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Chemistry and Microscopy of Wood Decay by Some Higher Ascomycetes

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Chemistry and Microscopy of Wood Decay by Some Higher Ascomycetes Vol. 43 (1989) No. 1 Chemistry and Microscopy of Wood Decay by Ascomycetes 11 Holzforschung OFFPRINT 43 (1989) 11-18 Chemistry and Microscopy of Wood Decay by Some Higher Ascomycetes By Thomas Nilsson and Geoffrey Daniel Department of Forest Products, The Swedish University of Agricultural Sciences, Box 7008, S-75007 Uppsala, Sweden and T. Kent Kirk and John R. Obst Forest Products Laboratory1), Forest Service, U.S. Department of Agriculture, Madison, Wisconsin 53705-2398, U.S.A. Keywords Summary White-rot Chemical and microscopic features of wood decay by several ascomycetes in axenic culture are described. Soft-rot The tested ascomycetes caused significant weight losses in birch wood. Daldinia concentrica was espe- Lignin biodegradation cially active, causing a weight loss of 62.9% after 2 months. While lignin and carbohydrates were both Syringyl:guaiacyl ratio degraded, carbohydrates were preferentially attacked. As measured by alkaline nitrobenzene oxidation, Daldinia syringylpropane units of the lignin were removed selectively. All of the tested ascomycetes eroded fiber Hypoxylon cells walls beginning from the lumen. Some also caused cavities in the secondary walls, typical of soft-rot Xylaria decay. Pine wood was generally resistant to decay; some of those species which were capable of forming Libertella soft-rot cavities in birch caused significant weight loss in pine. Alstonia scholaris, a tropical hardwood Alstonia scholaris with a guaiacyl-rich lignin, was resistant to degradation. Betula verrucosa Populus tremula Pinus sylvestris Introduction remnants of the middle lamella were seen. The vessels, ray paren- chyma, and vertical parenchyma in lime showed no signs of degra- Ascomycetes and Fungi imperfecti generally cause dation even in advanced decay. In elm the vessels and ray paren- soft-rot decay of wood (Savory 1954; Corbett 1965; chyma in the earlywood but not latewood resisted degradation. Duncan and Eslyn 1966; Nilsson 1973). Xylariaceous From staining experiments the author concluded that removal of ascomycetes from genera such as Daldinia, Hypoxy­ all substance from the fiber walls in both woods was preceded by localized delignification. Later, larger areas of the walls were delig- lon, and Xylaria have often been regarded as white- nified. The decayed wood was lighter in color than the normal rot fungi (Campbell and Wiertelak 1935; Cartwright wood, indicating a bleaching effect. and Findlay 1946; Kirk 1971; Rogers 1979). Past re- Laboratory decay studies with axenic cultures were search on wood decay by the ascomycetes has been later conducted by several investigators. Blaisdell limited; only one report was found on chemical ana- (1939) found significant weight losses in three differ- lyses of wood decayed in axenic culture. The purpose ent hardwoods by Daldinia concentrica, Hypoxylon of the present study was to describe the chemical and sp., and Xylaria sp. Cartwright and Findlay (1946) re- microscopic features of attack of woods by several ported that x. polymorpha caused a weight loss of higher ascomycetes in axenic culture. 14% in beech wood blocks after 4 months. Similarly, The earliest reports were of chemical and microscopic studies of decay of aspen wood to 16% weight loss by X. digitata naturally decayed hardwoods. Thus Campbell and Wiertelak H. rubiginosum (1935) analyzed lime (Tilia vulgaris) wood decayed by Ustulina vul­ and after 3 months was reported by garis Tul. (= Hypoxylon deustum (Hoffm.: Fr.) Grev.). They esti- Rajagopalan (1966). Weight losses from 10 to 26% in mated from chemical analyses that about one-third of the cellulose aspen and red oak woods in 3 months by D. concen­ and one-fourth of the lignin had been lost through fungal attack. trica, H. atropunctatum, H. mediterraneum, H. Wilkins (1936, 1939) gave detailed microscopic descriptions of ad- pruinatum, and H. punctulatum were reported by vanced decay by U. vulgaris in standing trees of lime and elm. At- tack in lime, more pronounced in latewood than in earlywood fi- Merrill, French, and Wood (1964). Weight losses by a bers, led to a thinning of secondary cell walls. The middle lamella white-rot fungus and a brown-rot fungus included for in both lime and elm persisted, but in the most advanced decay only comparison were 50 to 70%. Microscopic examina- tion of aspen blocks decayed by H. pruinatum re- 1) The Laboratory is maintained in cooperation with the University vealed an erosion form of attack in fiber, vessel, and of Wisconsin. ray parenchyma walls; the middle lamella was not af- Holzforschung / Vol. 43 / 1989 / No. 1 1989 Walter de Gruyter · Berlin · New York 12 T. Nilsson, G. Daniel, T.K. Kirk and J.R. Obst Holzforschung fected, and no soft-rot cavities were reported. No Wood species chemical analyses of the laboratory-decayed woods The