centre, Private Bag 3123, Hamilton 3240, New Zealand; E-Mail: [email protected] 2 AgResearch Limited, Grasslands Research Centre, Private Bag 11008, Palmerston North 4442, New Zealand; E-Mails: [email protected] (W.J.K.); [email protected] (G.T.A.); [email protected] (C.D.M.) 3 School of Biological Sciences, The University of Auckland, Private Bag 92019, Auckland 1142, New Zealand; E-Mail: [email protected] * Author to whom correspondence should be addressed; E-Mail: [email protected]; Tel.: +64-7-838-5314; Fax: +64-7-838-5012. Academic Editor: Rainer Breitling Received: 10 April 2015/ Accepted: 17 June 2015 / Published: 6 July 2015 Abstract: We used a previously developed simulation model of a plant cell wall and its enzymatic degradation to compare the abilities of two hemicelluloses, glucuronoarabinoxylan (GAX) and xyloglucan (XG), to protect cellulose microfibrils (CMFs) from attack by cellulose-degrading enzymes. Additionally, we investigated the effect of XG abundance on the degradation rate of CMFs in the presence of the same enzymes. Simulations were run using hypothetical cell-wall compositions in which the numbers and arrangement of CMFs and (1,3;1,4)-β-glucan were kept constant, but the proportions of GAX and XG were altered. Scenarios considered walls with low and equal proportions of either GAX or XG, and also low, medium and high proportions of XG in the absence of GAX. The rate of CMF degradation was much lower in walls with GAX than walls with XG, except for early in the simulation when the reverse held, suggesting that XGs were protecting CMFs by competitive inhibition. Increasing XG content reduced both the degradation rate of CMFs and the percent of XG degraded, indicating that activity of enzymes decreased with XG density despite XG being degradable. Glucose oligosaccharide breakdown products were analysed on the basis of the originating polysaccharide and their Computation 2015, 3 337 degree of polymerisation (DP). The presence of GAX as opposed to equal amounts of XG had some significant effects on the amount and profile of breakdown products from XG and CMFs. Keywords: plant cell wall; lignocellulose; enzymatic degradation; enzymes; accessibility; competitive inhibition; rumen; second-generation biofuels, agent-based modelling; simulation model 1. Introduction Hemicelluloses are a group of plant cell-wall polysaccharides characterised by having a 1,4-β-linked backbone with an equatorial configuration [1]. They include heteroxylans, xyloglucans, 1,3;1,4-β-glucans and heteromannans, and often interact with cellulose (1,4-β-glucan), which is the most abundant polymer in plant cell walls [2] and occurs as crystalline cellulose microfibrils (CMFs) formed by the lateral association of parallel cellulose molecules [3,4]. The hemicellulose xyloglucan (XG) can account for 20%–25% of primary cell walls in eudicotyledons, such as forage legumes (family Fabaceae) and brassicas (family Brassicaceae), but accounts for only a small proportion (~2%–5%) of primary cell walls in the grass family (Poaceae) [2,5]. The predominant hemicellulose in both primary and secondary cell walls in this family is instead the heteroxylan glucuronoarabinoxylan (GAX), with the primary cell wall also containing varying, but often small proportions of 1,3;1,4-β-glucan [2]. Another heteroxylan, 4-O-methylglucuronoxylan (4-O-MeX), is the predominant hemicellulose in the secondary cell walls of the eudicotyledons. Despite the structural differences between the different hemicelluloses, they have traditionally all been assumed to have the same main role of providing structural strength to the cell wall by tethering the CMFs [1] as well as protecting CMFs from degradation by enzymes secreted by pathogenic microorganisms. This protective role of hemicelluloses is also probably important in reducing the enzymatic degradation of cellulose in plant cell walls (lignocellulose) in two applied contexts. First, in the rumens of ruminant animals which are populated by microorganisms that produce cell-wall degrading enzymes, and second, in the saccharification step in the production of second generation biofuels from plant biomass rich in cell walls, where enzyme mixtures including cellulases are used, particularly from the cellulolytic brown-rot fungus Trichoderma reesei (recently re-named Hypocrea jecorina). In the latter context, it has been found that pre-treatments of biomass that remove hemicelluloses (and lignin) can enhance the enzymatic hydrolysis of cellulose [6–8]. However, recent in vitro work with composites of hemicelluloses and CMFs has demonstrated that GAX and XG interact differently with CMFs [9], with GAX-CMF composites being