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Characterization of Schinopsis Haenkeana Wood Decayed by Phellinus Chaquensis (Basidiomycota, Hymenochaetales)

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Characterization of Schinopsis Haenkeana Wood Decayed by Phellinus Chaquensis (Basidiomycota, Hymenochaetales) IAWA Journal, Vol. 33 (1), 2012: 91–104 CHARACTERIZATION OF SCHINOPSIS HAENKEANA WOOD DECAYED BY PHELLINUS CHAQUENSIS (BASIDIOMYCOTA, HYMENOCHAETALES) María Luján Luna1, 2, *, Mónica A. Murace3, Gerardo L. Robledo4 and Mario C. N. Saparrat5,6,7 SUMMARY Schinopsis haenkeana is a native tree to the Chaco Serrano Forests in Argentina. The white-rot fungus Phellinus chaquensis degrades its wood, causing a white- rot type of decay. The objective of this study was to investigate the structural alterations caused by P. chaquensis in S. hankeana decayed naturally and in vitro. Sound living branches with decay and basidiocarps of P. chaquensis were sampled from the field and in vitro decay tests were performed according to the ASTM D-2017-81 standard method. Naturally decayed branches exhibited an innermost discolored zone with white-rot decay and an outer yellowish-white portion of sound sapwood. Using LM and SEM, degraded tissue displayed diag- nostic characters of selective delignification and simultaneous decay. Findings indicate that P. chaquensis causes a mottled pattern of decay (selective deligni- fication plus simultaneous decay) inS. haenkeana wood. Other features such as accumulation of extractives, profuse deposition of crystals and tyloses, typical of Schinopsis spp. heartwood, were additionally observed. In laboratory degraded material, signs of selective delignification and incipient stages of simultaneous decay were noticeable only microscopically. Chemical analysis revealed an oxi- dative alteration of aromatic moieties in naturally decayed samples which might be related to the accumulation of phenols as a response to fungal attack when compared to sound samples. Naturally degraded sapwood exhibits anatomical and chemical modifications that indicate the development of discolored wood derived from the host-pathogen interaction. Key words: Schinopsis haenkeana, Phellinus chaquensis, anatomy, white-rot de- cay, pathological heartwood, Fourier transform infrared spectroscopy, aromatic moieties. 1) Cátedra Morfología Vegetal, Facultad de Ciencias Naturales y Museo, Universidad Nacional de La Plata, Paseo del Bosque s/n, (1900) La Plata, Argentina. 2) Comisión de Investigaciones Científicas de la Provincia de Buenos Aires (CIC-BA). 3) Cátedra Protección Forestal, Facultad de Ciencias Agrarias y Forestales, UNLP, CC 31, (1900) La Plata, Argentina. 4) Laboratorio de Micología, IMBIV, CONICET, Universidad Nacional Córdoba, CC 495, (5000) Córdoba, Argentina. 5) INFIVE, UNLP- CCT La Plata CONICET, CC 327, (1900) La Plata, Argentina. 6) Instituto Spegazzini (FCNyM-UNLP) 53 # 477, (1900) La Plata, Argentina. 7) Cátedra Microbiología Agrícola, Facultad de Ciencias Agrarias y Forestales, UNLP, 60 y 119, (1900) La Plata, Argentina. *) Corresponding author [E-mail: [email protected]]. Associate Editor: Susan A. Anagnost Downloaded from Brill.com09/26/2021 11:49:19AM via free access 92 IAWA Journal, Vol. 33 (1), 2012 INTRODUCTION Schinopsis haenkeana Engl. (“orco quebracho”, “quebracho del cerro”) is a native tree, belonging to the Anacardiaceae, characteristic of the Chaco Serrano forests from central Argentina and considered vulnerable because of a decline in area of occupancy and the potential levels of exploitation (IUCN 2011). In this country the timbers of Schinopsis spp. are widely utilized because of their hardness and high durability, which are associated with high tannin contents (Roth & Giménez de Bolsón 1997). The most common uses are: fence posts, telegraph poles, bridges, railways, as well as tannin extraction and fire-wood. The white-rot fungus Phellinus chaquensis (Iaconis & J. E. Wright) J. E. Wright & J. R. Deschamps (Basidiomycota, Hymenochaetales) has been reported growing on Caesalpinia paraguariensis (D. Parodi) Burkart and Schinopsis spp. wood in the Chaco Forests (Iaconis & Wright 1953) and is the only heart-rot agent known for Schinopsis haenkeana (Robledo & Urcelay 2009). As a consequence of decay, infected trees are often hollow. Although fungal attack negatively impacts the potential applications of this wood by humans, it is considered an important factor in ecosystem dynamics, because many birds and mammals utilize the cavities formed during decay as a refuge and for nesting (Robledo & Urcelay 2009). Several species of Phellinus are known to cause white heart-rot in hardwoods as well as softwoods around the world (Blanchette 1980; Larsen & Cobb-Poulle 1990; Allen et al. 1996; Callan 1998; Ryvarden 2004). In Argentina, some Phellinus species are reported to have a wide range of substrates, i.e. Phellinus rimosus complex on Cedrela fissilis, Acacia