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Fate of Non-Structural Carbohydrates Throughout the Stem of Teak

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Fate of Non-Structural Carbohydrates Throughout the Stem of Teak Available online at http://www.ifgdg.org Int. J. Biol. Chem. Sci. 12(3): 1102-1112, June 2018 ISSN 1997-342X (Online), ISSN 1991-8631 (Print) Original Paper http://ajol.info/index.php/ijbcs http://indexmedicus.afro.who.int Heartwood formation process in teak (Tectona grandis L. f): fate of non-structural carbohydrates and characterization of forsythoside B Bobelé Florence NIAMKE1*, Augustin Amissa ADIMA1, Kati-Coulibaly SERAPHIN2, Nadine AMUSANT3 and Christian JAY-ALLEMAND4 1 Laboratoire de Procédés Industriels de Synthèse de l’Environnement et de l’Energie Nouvelle (LAPISEN), Unité Chimie des Eaux et des Substances Naturelles, Institut National Polytechnique Houphouët Boigny, B.P. 1313 Yamoussoukro, Côte d’Ivoire. 2 Laboratoire de Nutrition et Pharmacologie de l'UFR Biosciences 22 BP 582 Abidjan 22, Côte d'Ivoire. 3Unité Mixte de Recherche (UMR) ECOFOG1, 2091 Route de Baduel - BP 792 - 97337 Cayenne cedex. Guyane, France. 4 Laboratoire de Biochimie et de Physiologie Végétale, Université de Montpellier II, UMR 1208 IATE (UM, INRA, SupAgro) CC024, Place Eugène Bataillon, 34095 Montpellier Cedex 05 France. * Corresponding author; E-mails: [email protected] / [email protected] ABSTRACT Heartwood formation is an important process in perennial plants as trees. Non-structural carbohydrates (NSC) storage in wood is an important feature of heartwood formation properties before and after wood drying. To better understand how NSC are stored and transformed in teak wood, their radial distribution was studied before and after chemical hydrolysis by using a spectrophotometric method coupled to enzymatic assays. In sapwood, NSC (starch, glucose, fructose and sucrose) and condensed NSC (mainly phenolic glucosides) were strongly accumulated respectively 100.9 5.9 and 30.2 5.5 mol.g.dw-1. They decreased abruptly (5-10 folds lower) from sapwood to heartwood. However, if the proportion of condensed glucose was 3 folds higher than that of non-condensed glucose in the sapwood, this ratio increases 20 folds in the heartwood indicating that glucosylation process could occur. The forsythoside B, a trisaccharide of caffeic acid present in sapwood, was likely hydrolyzed during heartwood formation. Our results show that high proportions of starch and soluble NSC (glucose, fructose and sucrose; 80%) and condensed NCS (essentially glucose after chemical hydrolysis of the extract; 20%) were mobilized in the sapwood and used in the transition zone leading to their abrupt depletion in the heartwood where condensed NSC dominated (75-90%). The unmetabolized glucose was likely stored mainly by etherification. Significant correlations between glucose contents (before and after hydrolysis) and natural durability (-0.43≤ R≤ -0.67) were found. These results indicate that high levels of NSC mobilization in the stem could be a relative efficient strategy of teak for enhancing its heartwood natural durability. © 2018 International Formulae Group. All rights reserved. Keywords: Teak, sapwood, non-structural carbohydrate, hydrolysis, natural durability. INTRODUCTION species and is one of the most valuable Teak (Tectona grandis L. f.) is tropical timbers on the international market indigenous to India, Myanmar, Thailand and (Bhat and Ma, 2004). Teak has long been Laos. It was introduced in the tropics as exotic known to possess highly durable wood, due © 2018 International Formulae Group. All rights reserved. 4091-IJBCS DOI: https://dx.doi.org/10.4314/ijbcs.v12i3.2 B. F. NIAMKE et al. / Int. J. Biol. Chem. Sci. 12(3): 1102-1112, 2018 largely to the presence of extractives in the Reserve materials are present in the heartwood, e.g. anthraquinones and wood as lipids and non-structural naphthoquinones (Niamké et al., 2011). In carbohydrates. Storage of non-structural contrast to heartwood, teak sapwood is less carbohydrates (NSC), principally sugars and durable mainly due to the presence of reserves starch (Hoch et al., 2003) is very important for material and its low content of extractives woody species. Indeed, these reserves were (Niamké et al., 2011). This difference of used in long-lived organisms for their natural durability between sapwood and perennity by fighting against biotic and heartwood has been suggested to be strongly abiotic stress, including pests, disturbance, related to the storage of NSC which are and drought (Dietze et al., 2014). NSC play responsible for the biosynthesis of toxic many functional roles such as nutrient secondary substances (Hamada et al., 2017). transport, carbon providing for energy In addition, it is shown that high content of metabolism and osmoregulation, and the sucrose and other sugars in wood promotes its biosynthesis of