Wood-decay type and fungal guild dominance across a North American log transplant experiment

We incubated 196 large-diameter aspen (Populus tremuloides), birch (Betula papyrifera), and pine (Pinus taeda) logs on the FACE Wood Decomposition Experiment encompassing eight climatically-distinct forest sites in the United States. We sampled dead wood from these large-diameter logs after 2 to 6 y of decomposition and determined wood rot type as a continuous variable using the lignin loss/density loss ratio (L/D) and assessed wood-rotting fungal guilds using high-throughput amplicon sequencing (HTAS) of the ITS-2 marker. We found L/D values in line with a white rot dominance in all three tree species, with pine having lower L/D values than aspen and birch. Based on HTAS data, white rot fungi were the most abundant and diverse wood-rotting fungal guild, and soft rot fungi were more abundant and diverse than brown rot fungi in logs with low L/D values. For aspen and birch logs, decay type was related to the wood density at sampling. For the pine logs, decay type was associated with the balance between white and brown/soft rot fungi abundance and OTU richness. Our results demonstrate that decay type is governed by biotic and abiotic factors, which vary by tree species.     

The effects of wood decay type on the growth of bryophyte gametophytes

Deadwood is an important habitat for bryophytes in boreal and subalpine forests. The type of decay in wood (white, brown, and soft rot) caused by fungal colonizers has been revealed to affect bryophyte communities. However, little is known about the effects of decay type on the growth of bryophytes. We tested the effect of wood decay type on gametophyte growth for two common bryophyte species, Scapania bolanderi Austin and Pleurozium schreberi (Brid.) Mitt., which dominate the logs in subalpine coniferous forest on Mt. Ontake, in central Honshu, Japan. We used pot culture experiments in an open-sky nursery field. After eight months of cultivation, the growth of S. bolanderi was larger on brown rot wood than white rot wood, but the growth of P. schreberi was not. Mixed cultures of the two species also showed greater growth on brown rot wood. However, growth of S. bolanderi was significantly smaller than P. schreberi in mixed culture. These results suggest that brown rot wood enhances growth of S. bolanderi, but growth may be reduced under competition from P. schreberi. The results are in agreement with the field observation that brown rot wood has a positive association with S. bolanderi coverage on deadwood.     

Diversity and decay ability of basidiomycetes isolated from lodgepole pines killed by the mountain pine beetle

When lodgepole pines (Pinus contorta Douglas ex Louden var. latifolia Engelm. ex S. Watson) that are killed by the mountain pine beetle (Dendroctonus ponderosae) and its fungal associates are not harvested, fungal decay can affect wood and fibre properties. Ophiostomatoids stain sapwood but do not affect the structural properties of wood. In contrast, white or brown decay basidiomycetes degrade wood. We isolated both staining and decay fungi from 300 lodgepole pine trees killed by mountain pine beetle at green, red, and grey stages at 10 sites across British Columbia. We retained 224 basidiomycete isolates that we classified into 34 species using morphological and physiological characteristics and rDNA large subunit sequences. The number of basidiomycete species varied from 4 to 14 species per site. We assessed the ability of these fungi to degrade both pine sapwood and heartwood using the soil jar decay test. The highest wood mass losses for both sapwood and heartwood were measured for the brown rot species Fomitopsis pinicola and the white rot Metulodontia and Ganoderma species. The sap rot species Trichaptum abietinum was more damaging for sapwood than for heartwood. A number of species caused more than 50% wood mass losses after 12 weeks at room temperature, suggesting that beetle-killed trees can rapidly lose market value due to degradation of wood structural components.     

Substrate affinities of wood decay fungi are foremost structured by wood properties not climate

Wood decomposing fungi differ in their substrate affinities, but to what extent factors like wood properties influence host specialization, compared to climate, is largely unknown. In this study, we analysed British field observations of 61 common wood decay species associated with 41 tree and shrub genera. While white rot fungi ranged from low-to high-substrate affinity, brown rot fungi were exclusively mid-to high-affinity. White rot fungi associated with dead fallen wood demonstrated the least substrate affinity. The composition of wood decomposer fungi was mostly structured by substrate properties, sorted between angiosperms and conifers. Any relationships with temporal and regional climate variability were of far less significance, but did predict community-based and substrate-usage host shifts, especially for fungi on fallen deadwood. Our results demonstrate that substrate shifts by wood-decay fungi will depend primarily upon their degree of affinity to, and the distribution of, related woody genera, followed less at regional levels by climate impacts.     

Carbon and nitrogen acquisition strategies by wood decay fungi influence their isotopic signatures in Picea abies forests

We examined whether sporocarp carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N values) reflected different functional strategies in 15 species of wood decay fungi. In Finnish Picea abies forests, we compared sporocarp δ¹³C and δ¹⁵N against log diameter, proximity to ground, and three wood decay types, specifically brown rot, nonselective white rot, and selective white rot (targeting hemicellulose and lignin preferentially). In regression analysis (adjusted r² = 0.576), species accounted for 31% of variability in δ¹³C, with factors influencing wood δ¹³C accounting for the remainder. Brown rot fungi and three white rot fungi that selectively attacked hemicellulose (Heterobasidion parviporum, Phellopilus nigrolimitatus, and Trichaptum abietinum) were higher in δ¹³C than nonselective white rot fungi. This was attributed to greater assimilation of ¹³C-enriched pentoses from hemicellulose by these fungi. The pathogenic white rot fungus Heterobasidion parviporum had higher δ¹⁵N with proximity to ground and increasing log diameter. This suggested that ¹⁵N-enriched soil N contributed to decomposing logs and that Heterobasidion growing from a bigger resource base had increased access to soil N. These isotopic patterns accordingly reflected both functional diversity of wood decay fungi and site-specific factors.     

