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Effects of Calcium-Based Materials and Iron Impurities on Wood Degradation by the Brown Rot Fungus <Italic>Serpula Lacryma

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Effects of Calcium-Based Materials and Iron Impurities on Wood Degradation by the Brown Rot Fungus <Italic>Serpula Lacryma See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/249925305 Effects of calcium-based materials and iron impurities on wood degradation by the brown rot fungus Serpula lacrymans Article in Holzforschung · January 2010 DOI: 10.1515/HF.2010.009 CITATIONS READS 15 309 1 author: Jonathan Schilling University of Minnesota Twin Cities 46 PUBLICATIONS 2,415 CITATIONS SEE PROFILE Some of the authors of this publication are also working on these related projects: Genomic comparison of lignocellulose-degrading successions by wood-decaying fungi View project FUNGuild View project All content following this page was uploaded by Jonathan Schilling on 01 April 2015. The user has requested enhancement of the downloaded file. Article in press - uncorrected proof Holzforschung, Vol. 64, pp. xxx-xxx, 2010 • Copyright ᮊ by Walter de Gruyter • Berlin • New York. DOI 10.1515/HF.2010.009 Effects of calcium-based materials and iron impurities on wood degradation by the brown rot fungus Serpula lacrymans Jonathan S. Schilling* frequent and economically important indoor decay fungi in Department of Bioproducts and Biosystems Engineering, temperate climates, particularly in Europe (Schmidt 2000). University of Minnesota, Saint Paul, Its ability to grow long distances between carbon sources MN, USA over non-nutritional materials is distinctive and has been subject to extensive research and review (Thornton and John- *Corresponding author. son 1986; Jennings and Bravery 1991; Henry 2003). In most Department of Bioproducts and Biosystems Engineering, cases, S. lacrymans can penetrate and degrade building mate- 2004 Folwell Avenue, 108 Kaufert Lab, University of rials, such as mortar and plaster (Bech-Andersen 1987; Low Minnesota, Saint Paul, MN 55108, USA Phone: q1-612-624-1761 et al. 2000), but it is unresolved how these associations affect Fax: q1-612-625-6286 the fungus physiologically. E-mail: [email protected] A commonality among many non-woody building mate- rials is a high level of calcium (Ca). Bech-Andersen (1985) suggested first that Ca2q neutralizes oxalic acid and allows Abstract better pH control during wood decay by S. lacrymans. Fol- low-up studies showed electron micrographs of S. lacrymans Calcium-containing materials have been implicated in pro- degrading Ca-containing materials (Bech-Andersen 1987; moting wood degradation by Serpula lacrymans, but mech- Low et al. 2000) and producing Ca-oxalate away from mate- anisms remain unresolved. In this study, S. lacrymans and rial surfaces (Bech-Andersen 1991). Others observed faster Serpula himantioides degraded pine sapwood in agar-block linear growth rates on Ca-containing materials versus on microcosms with one of four treatments: calcium-free, 5 mM wood (Thornton and Johnson 1986). In a controlled labora- agar CaCl , high-purity gypsum (CaSO ), and gypsum 2 4 tory trial, Palfreyman et al. (1996) demonstrated that increas- amended with 1% FeSO . Calcium and iron availability were 4 ing Ca(NO ) levels in minimal nutrient medium hastened limited in minimal nutrient agar. At week 5, pine degradation 3 2 decay of pine sapwood in agar-block microcosms. The abso- was significantly higher for S. lacrymans in iron-amended lute requirement for Ca by S. lacrymans has been written gypsum treatments than other treatments, and the respective into general texts (Bech-Andersen 1991; Ridout 2000), and agar oxalate levels were also higher. Oxalate solubility was a survey written by Krzyzanowski et al. (1999) for building lowest in pure calcium microcosms. Scanning electron practitioners recommended avoiding Ca-containing materi- microscopy showed hyphae in contact with gypsum and pre- als, such as plaster, to avoid this destructive fungus. cipitation of calcium oxalate. At week 15, wood degradation However, there are some weaknesses in these conclusions by S. lacrymans was severe ()60%) in both calcium-free and recommendations. As discussed by Palfreyman et al. and iron-amended treatments, but was significantly less in f (1996), effects on degradation rates could not be conclusive- pure calcium treatments ( 45%). Cation analysis in week 2q - 15 wood revealed higher calcium and iron levels in treat- ly linked to Ca additions versus its counter ion NO3 . ments containing those element additions. Serpula himan- Nitrogen levels are very low in wood, and Watkinson et al. tioides had decayed wood equally among treatments at both (1981) have shown that nitrate, added as Na(NO3)2, signifi- harvests. Results demonstrate that calcium has an inhibiting cantly enhances hyphal growth and promotes cellulolysis in effect – and not a promoting effect as hitherto believed – S. lacrymans. Palfreyman et al. (1996) also found less decay on wood degradation by S. lacrymans. It appears that oxalate than in untreated controls when the Ca source was CaCl2. and iron