toxic extractives for defense rapid colonization by fungi (Okahisa et al., mechanisms (Kampe and Magel, 2013), they 2006; Ja’afaru and Onilude, 2014), indicating are highly relevant in the context of tree that sugars status of a tissue could be an resilience to many global change factors. indicator for its susceptibility to pathogens, However, it remains unclear for how these especially to fungi attacks. reserves are stored in the wood. The differences in natural durability Therefore, we are interested by the between sapwood and heartwood are related mobilization of NSC and their possible to heartwood formation. Heartwood formation relationship to natural durability in teak. In is an important process in perennial plants as teak, previous studies have shown that NSC trees. It is the ultimate process leading to the (starch, glucose, fructose and sucrose) and fats death of living sapwood tissues due to internal were highly accumulated in sapwood phenomenon depending on the cycle of tree (Nobuchi et al., 2005). Moreover, Windeissen life involving both water and mechanical et al. (2003) found that hydrolyzed methanol gradients (Berthier et al., 2001; Kokutse et al., extracts of all parts of Panama teak wood 2009). During heartwood formation, several contained mono and disaccharides and sugar widespread reactions (oxidation, hydrolysis, derivatives. They suggested that a part of teak methylation and glycosylation) occurred and extractives occurred in condensed forms or as lead to the improvement of wood properties glycosides with more than two sugar such as natural durability and colour. components. Glycosylation (the conjugation of a sugar In the present work, we attempted to moiety) lead to modifications of chemical better understand the potentialities of structures and is catalysed by a family of perennial plant to mobilize and use their NSC enzymes called glycosyltransferases. It plays in the wood to strengthen their defence system many roles in plant: (1) it contributes to the and improve their natural durability. For that, high chemical diversity among plant the radial distribution of NSC was examined secondary metabolites, (2) it is implied in the before and after chemical hydrolysis of wood control of the compartmentalization of samples. Soluble NSC and starch, and metabolites, stabilization, enhancement of condensed NSC were quantified through the water solubility and analysis of major soluble NSC in both deactivation/detoxification of natural sapwood and heartwood of trees from products, leading to the regulation of Malaysian plantation-growth teak. Their metabolic homeostasis, detoxification of natural durability was already determined xenobiotics and the biosynthesis, storage and (Niamké et al. 2011). The content of NSC was transport properties of secondary metabolites determined by spectrophotometric method (Yonekura-Sakakibara, 2009). after enzymatic assays and the major phenolic compounds were analyzed by HPLC. 1103 B. F. NIAMKE et al. / Int. J. Biol. Chem. Sci. 12(3): 1102-1112, 2018 MATERIALS AND METHODS aqueous) were analyzed by HPLC and the Materials aqueous phase was used for NSC Six trees of teak from Malaysia, quantification as described above. The acid ranging in age from 5.5 to 10 years were used hydrolysis of rutin (quercetin rutinoside) was and sampled as previously described by used as control. Niamké et al. (2010). All the wood slices were conditioned in a climatic room (humidity: Basic hydrolysis 65% ± 5%, temperature 20 °C ± 1 °C) until Acetonic extract (750 µL) was mixed their use. From one radius, four sections of with 750 µL of 2 N sodium hydroxide; wood (25 x 25 x 10 mm, R,T,L) were taken nitrogen was added and the mixture was successively from sapwood to the pith into conserved at 4 °C during 2 h. After sapwood (SW), outer heartwood (OHW), neutralisation until pH 2, the extract was dried middle heartwood (MHW), inner heartwood under nitrogen. The dried extract was (IHW). The wood samples were ground to 0.5 dissolved in an aqueous methanolic solution mm of size of the particles in a Retsch ZM (50%). The hydro-methanolic extract was 200 mill (Oise, France). filtered and the final solution was analyzed for non-structural carbohydrates as described Non-Structural Carbohydrates extraction above. The basic hydrolysis of chlorogenic and quantification acid (cafeoyl-quinic acid) was used as control. Non-structural carbohydrates (NSC- starch, glucose, fructose and sucrose) were Isolation, purification and identification of extracted and measured enzymatically by UV the phenolic compound H1 methods (spectrophotometer Bio-Tek Grounded dry matter (12.8 g) of Instruments, Colmar, France) according to sapwood were extracted with 140 mL of Niamké et al. (2010). Glucose and fructose acetone/water (80:20, v/v) at 4 °C and pH = 2 were determined directly, and sucrose (on the (see extraction
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