mechanically less strong than XG-CMF composites [10]. There may also be differences between GAX and XG in their abilities to protect CMFs from degradation by cellulases. Two mechanisms of protection have been proposed: First, by the hemicellulose itself or oligosaccharides produced by its enzymatic degradation acting as an inhibitor of cellulose degradation, and second, by the hemicellulose forming a physical barrier limiting accessibility of cellulases to the CMFs. There is evidence that GAX and oligosaccharides (xylooligosaccharides) obtained from this by xylanase activity bind to the active sites Computation 2015, 3 338 of cellulases and competitively inhibit them [11–15]. This inhibition was found to be particularly marked for cellobiohydrolases, which are exo-cellulases that successively cleave cellobiose units from the reducing end (type I) or non-reducing end (type II) of cellulose or cellooligosaccharides. Importantly, the most abundant cellulase produced by H. jecorina, Cel7A, is a cellobiohydolase I. X-ray crystallographic studies showed that xylooligosaccharides bind to Cel7A at the entrance to the substrate-binding tunnel [15]. XG is hydrolysed by cellulases, but whether it or the oligosaccharides produced inhibit these cellulases appear not to have been investigated. Furthermore, there is no evidence to indicate whether GAX or XG forms a better physical barrier to the access of cellulases to CMFs. Here we report a modelling study in which we compared the effectiveness of GAX and XG in protecting CMFs from degradation by cellulases, and further investigated the role of XG density in protecting CMFs in cell walls that lack GAX or other heteroxylans. In this study, we created four hypothetical “cell walls”, similar to the cellulose-hemicellulose composites reported in [10] and used simulations of cellulase activity on these. 2. Methods We modelled the four composite cell walls by adapting a previously published 3-D model of a perennial ryegrass cell wall (RGCW) and its enzymatic degradation [16]. All four cell walls had several elements in common with RGCW, namely nine CMFs arranged in parallel (with the cellulose molecules 60 glucosyl residues long) with 35 (1,3;1,4)-β-glucans attached, all in their original configurations. The first model cell wall (Low XG) was created as a control, and contained 93 XGs, which were in the RGCW model [16]. The second model cell wall (Low GAX) had no XGs, but contained 93 GAXs from the RGCW in their original configurations. These two model cell walls allowed a comparison to be made of the abilities of XG and GAX to protect cellulose from degradation. The third and fourth model cell walls were designed to investigate the effect of XG density on cellulose degradation. These walls contained no GAXs and were formed by inserting 76 additional XGs into the Low XG model to obtain a cell wall with 169 XGs (Medium XG) and then a further 76 XGs to obtain a cell wall with 245 XGs (High XG). The number 245 was chosen to match the number of GAXs in the RGCW, with the Medium XG cell wall being intermediate between the two. The GAXs in Low GAX had an average degree of polymerisation (DP) of 27, with standard deviation (SD) of 9, and no glycosidic linkages that could be degraded by a cellulase/β-glucanase. The DP of the XG backbones averaged 29 for both Low and Medium XG and 30 for High XG (with an SD of 7 for all three). Low, Medium and High XG contained respectively 2742, 5104 and 7206 XG-sourced glucoses and respectively 2649, 4936 and 6961 glycosidic bonds degradable by a cellulase/β-glucanase. The CMFs in each cell wall contained 15,120 glucoses and 14,868 glycosidic bonds (1680 glucoses and 1652 glycosidic bonds per CMF). The (1,3;1,4)-β-glucans in each wall contained 1028 glucoses and 993 glycosidic bonds, with average DPs and numbers of glycosidic linkages of 29 and 28 respectively, both with SDs of 3. Dimensions were specified in terms of the radius of a glucose molecule, rg. As a reference, the Stoke’s radius of glucose is 0.365 nm [17]. Each wall had an approximate dimension of 120 rg deep (in the orientation of the CMF direction), 80 rg wide and 80 rg high. The volume not occupied by CMFs Computation 2015, 3 339 3 was approximately 690,000 rg . The total number of residues contained in all hemicelluloses of different types was 5311 for the Low GAX wall, and 5048, 8516 and 11,592 for the Low, Medium and High XG walls. Assuming each residue to be similar in size to a glucose molecule, the non-CMF residues occupied about 3.2% of the volume of the non-CMF space in the Low GAX wall, and about 3.1%, 5.2% and 7.0% of the volume of the non-CMF space in the Low, Medium and High XG walls.