play a role in stimulating wood degradation by this Steenkjær-Hastrup et al. (2005) found CaCl2 to be somewhat destructive fungus. inhibitory of wood degradation in S. lacrymans decay micro- cosms, and neither CaCl2 nor pure gypsum (CaSO4Ø2H2O) Keywords: copper tolerance; dry rot; Fenton; himantioides; have promoted wood decay by the ‘sister’ taxon S. himan- oxalic acid. tioides or by other wood degrading species (Schilling and Bissonnette 2008). Crystallization with Ca could be inhibi- tory if the soluble oxalate anion plays an active role during Introduction wood degradation by S. lacrymans, including mobilizing iron (Fe) as often theorized (e.g., Varela and Tien 2003). Serpula lacrymans is a destructive brown rot wood-degrad- Iron impurities in Ca-containing building materials might ing fungus (Kauserud et al. 2007). It is one of the most instead be responsible for any enhancement of wood degra- 2010/75 Article in press - uncorrected proof 2 J.S. Schilling dation by S. lacrymans. Iron is theorized to play a central (‘‘CaCl2’’ treatment). The other pure Ca treatment was a gypsum role in the oxidative Fenton-based mechanisms of brown rot disc (11.5 mm diameter) (‘‘CaSO4’’ treatment) added adjacent but fungi (Goodell et al. 1997; Hyde and Wood 1997; Kerem et not touching wood blocks on top of mesh supports. Gypsum discs al. 1998). Like other brown rot fungi, S. lacrymans secretes were formed inside a cylinder of aluminum foil (shaped around a ࠻ ) high levels of oxalic acid during growth (Akamatsu et al. 7 cork borer) resting on wax paper by adding 0.2 g 99% pure CaSO hemihydrate (Sigma) and then adding 200 ml deionized dis- 2004), reduces wood mechanical strength early during the 4 tilled water. After mixing, discs were allowed to solidify overnight, degradation process (Doi and Nishimoto 1986), and produces gently sanded on the edges, and autoclaved prior to adding to micro- Fe-reducing chelators linked to oxidative depolymerization q cosms. The Fe ‘impurity’ discs (‘‘Fe CaSO4’’ treatment) were cre- of cellulose (Shimowaka et al. 2004). Extracellular oxalate ated in the same manner, adding 200 ml of a fresh 1% (w/v) solution 3q coordinates well as a soluble complex with Fe (Henry of FeSO4Ø7H2O (Mallinckrodt Chemicals, Phillipsburg, NJ, USA) 2003), potentially facilitating Fe availability for Fenton and deionized water to CaSO4 powder, matching the sulfate counter chemistry in early stages of the brown rot mechanism (Goo- ion to isolate effects of a high Fe impurity level. A 1% level is dell et al. 1997). Iron has been reported to promote S. lacry- relevant to real-world levels, and was chosen to approximate levels mans wood degradation (Paajanen 1993), and Low et al. in materials tested by Low et al. (2000) that ranged from 0.3% to (2000) showed Fe translocation by S. lacrymans from plaster 4.2% Fe. Microcosms (ns6 per treatment) were inoculated with two 1 cm2 and stone into wood and observed ‘rusty’ hyphae on high- blocks from 2-week-old fungal cultures and incubated in the dark iron materials. at constant temperature (208C). Five replicates were used to com- The hypothesis for this work was that Ca would not pare wood weight loss. The sixth replicate was used for electron enhance S. lacrymans wood decay and that by immobilizing microscopy. Inoculum, blocks, and gypsum discs were arranged in oxalate, Ca might inhibit Fe sequestration and interfere with an ‘X’ configuration, with two adjacent blocks on mesh, a center the brown rot wood degradation mechanism. The aim of this gypsum disc (if applicable) on mesh, and two adjacent sources of work was to isolate effects of Ca and to test contributing inoculum on agar, with no contact among components. Control effects of Fe as an impurity in Ca-containing materials. microcosms with no fungal inoculum (‘‘control’’ treatment) served to monitor any contamination (none was detected), to monitor wood moisture contents, and to serve as ‘time zero’ material. Materials and methods Harvesting and microscopy preparation Fungal strains Out of five of the six replicate treatment microcosms, one of the Serpula lacrymans (Wulfen: Fries) Schroeter strain EMPA 65 two blocks was aseptically removed at week 5, with the final block (ATCC 32750) and Serpula himantioides (Fr: Fr) P. Karst (ATCC harvested at week 15. Blocks were oven-dried for 48 h at 1038C 36335) were used in this study. The latter was originally deposited and weighed, as before. Decay progress was calculated as a percent as S. lacrymans, but has twice been positively identified as S. of the original mass lost during degradation (weight loss). Blocks himantioides by amplification of the internal transcribed spacer were then ground to 20-mesh in a Wiley mill and powder was stored (ITS) region and BLAST searches of sequences in GenBank over dessicant. Non-inoculated control blocks were treated similarly, (Steenkjær-Hastrup et al. 2005; Kauserud et al. 2006). Both isolates except fresh weights were also recorded to allow calculation of w were maintained on 20 ml yeast malt agar (2% agar, ATCC medium moisture content on a standard dry-weight basis (water weight/oven = x No.
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