following wood species were used in the experiments: birch were reported by any of the foregoing authors. In (Betula verrucosa Ehrh.), Scots pine (Pinus sylvestris L.), Euro- other work, however, Kistler and Merrill (1968) pean aspen (Populus tremula L.), and a tropical hardwood Alstonia scholaris (L.) R.Br. The latter has an exceptionally high lignin con- found a lignin loss of 25% at a weight loss of 25% in tent of approximately 30% and, as shown here, a guaiacyl-rich lig- red oak decayed by Strumella coryneoidea. Weight nin. losses up to 40% were also observed. The fungus For the majority of the decay experiments, sapwood blocks of birch caused a gradual thinning of the secondary cell walls. and pine measuring 5 × 15 × 30 mm (5- × 15-mm transverse face) The xylariaceous ascomycetes occur primarily on an- were employed. In one test involving Daldinia concentrica and Tra­ metes versicolor the block size was 10 × 20 × 20 mm (20- × 20-mm giosperm wood (Rogers 1979), and apparently no one transverse face). has studied the effects of these fungi on gymnosperm woods. The group is important in degrading woody Decay materials on the forest floor. The wood blocks were dried at 105°C and weighed. They were then buried, transverse face downwards, in a layer of moist commercial In the present study we conducted both chemical and potting soil in 100-ml Erlemeyer flasks. The flasks were equipped microscopic analyses of birch wood decayed by sev- with cotton plugs, sterilized by autoclaving, cooled, inoculated with eral species of the Xylariaceae. Pine, aspen, and suspensions of mycelium, and incubated for 2 and 4 months at Alstonia woods were included for comparison with 25°C. The initial moisture content of the wood blocks was certain fungi. 30-40%.In a separate experiment with D. concentrica and T. ver­ sicolor on aspen birch, and Alstonia scholaris wood blocks, the in- cubation period was 10 weeks. After incubation, the wood blocks were removed and cleaned of Material and Methods adhering soil and mycelium. They were then dried at 105°C to con- Fungal species stant weight. Weight losses were calculated as percentage of the original weight. The studied species and their origins are given in Table 1. Table 1. Species and strains of fungi studied. and their origins Species Strain Origin Ascomycetes Daldinia concentrica (Bolt.: Fr.) Ces. and De Not C62 Forintek1) D. eschscholzii (Ehrenb.) Rehm Candoussau J.D.R.2) D. occidentalis Child 9597 Petrini3) Hypoxylon confluens (Tode: Fr.) Westend. CBS335.70 CBS4) H. deustum (Hoffm.: Fr.) Grev. CBS 723.69 CBS (Synonym: Ustulina vulgaris Tul.) H. fuscum (Pers.: Fr.) Fr. C144 Forintek H. mammatum (Wahlenb.) J.H. Miller 324A, 238A5) Forintek H. multiforme (Fr.) Fr. CBS 856.72 CBS H. nummularium Bull.: Fr. CBS 969.70 CBS H. rubiginosum Pers.: Fr. fide Miller C194, 328A6) Forintek Libertella betulina Desm. [anamorph of BH SLU7) Diatrype stigma (Hoffm.: Fr.) Fr.] Xylaria acuta Peck Cooke J.D.R. X. arbuscula Sacc. Samuels J.D.R. X. carpophila (Pers.: Fr.) Fr. 9612 Petrini X. feejeensis (Berk.) Fr. 9613 Petrini X. hypoxylon (L.: Fr.) Grev. 74426 SLU X. longipes Nits. CBS 148.73 CBS X. polymorpha (Pers.: Fr.) Grev. Proffer J.D.R. Basidiomycete Trametes versicolor (Fr.) Pilat (= Coriolus versicolor (Fr.) Quel.) 1) Obtained from Keith A. Seifert, Forintek Canada Corp., Ottawa, Canada. 2) Obtained from Jack D. Rogers, Department of Plant Pathology, Washington State University, Pullman, Washington, U.S.A. 3) Obtained from Liliane Petrini, Benglen, Switzerland. 4) Obtained from Centraalbureau voor Schimmelcultures, Baarn, The Netherlands. 5) Received as Hypoxylon pruinatum. 6) Received as Hypoxylon perforatum. 7) Obtained from the culture collection at Department of Forest Products, Swedish University of Agricultural Sciences, Uppsala, Sweden Vol. 43 (1989) No. 1 Chemistry and Microscopy of Wood Decay by Ascomycetes 13 For microscopy, another type of decay procedure was used (Nilsson Table 2. Average *) percentage weight loss of birch and pine wood 1973). Sterilized birch wood blocks (5 × 5 × 10 mm) were placed decayed for 2 and 4 months by 10 species of ascomycetes on mycelia growing on malt extract or cellulose/mineral salts medium on agar slopes. Test fungus Average weight loss, % In a separate experiment, thin transverse sections (approx. 10 µm) of pine sapwood were exposed directly to the mycelium of D. con­ 2 months 4 months centrica growing on malt agar plates. Birch Pine Birch Pine Transmission electron microscopy (T.E.M.) Daldinia concentrica 62.9 2.5 76.6 2.4 Small pieces of birch wood from blocks degraded by either D. con­ Hypoxylon fuscum 20.1 2.4 40.3 2.3 centrica X. polymorpha or were fixed at room temperature in 3% H. mammatum (324A) 20.4 2.6 39.9 4.7 v/v glutaraldehyde and 2% paraformaldehyde in 0.1M sodium H. mammatum (238A) 23.7 2.2 45.7 1.4 cacodylate buffer (pH 7.2). After washing in buffer (3 × 30 min) H. multiforme 22.7 1.4 49.0 2.5 the samples were post-fixed in 1% w/v osmium tetroxide (3 hr, H.
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