Recommended publications
  • Plantphysiology134-1.Pdf
    The Galactose Residues of Xyloglucan Are Essential to Maintain Mechanical Strength of the Primary Cell Walls in Arabidopsis during Growth1 Marı´a J. Pen˜a2, Peter Ryden, Michael Madson, Andrew C. Smith, and Nicholas C. Carpita* Department of Botany and Plant Pathology, Purdue University, West Lafayette, Indiana 47907 (M.J.P., M.M., N.C.C.); and the Institute of Food Research, Norwich Research Park, Colney, Norwich NR4 7UA, United Kingdom (P.R., A.C.S.) In land plants, xyloglucans (XyGs) tether cellulose microfibrils into a strong but extensible cell wall. The MUR2 and MUR3 genes of Arabidopsis encode XyG-specific fucosyl and galactosyl transferases, respectively. Mutations of these genes give precisely altered XyG structures missing one or both of these subtending sugar residues. Tensile strength measurements of etiolated hypocotyls revealed that galactosylation rather than fucosylation of the side chains is essential for maintenance of wall strength. Symptomatic of this loss of tensile strength is an abnormal swelling of the cells at the base of fully grown hypocotyls as well as bulging and marked increase in the diameter of the epidermal and underlying cortical cells. The presence of subtending galactosyl residues markedly enhance the activities of XyG endotransglucosylases and the accessi- bility of XyG to their action, indicating a role for this enzyme activity in XyG cleavage and religation in the wall during growth for maintenance of tensile strength. Although a shortening of XyGs that normally accompanies cell elongation appears to be slightly reduced, galactosylation of the XyGs is not strictly required for cell elongation, for lengthening the polymers that occurs in the wall upon secretion, or for binding of the XyGs to cellulose.
    [Show full text]
  • Cell Wall Loosening by Expansins1
    Plant Physiol. (1998) 118: 333–339 Update on Cell Growth Cell Wall Loosening by Expansins1 Daniel J. Cosgrove* Department of Biology, 208 Mueller Laboratory, Pennsylvania State University, University Park, Pennsylvania 16802 In his 1881 book, The Power of Movement in Plants, Darwin alter the bonding relationships of the wall polymers. The described a now classic experiment in which he directed a growing wall is a composite polymeric structure: a thin tiny shaft of sunlight onto the tip of a grass seedling. The weave of tough cellulose microfibrils coated with hetero- region below the coleoptile tip subsequently curved to- glycans (hemicelluloses such as xyloglucan) and embedded ward the light, leading to the notion of a transmissible in a dense, hydrated matrix of various neutral and acidic growth stimulus emanating from the tip. Two generations polysaccharides and structural proteins (Bacic et al., 1988; later, follow-up work by the Dutch plant physiologist Fritz Carpita and Gibeaut, 1993). Like other polymer compos- Went and others led to the discovery of auxin. In the next ites, the plant cell wall has rheological (flow) properties decade, another Dutchman, A.J.N. Heyn, found that grow- intermediate between those of an elastic solid and a viscous ing cells responded to auxin by making their cell walls liquid. These properties have been described using many more “plastic,” that is, more extensible. This auxin effect different terms: plasticity, viscoelasticity, yield properties, was partly explained in the early 1970s by the discovery of and extensibility are among the most common. It may be “acid growth”: Plant cells grow faster and their walls be- attractive to think that wall stress relaxation and expansion come more extensible at acidic pH.
