Performance of Bark Beetle Damaged Norway Spruce Wood Against Water and Fungal Decay
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PEER-REVIEWED ARTICLE bioresources.com Performance of Bark Beetle Damaged Norway Spruce Wood Against Water and Fungal Decay Davor Kržišnik, Boštjan Lesar, Nejc Thaler, and Miha Humar * Norway spruce is one of the most important wood species in Central Europe. Unfortunately, bark beetles have prominently attacked spruce trees. Bark beetles colonize wood in symbiosis with ophiostomatoid fungi, which is visible in prominent blue staining. This reduces the commercial value of the infested wood. The relevant properties of blue stained wood were therefore determined: bending and compression strength, sorption properties, DVS analysis, water uptake, and durability against wood decay fungi. This information was applied in the Meyer-Veltrup model for assessment of material resistance. Scanning electron microscopy analysis confirmed severe infestation of blue stained wood, which was also evident from the color of the specimens. The mechanical properties were almost unaffected, as were the sorption properties. However, the durability and water exclusion efficacy of blue stained wood were considerably decreased, which indicates that decay can be expected to appear faster on blue stained wood than on control-untreated wood specimens. Keywords: Bark beetle; Blue stain; Durability; Norway spruce; Sap stain; Water performance; Wood decay fungi Contact information: University of Ljubljana, Biotechnical Faculty, Jamnikarjeva 101, 1000 Ljubljana, Slovenia; *Corresponding author: [email protected] INTRODUCTION Norway spruce (Picea abies) is one of the most important wood species in Central Europe. The growing of spruce wood has been promoted even in regions in which this tree species is not naturally present. Spruce in these locations is fairly susceptible to various abiotic and biotic factors (Repe et al. 2013). The health of the forest thus decreases and the percentage of sanitary felling increases. Damage caused by insects, mostly bark beetles (Coleoptera, Scolytinae), is the most important reason for sanitary felling of Norway spruce in Central Europe, followed by pathogenic fungi and windthrow (Repe et al. 2013). The most devastating effects of bark beetles on spruce trees in Central Europe are associated with Ips typographus, Pityogenes chalcographus, Ips amitinus, and Polygraphus poligraphus (Kirisits 2001; Repe et al. 2015). Ips typographus is predominately aggressive towards Norway spruce. This beetle develops on weakened or freshly harvested trees. The damage caused by bark beetles has even increased due to global warming in the last decade (Repe et al. 2013). Bark beetles are well-known vectors of ophiostomatoid fungi. These fungi are introduced into a new host tree during bark-beetle attacks and the construction of their egg galleries (Harrington 2005; Jankowiak et al. 2009). Bark beetles carry the fungal spores on specialized structures called mycangium, or freely on their bodies. The spores can also be eaten and passed through the digestive tract (Paine et al. 1997). Wood colonization by ophiostomatoid fungi can lead to blue to grey discoloration of the sapwood, which is Kržišnik et al. (2018). “Blue stained wood,” BioResources 13(2), 3473-3486. 3473 PEER-REVIEWED ARTICLE bioresources.com frequently referred to as blue stain or sap stain (Humar et al. 2008; Ratnasingam et al. 2016). Discoloration is a result of the micro-organismal pigment, melanin (Zink and Fengel 1989; Hernandez et al. 2016). Blue stain is a blue, grey, or black striped discoloration on wood. Fungi causing blue stain consume nutrients in the parenchyma cells of sapwood. However, in certain cases, enzymes associated with degradation have been determined; for instance, mannanase, pectinase, and amylase have been detected (Schirp et al. 2003). Although blue stain fungi belong to the same group of fungi as soft rot fungi (Troya et al. 1990), it is generally assumed that blue stain fungi do not cause any minor cell wall attack, so strength properties are hardly affected (Schmid 2006; Humar et al. 2008). It is presumed that fungal hyphae grow on the internal face of cell walls, without any enzymatical alteration on the surface (Liese 1964). However, some hyphae of blue stain fungi have also been seen within parenchyma cell walls (Liese 1964), or even between cell walls in the middle lamella region, which can affect the mechanical properties. Furthermore, blue stain fungi produce intra- and extracellular enzymes, some of which can degrade polysaccharides and pectins. Enzymes that degrade lignin are also present (Troya et al. 1990; Sharpe and Dickinson 1992). Ophiostomatoid fungi, as well as other blue stain and sap stain fungi, affect wood permeability. These fungi are thus frequently used for bio-incising, a biotechnological method designed to enhance the permeability of refractory wood species by incubation with wood-inhabiting fungi or bacteria (Lehringer et al. 2010; Thaler et al. 2012). The most frequently used organism for a variety of biotechnological processes in the wood processing industry is called