spp., and Prosopis spp., meanwhile other species are only found on a few or one substrate, i.e. P. andinopatagonicus on Nothofagus spp. and Phellinus tabaquilio on Polylepis australis (Wright & Blumenfeld 1984; Deschamps & Wright 2000; Vizcarra Sánchez 2004; Rajchenberg 2006; Robledo & Urcelay 2009). The term white rot has been traditionally used to describe a type of decay where lignin, cellulose and hemicelluloses are degraded and the wood assumes a bleached appearance. Two patterns of white rot are generally recognized: selective delignification and simultaneous decay (Blanchette 1980; Schwarze et al. 2000). In selectively decay- ed woods, lignin and hemicellulose are preferentially removed causing delignification and separation among cells at the middle lamella. In simultaneous decay, all cell wall components are degraded (i.e. lignin, cellulose and hemicelluloses) thus causing the erosion of the cell wall adjacent to hyphae (Blanchette & Reid 1986; Anagnost 1998). Wood degradation may be dependent on the fungal species and its variation among strains, particular substrate characteristics (wood), nutrient availability and enviromental conditions (Blanchette 1991; Perestelo et al. 1999). Since information about the type of decay caused by P. chaquensis in “orco-quebra- cho” wood is very limited (Luna et al. 2007; Robledo & Urcelay 2009), this study was done to investigate the structural alterations caused by P. chaquensis in S. hankeana samples from natural forests and in-vitro decayed wood. Downloaded from Brill.com09/26/2021 11:49:19AM via free access Luján Luna et al. — Wood decay of Schinopsis 93 Figure 1–3. Schinopsis haenkeana attacked by Phellinus chaquensis. – 1: General view of a specimen in the Chaco Serrano Forest. – 2: Detail of trunk with basidiomata. – 3: Detail of basidioma. MATERIALS AND METHODS Living branches (15–20 cm diameter) of Schinopsis haenkeana, both with basidiomata of Phellinus chaquensis and without fungal fructifications (Fig. 1–3), were collected at Punilla Department, Córdoba Province, Argentina (30º 49' S, 64º 32' W). Sound branches were used as controls and to conduct in-vitro decay testing accord- ing to the ASTM D-2017-81 standard method. Laboratory decay tests on dead wood were undertaken in order to distinguish host (living wood) response to fungal attack. Downloaded from Brill.com09/26/2021 11:49:19AM via free access 94 IAWA Journal, Vol. 33 (1), 2012 All the materials (sound, naturally decayed and in-vitro decayed woods) were used for the anatomical and chemical analysis. Fungal isolation, growth and oxidative enzyme activity on agar plates Fungal mycelium was isolated from the context tissues of P. chaquensis basidioma by inoculating on 2% (w v-1) selective malt extract agar (MEA) medium supplemented with chloramphenicol (50 mg l-1) and benomyl (1–3 mg l-1) and adjusted to pH 7. The strain obtained in axenic culture was subcultured on MEA slants as stock cultures, be- ing deposited in the Culture Collection of Instituto Spegazzini, Universidad Nacional de La Plata, Argentina (LPSC 1087). The strain was identified on the basis of its cultural features, such as growth and oxidative enzyme activity on modified Czapek Dox agar (2%, w v-1) basal media with low (1.1 mM) and high (10.9 mM) ammonium tartrate levels, supplemented with guaiacol (1 mM) according to Saparrat et al. (2009). Description of the fungus and its in-vitro cultural features were based on Iaconis & Wright (1953) and Robledo & Urcelay (2009). Anatomical study Materials were analyzed under stereomicroscopy, light microscopy (LM) and scan- ning electron microscopy (SEM). For LM, samples were fixed in formaldehyde-acetic acid-alcohol (FAA), dehydrated through an ethanol series and embedded in paraffin. Sections (8–12 μm thick) were double stained with safranin-fast green (D’Ambroggio de Argüeso 1986). Some sections were left unstained and viewed between crossed polar- izing filters. LM studies were made employing a Nikon Photolab 2 light microscope. For SEM, transverse and longitudinal sections were obtained. Samples were at- tached without pretreatment to aluminum stubs using double sticky tape, air dried and sputter-coated with gold-palladium. Observations were made in a JEOL, JSM-35 CF scanning electron microscope. The anatomical features were described according to Roth and Giménez de Bolsón (1997) and Giménez et al. (2000). The types of decay were identified following the diagnostic characters proposed by Anagnost (1998) and Schwarze (2007). Fourier transform infrared (FT-IR) spectroscopy The chemical composition of sound and decayed samples was analyzed by means of the potassium bromide (KBr) method on a Bruker Fourier-transform IR spectro- photometer. Wood samples were ground to sawdust (< 0.4 mm) in a refrigerated Janke and sieved in a Kunkel mill. Pellets, consisting of 200 mg KBr and c. 3 mg ground wood, were pressed and scanned from
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