    [Show full text]
  • 1589168583 289 16.Pdf
    Plant Physiology and Biochemistry 136 (2019) 155–161 Contents lists available at ScienceDirect Plant Physiology and Biochemistry journal homepage: www.elsevier.com/locate/plaphy Research article Molecular insights of a xyloglucan endo-transglycosylase/hydrolase of radiata pine (PrXTH1) expressed in response to inclination: Kinetics and T computational study Luis Morales-Quintanaa,b, Cristian Carrasco-Orellanaa, Dina Beltrána, ∗ María Alejandra Moya-Leóna, Raúl Herreraa, a Functional Genomics, Biochemistry and Plant Physiology, Instituto de Ciencias Biológicas, Universidad de Talca, Campus Lircay s/n, Talca, Chile b Multidisciplinary Agroindustry Research Laboratory, Instituto de Ciencias Biomédicas, Universidad Autónoma de Chile, 5 poniente #1670, Talca, Chile ARTICLE INFO ABSTRACT Keywords: Xyloglucan endotransglycosylase/hydrolases (XTH) may have endotransglycosylase (XET) and/or hydrolase Plant cell wall (XEH) activities. Previous studies confirmed XET activity for PrXTH1 protein from radiata pine. XTHs could Xyloglucan endotransglycosylase/hydrolases interact with many hemicellulose substrates, but the favorite substrate of PrXTH1 is still unknown. The pre- Enzymatic parameters diction of union type and energy stability of the complexes formed between PrXTH1 and different substrates Xyloglycans (XXXGXXXG, XXFGXXFG, XLFGXLFG and cellulose) were determined using bioinformatics tools. Molecular Docking, Molecular Dynamics, MM-GBSA and Electrostatic Potential Calculations were employed to predict the binding modes, free energies of interaction and the distribution of electrostatic charge. The results suggest that the enzyme formed more stable complexes with hemicellulose substrates than cellulose, and the best ligand was − the xyloglucan XLFGXLFG (free energy of −58.83 ± 0.8 kcal mol 1). During molecular dynamics trajectories, hemicellulose fibers showed greater stability than cellulose. Aditionally, the kinetic properties of PrXTH1 en- zyme were determined.
    [Show full text]
  • Plant Xyloglucan Xyloglucosyl Transferases and the Cell Wall Structure: Subtle but Significant
    molecules Review Plant Xyloglucan Xyloglucosyl Transferases and the Cell Wall Structure: Subtle but Significant Barbora Stratilová 1,2 , Stanislav Kozmon 1 , Eva Stratilová 1 and Maria Hrmova 3,4,* 1 Institute of Chemistry, Centre for Glycomics, Slovak Academy of Sciences, Dúbravská cesta 9, SK-84538 Bratislava, Slovakia; [email protected] (B.S.); [email protected] (S.K.); [email protected] (E.S.) 2 Faculty of Natural Sciences, Department of Physical and Theoretical Chemistry, Comenius University, Mlynská Dolina, SK-84215 Bratislava, Slovakia 3 School of Life Science, Huaiyin Normal University, Huai’an 223300, China 4 School of Agriculture, Food and Wine, University of Adelaide, Glen Osmond, SA 5064, Australia * Correspondence: [email protected] or [email protected]; Tel.: +61-8-8313-7181 Academic Editor: László Somsák Received: 25 October 2020; Accepted: 26 November 2020; Published: 29 November 2020 Abstract: Plant xyloglucan xyloglucosyl transferases or xyloglucan endo-transglycosylases (XET; EC 2.4.1.207) catalogued in the glycoside hydrolase family 16 constitute cell wall-modifying enzymes that play a fundamental role in the cell wall expansion and re-modelling. Over the past thirty years, it has been established that XET enzymes catalyse homo-transglycosylation reactions with xyloglucan (XG)-derived substrates and hetero-transglycosylation reactions with neutral and charged donor and acceptor substrates other than XG-derived. This broad specificity in XET isoforms is credited to a high degree of structural and catalytic plasticity that has evolved ubiquitously in algal, moss, fern, basic Angiosperm, monocot, and eudicot enzymes. These XET isoforms constitute gene families that are differentially expressed in tissues in time- and space-dependent manners during plant growth and development, and in response to biotic and abiotic stresses.