Cartapip 97. It is an albino strain of Ophiostoma piliferum (Mai et al. 2004). An increase in wood permeability is presumably induced by the selective degradation of pit membranes in bordered and half-bordered pits, entailing only negligible changes in the tracheid cell wall. The influence of micro-organisms on the permeability of wood has been known for some decades (Mai et al. 2004). The main aim of this study was to elucidate how decreased moisture performance influences the service life of blue stained wood in aboveground applications. Water performance can be described as an ability of the wood to restrain water uptake during rainfall events and to release the water as fast as possible in order to keep moisture content below threshold required for fungal decay. The material resistance of wood in an aboveground application is a function of inherent durability (as a consequence of the presence of biocides and/or biologically active extractives) and water performance (Meyer-Veltrup et al. 2017). The inherent durability of spruce is low, so good water performance efficacy considerably influences the material resistance of wood outdoors. Because the water performance of blue stained wood is presumably considerably reduced, it is of great commercial importance in forecasting the service life of blue stained wood. Material resistance is a key piece of data for the majority of the service life prediction models. However, information on material resistance is not sufficient to estimate service life, as service life in addition to material resistance depends on micro and macro climate, as well as conditions of use. EXPERIMENTAL In the frame of the present study, basic properties of blue stained and control wood were determined. These properties included mechanical properties (bending and compression strength), color, sorption properties, water uptake, and durability against Kržišnik et al. (2018). “Blue stained wood,” BioResources 13(2), 3473-3486. 3474 PEER-REVIEWED ARTICLE bioresources.com wood decay fungi. Some of these data were applied in a material resistance model (Meyer- Veltrup et al. 2017). This approach enables comprehensive assessment of quality of blue stained wood. Materials The research was performed on Norway spruce (Picea abies) wood. The wood originated from central Slovenia and had grown between 500 and 700 m above sea level. The diameter of the trees was between 35 and 45 cm. There were two types of wood used; control samples were made of undamaged spruce, without visible signs of decay or blue staining. The second set of specimens was made of wood from spruce trees that had been felled due to attack by European spruce bark beetle (Ips typographus) and were severely infested with Ophiostomatoid fungi, which causes blue staining. This material in the present study is therefore identified as blue stained wood. The density of the samples and ring width of the two sets of specimens were comparable. Color of Wood The color of the specimens was determined from scanned specimens and expressed in the CIE L*a*b* system (Humar et al. 2008). The L*-axis represents the lightness, while, a* and b* are the chromaticity coordinates. In the CIE L*a*b* coordinates, +a* stands for red, -a* for green, +b* for yellow, -b* for blue, and L* varies from 100 (white) to zero (black). Any color can thus be characterized (Brock et al. 2000). Mechanical Properties Static three-point bending tests were performed according to the EN 310 standard (CEN 1993) on a Zwick-Roell Z005 universal testing machine (Zwick-Roell, Ulm, Germany) to obtain the modulus of elasticity (MoE - bending) and bending strength (Fm – bending strength). In total, 20 test specimens (10 replicates for each specimen type) with dimensions 100 mm × 20 mm × 5 mm were prepared and conditioned in a standard climate (relative humidity = 65 ± 5%; temperature = 20 ± 2 °C) until a constant mass of the specimens was achieved. The specimens were tested for MoE and Fm immediately after incubation in the standard climate. Compressive strength (Fm - compression) was determined according to the ASTM D1037-12 (2012) standard on a Zwick-Roell Z100 universal testing machine (Zwick-Roell, Ulm, Germany). In total, 20 test specimens (10 for each specimen type) with dimensions 50 mm × 20 mm × 20 mm were prepared and incubated in a standard climate until a constant mass of the specimens was achieved. The specimens were tested for compressive strength immediately after incubation in the standard climate. After the test, the compressive strength (Fm) was calculated. Specimens used for mechanical properties were also used for determination of density. Density was determined after conditioning in a standard climate (RH = 65 ± 5 %; T = 20 ± 2 °C). In order to calculate density, all three dimensions of specimens were measured, and their mass was then determined. Density was determined on all 40 specimens. SEM Analysis Scanning electron microscopy (SEM) was performed to confirm the presence of fungal hyphae in the wood. Specimens were prepared, ensuring that they were oriented in all three anatomical planes, cutting the surface of each anatomical plane with a sliding Kržišnik et al.