    [Show full text]
  • Dietary Fibre from Whole Grains and Their Benefits on Metabolic Health
    nutrients Review Dietary Fibre from Whole Grains and Their Benefits on Metabolic Health Nirmala Prasadi V. P. * and Iris J. Joye Department of Food Science, University of Guelph, Guelph, ON N1G 2W1, Canada; [email protected] * Correspondence: [email protected] Received: 31 August 2020; Accepted: 30 September 2020; Published: 5 October 2020 Abstract: The consumption of whole grain products is often related to beneficial effects on consumer health. Dietary fibre is an important component present in whole grains and is believed to be (at least partially) responsible for these health benefits. The dietary fibre composition of whole grains is very distinct over different grains. Whole grains of cereals and pseudo-cereals are rich in both soluble and insoluble functional dietary fibre that can be largely classified as e.g., cellulose, arabinoxylan, β-glucan, xyloglucan and fructan. However, even though the health benefits associated with the consumption of dietary fibre are well known to scientists, producers and consumers, the consumption of dietary fibre and whole grains around the world is substantially lower than the recommended levels. This review will discuss the types of dietary fibre commonly found in cereals and pseudo-cereals, their nutritional significance and health benefits observed in animal and human studies. Keywords: dietary fibre; cereals; pseudo-cereals; chronic diseases 1. Introduction Consumers worldwide are interested in a healthy diet. Whole grain products, encompassing both cereals and pseudo-cereals, should constitute an important part of this healthy diet. The consumption of whole grain products is considered to have a beneficial effect on risk reduction of non-communicable diseases (NCD), including cardiovascular diseases, cancers, gastrointestinal disorders and type 2 diabetes [1–3].
    [Show full text]
  • Plant and Microbial Xyloglucanases
    Plant and microbial xyloglucanases: Function, Structure and Phylogeny Jens Eklöf Doctoral thesis Royal Institute of Technology School of Biotechnology Division of Glycoscience Stockholm 2011 Plant and microbial xyloglucanases: Function, Structure and Phylogeny ISBN 978‐91‐7415‐932‐5 ISSN 1654‐2312 Trita‐BIO Report 2011:7 ©Jens Eklöf, Stockholm 2011 Universitetsservice US AB, Stockholm ii Jens Eklöf Jens Eklöf (2011): Plant and microbial xyloglucanases: Function, Structure and Phylogeny School of Biotechnology, Royal Institute of Technology (KTH), Stockholm, Sweden Abstract In this thesis, enzymes acting on the primary cell wall hemicellulose xyloglucan are studied. Xyloglucans are ubiquitous in land plants which make them an important polysaccharide to utilise for microbes and a potentially interesting raw material for various industries. The function of xyloglucans in plants is mainly to improve primary cell wall characteristics by coating and tethering cellulose microfibrils together. Some plants also utilise xyloglucans as storage polysaccharides in their seeds. In microbes, a variety of different enzymes for degrading xyloglucans have been found. In this thesis, the structure‐function relationship of three different microbial endo‐xyloglucanases from glycoside hydrolase families 5, 12 and 44 are probed and reveal details of the natural diversity found in xyloglucanases. Hopefully, a better understanding of how xyloglucanases recognise and degrade their substrate can lead to improved saccharification processes of plant matter, finding uses in for example biofuel production. In plants, xyloglucans are modified in muro by the xyloglucan transglycosylase/hydrolase (XTH) gene products. Interestingly, closely related XTH gene products catalyse either transglycosylation (XET activity) or hydrolysis (XEH activity) with dramatically different effects on xyloglucan and on cell wall characteristics.
    [Show full text]
  • 2.19 Storage Plant Polysaccharides: Xyloglucans, Galactomannans, Glucomannans
    2.19 Storage Plant Polysaccharides: Xyloglucans, Galactomannans, Glucomannans K. Nishinari and M. Takemasa, Osaka City University, Osaka, Japan H. Zhang, Shanghai Jiao Tong University, Shanghai, China R. Takahashi, Gunma University, Gunma, Japan ß 2007 Elsevier Ltd. All rights reserved. 2.19.1 Introduction 614 2.19.2 Sources and Production of Xyloglucan, Galactomannan, and KGM 615 2.19.2.1 Xyloglucan 615 2.19.2.2 Galactomannans 616 2.19.2.2.1 Locust bean gum 616 2.19.2.2.2 Guar gum 617 2.19.2.2.3 Fenugreek gum 617 2.19.2.2.4 Tara gum 617 2.19.2.3 Konjac glucomannan (KGM) 617 2.19.3 Chemical Structure 617 2.19.3.1 Xyloglucan 617 2.19.3.2 Galactomannans 618 2.19.3.2.1 Modification of galactomannans 619 2.19.3.2.2 Purification of galactomannans 619 2.19.3.2.3 Composition analysis of galactomannans 620 2.19.3.3 Konjac Glucomannans 620 2.19.4 Solution Properties 621 2.19.4.1 Xyloglucan 622 2.19.4.2 Galactomannans 623 2.19.4.3 Konjac Glucomannans 625 2.19.5 Gelation of a Single Polysaccharide with and without Small Molecules 627 2.19.5.1 Gelation of Xyloglucan 627 2.19.5.1.1 Gelation of xyloglucan by addition of alcohol or sugar 628 2.19.5.1.2 Gelation of xyloglucan by removing galactose residues 628 2.19.5.1.3 Gelation of xyloglucan by addition of polyphenols 630 2.19.5.1.4 Gelation of xyloglucan by addition of iodide solution 631 2.19.5.1.5 Gelation of xyloglucan by addition of Congo red 631 2.19.5.2 Gelation of Galactomannan 633 2.19.5.3 Gelation of KGMs 634 2.19.5.3.1 Gelation of KGM in the presence of alkali 634 2.19.5.3.2 The cross-links
    [Show full text]
  • Studies on Recognition Mechanism for Branched Polysaccharides in Family 55 and 74 Glycoside Hydrolases
    Studies on recognition mechanism for branched polysaccharides in family 55 and 74 glycoside hydrolases (ファミリー55および74に属する糖質加水分解酵素における分岐多糖の認識メカニズム) Takuya Ishida 石田 卓也 Index 1. Introduction 1-1 Structures of polysaccharides in nature 1 1-2 Glycoside Hydrolases active on branched polysaccharides 3 1-2-1 classification by Carbohydrate Active enZymes (CAZy) data base 3 1-2-2 Catalytic mechanism 4 1-2-3 mode of action 5 1-2-4 protein folds and clan 7 1-3 Degradation of branched polysaccharides by GHs 10 1-4 Glycoside hydrolase family 74 and 55 1-4-1 Glycoside Hydrolase family 74 12 1-4-2 Glycoside Hydrolase family 55 15 1-5 Basidiomycete Phanerochaete chrysosporium 18 1-6 Aim of thesis 20 1-7 References 21 2. Characterization and X-ray crystallography of PcXgh74B 2-1 Introduction 28 2-2 Experimental procedure 30 2-3 Result 35 2-4 Discussion 48 2-5 Summary 53 2-6 References 54 3. X-ray crystallography of PcLam55A 3-1 Introduction 59 3-2 Experimental procedure 60 3-3 Result 63 3-4 Discussion 73 3-5 Summary 79 3-6 References 80 4. Conclusion 84 5. Acknowledgement Appendix Abbreviation BLAST basic local alignment search tool BMCC bacterial microcrystalline cellulose CAZy carbohydrate-active enzymes CBH cellobiohydrolase CBM carbohydrate binding modules CE carbohydrate esterase CMC carboxymethylcellulose DP degree of polymerization GH glycoside hydrolase GPC gel permeation chromatography GT glycosyl transferase HCA hydrophobic cluster analysis HPLC high performance liquid chromatography MS mass spectrometry NMR nuclear magnetic resonance PAHBAH p-hydroxybenzoic acid hydrazide PASC phosphoric acid swollen cellulose PL polysaccharides lyase RMSD root mean square deviation TXG xyloglucan from tamarind seed XGO4 oligosaccharides derived from TXG with DP of 7-9 XGO8 oligosaccharides derived from TXG with DP of 16-18 Enzymes AlgE4A A-module of mannuronan C-5-epimerase from Azotobacter vinelandii BsIFTase inulin fructotransferase from Bacillus sp.
    [Show full text]
  • Impacts of Dietary Xylans, Xylooligosaccharides, and Xylanases on Intestinal Health and Growth Performance of Monogastric Animals
    animals Review Friend or Foe? Impacts of Dietary Xylans, Xylooligosaccharides, and Xylanases on Intestinal Health and Growth Performance of Monogastric Animals Jonathan T. Baker, Marcos E. Duarte, Debora M. Holanda and Sung Woo Kim * Department of Animal Science, North Carolina State University, Raleigh, NC 27695, USA; [email protected] (J.T.B.); [email protected] (M.E.D.); [email protected] (D.M.H.) * Correspondence: [email protected] Simple Summary: Xylan is naturally present in typical feedstuffs fed to animals and has been shown to cause increased digesta viscosity reducing nutrient digestibility and growth. Xylooligosaccharides are sugar oligomers consisting of xylose units that can be extracted and purified from biomaterials for use as a prebiotic in monogastric feeds. Xylooligosaccharides can also be obtained from the hydrolysis of xylan either in the intestine of the animal or in-vitro through various techniques. The question of xylanase supplementation versus xylooligosaccharide supplementation as well as symbiosis of both on the intestinal health and performance of monogastric livestock is still up for debate. Xylanase inhibitors present in common cereal grains provide yet another obstacle to overcome and are found to be highly variable. As the fear of antibiotic resistance increases, novel approaches to improve growth performance and enhance intestinal health without the use of antibiotics also increase. The aim of this article is to review the structural difference and its impact on xylan in feeds, classification Citation: Baker, J.T.; Duarte, M.E.; Holanda, D.M.; Kim, S.W. Friend or and the use of various xylanases, as well as the production and use of xylooligosaccharides for the Foe? Impacts of Dietary Xylans, physiological effects on intestinal health and growth performance of monogastric animals.
    [Show full text]
  • Xyloglucan and Cellulose Form Molecular Cross-Bridges Connecting Root Border Cells in Pea (Pisum Sativum) T
    Plant Physiology and Biochemistry 139 (2019) 191–196 Contents lists available at ScienceDirect Plant Physiology and Biochemistry journal homepage: www.elsevier.com/locate/plaphy Short communication Xyloglucan and cellulose form molecular cross-bridges connecting root border cells in pea (Pisum sativum) T ∗ Marc Ropitauxa, , Sophie Bernarda,b, Marie-Laure Follet-Gueyea,b, Maïté Vicréa, Isabelle Boulognea, Azeddine Driouicha,b a Université de ROUEN, UFR des Sciences et Techniques, Laboratoire Glycobiologie et Matrice Extracellulaire Végétale, UPRES-EA 4358, Fédération de Recherche « Normandie-Végétal », FED 4277, F-76821, Mont-Saint-Aignan, France b Cell Imaging Platform (PRIMACEN-IRIB), Normandie Université, UNIROUEN, F-76821, Mont-Saint-Aignan, France ARTICLE INFO ABSTRACT Keywords: Pea (Pisum sativum) root cap releases a large number of living border cells that secrete abundant mucilage into Border cells the extracellular medium. Mucilage contains a complex mixture of polysaccharides, proteins and secondary Cellulose metabolites important for its structure and function in defense. Unlike xyloglucan and cellulose, pectin and Mucilage arabinogalactan proteins have been investigated in pea root and shown to be major components of border cell Polysaccharides walls and mucilage. In this study, we investigated the occurrence of xyloglucan and cellulose in pea border cells Root and mucilage using cytochemical staining, immunocytochemistry and laser scanning confocal microscopy. Our Xyloglucan data show that i) unlike cellulose, xyloglucan is highly present in the released mucilage as a dense fibrillary network enclosing border cells and ii) that xyloglucan and cellulose form molecular cross-bridges that tether cells and maintain them attached together. These findings suggest that secreted xyloglucan is essential for mucilage strengthening and border cell attachment and functioning.
    [Show full text]
  • Xyloglucan Endotransglycosylase, a New Wall-Loosening Enzyme Activity from Plants
    Biochem. J. (1992) 282, 821-828 (Printed in Great Britain) 821 Xyloglucan endotransglycosylase, a new wall-loosening enzyme activity from plants Stephen C. FRY,* Rachel C. SMITH, Kirstie F. RENWICK, David J. MARTIN, Sarah K. HODGE and Kathryn J. MATTHEWS Centre for Plant Science, Division of Biological Sciences, University of Edinburgh, The King's Buildings, Mayfield Road, Edinburgh EH9 3JH, U.K. 1. Cell-free extracts of all plants tested contained a novel enzyme activity (xyloglucan endotransglycosylase, XET) able to transfer a high-Mr portion from a donor xyloglucan to a suitable acceptor such as a xyloglucan-derived nonasaccharide (Glc4Xyl3GalFuc; XG9). 2. A simple assay for the enzyme, using [3H]XG9 and based on the ability of the [3H]polysaccharide product to bind to filter paper, is described. 3. The enzyme was highly specific for xyloglucan as the glycosyl donor, and showed negligible transglycosylation of other polysaccharides, including CM-cellulose. 4. The Km for XG9 was 50 tM; certain other 3H-labelled xyloglucan oligosaccharides also acted as acceptors, and certain non- radioactive xyloglucan oligosaccharides competed with [3H]XG9 as acceptor; the minimum acceptor structure was deduced to be: Xyl Xyl 4 4 Glc -+ Glc -+ Glc 5. The pH optimum was approx. 5.5 and the enzyme was less than half as active at pH 7.0. The enzyme was slightly activated by Ca2l, Mg2", Mn2+, spermidine, ascorbate and 2-mercaptoethanol, and inhibited by Ag+, Hg2+, Zn2+ and La3+. 6. XET activity was essentially completely extracted by aqueous solutions of low ionic strength; Triton X- 100, Ca2 , La3+, and Li' did not enhance extraction.
    [Show full text]
  • A Soil-Binding Polysaccharide Complex Released from Root Hairs Functions in 2 Rhizosheath Formation
    bioRxiv preprint doi: https://doi.org/10.1101/2021.04.15.440065; this version posted April 16, 2021. The copyright holder for this preprint (which was not certified by peer review) is the author/funder. All rights reserved. No reuse allowed without permission. 1 A soil-binding polysaccharide complex released from root hairs functions in 2 rhizosheath formation 3 4 Andrew F. Galloway1, Jumana Akhtar1, Emma Burak2, Susan E. Marcus1, Katie J. Field2, Ian 5 C. Dodd3*, Paul Knox1* 6 1Centre for Plant Sciences, Faculty of Biological Sciences, University of Leeds, Leeds LS2 7 9JT 8 2Department of Animal and Plant Sciences, University of Sheffield, Sheffield, S10 2TN 9 3Lancaster Environment Centre, Lancaster University, Lancaster, LA1 4YQ 10 * Author for correspondence (email: [email protected]) 11 Conflict of Interest Statement 12 The authors declare that there is no conflict of interest 13 14 A.F.G, J.A., E.B., K.J.F., I.C.D., and P.K. designed the research; E.B. carried out rhizosheath 15 analyses; A.F.G., J.A and E.B. carried out the isolation, soil-binding studies and 16 immunoanalyses of root exudates: S.E.M and P.K carried out seedling prints and 17 immunofluorescence studies. A.F.G and P.K. drafted the manuscript. All authors contributed 18 to and approved the final version of the article. 19 20 Keywords 21 Barley (Hordeum vulgare), root hairs, bald root barley (brb), root exudates, rhizosheath, soil, 22 polysaccharide, xyloglucan 23 24 One sentence summary 25 The root exudate of a root hairless mutant of barley, relative to wild type, has an altered pattern 26 of polysaccharide epitopes and lesser amounts of an acidic soil-binding polysaccharide 27 complex.
    [Show full text]