Wood decay under the microscope

Many aspects of the interactions between host wood structure and fungal activity can be revealed by high resolution light microscopy, and this technique has provided much of the information discussed here. A wide range of different types of decay can result from permutations of host species, fungal species and conditions within wood. Within this spectrum, three main types are commonly recognised: brown rot, white rot and soft rot. The present review explores parts of the range of variation that each of these encompasses and emphasizes that degradation modes appear to reflect a co-evolutionary adaptation of decay fungi to different wood species or the lignin composition within more primitive and advanced wood cell types. One objective of this review is to provide evidence that the terms brown rot, white rot and soft rot may not be obsolete, but rigid definitions for fungi that are placed into these categories may be less appropriate than thought previously. Detailed knowledge of decomposition processes does not only aid prognosis of decay development in living trees for hazard assessment but also allows the identification of wood decay fungi that can be used for biotechnology processes in the wood industry. In contrast to bacteria or commercial enzymes, hyphae can completely ramify through solid wood. In this review evidence is provided that wood decay fungi can effectively induce permeability changes in gymnospermous heartwood or can be applied to facilitate the identification of tree rings in diffuse porous wood of angiosperms. The specificity of their enzymes and the mild conditions under which degradation proceeds is partly detrimental for trees, but also make wood decay fungi potentially efficient biotechnological tools.     

Temperature effects on hyphal growth of wood-decay basidiomycetes isolated from Pinus densiflora deadwood

Hyphal growth rates were tested on malt extract agar plates at eight different temperatures (5–40?°C) using 36 isolates of 17 basidiomycete species obtained from Pinus densiflora deadwood in Japan. All isolates of four brown rot species showed optimum growth at 30?°C, whereas the optimum growth temperature of white rot species varied from 20?°C to 30?°C. Analysis using a dataset from four cooler sites showed that brown rot fungi grew more rapidly than white rot fungi at higher temperatures (25?°C, 30?°C, and 35?°C). These results suggest that the hyphal growth of brown rot fungi might be physiologically adapted to higher temperatures than those of white rot fungi among the fungal species inhabiting deadwood of P. densiflora in Japan.     

Decay resistance of ash,beech and maple wood modified with N-methylol melamine and a metal complex dye

This study evaluated the decay resistance of ash (Fraxinus excelsior L.), beech (Fagus sylvatica L.), and maple (Acer platanoides L.) wood impregnated by a full cell process with N-methylol melamine (NMM) and combined NMM-metal complex dye (NMM-BS) in aqueous solutions. Basidiomycete decay testing involved incubation with Coniophora puteana (brown rot) and Trametes versicolor (white rot) according to a modified EN 113 (1996) standard, while for the soft rot fungal resistance was evaluated following the standard ENv 807 (2001). NMM and NMM-BS modifications at a WPG range of 7–11% provided decay protection against brown rot resulting in a mass loss less than the required limit (3%). The NMM and NMM-BS modified wood showed increased resistance to white rot decay; however, a higher WPG is needed to prohibit attack from this hardwood specific fungus. The metal-complex dye alone revealed biocidal effects against basidiomycetes. An increased WPG in NMM or NMM-BS had a positive impact against soft rot decay and the lowest mass losses after 32 weeks of exposure were obtained with NMM modification at about 18–21% WPG. NMM modification at this WPG range, however, was not sufficient to protect the wood from soft rot decay. The wood of beech and maple showed slightly higher resistance to all decay types than ash, probably due to the poorer degree of modification of the latter.     

Qualitative and quantitative changes of beech wood degraded by wood-rotting basidiomycetes monitored by Fourier transform infrared spectroscopic methods and multivariate data analysis

Beech wood (Fagus sylvatica L.) veneers were cultivated with white and brown rot fungi for up to 10 weeks. Fungal wood modification was traced with Fourier transform near infrared (FT-NIR) and Fourier transform mid infrared (FT-MIR) methods. Partial least square regression (PLSR) models to predict the total lignin content before and after fungal decay in the range between 17.0% and 26.6% were developed for FT-MIR transmission spectra as well as for FT-NIR reflectance spectra. Weight loss of the decayed samples between 0% and 38.2% could be estimated from the wood surface using individual PLSR models for white rot and brown rot fungi, and from a model including samples subjected to both degradation types.     

Differences in crystalline cellulose modification due to degradation by brown and white rot fungi

Wood-decaying basidiomycetes are some of the most effective bioconverters of lignocellulose in nature, however the way they alter wood crystalline cellulose on a molecular level is still not well understood. To address this, we examined and compared changes in wood undergoing decay by two species of brown rot fungi, Gloeophyllum trabeum and Meruliporia incrassata, and two species of white rot fungi, Irpex lacteus and Pycnoporus sanguineus, using X-ray diffraction (XRD) and ¹³C solid-state nuclear magnetic resonance (NMR) spectroscopy. The overall percent crystallinity in wood undergoing decay by M. incrassata, G. trabeum, and I. lacteus appeared to decrease according to the stage of decay, while in wood decayed by P. sanguineus the crystallinity was found to increase during some stages of degradation. This result is suggested to be potentially due to the different decay strategies employed by these fungi. The average spacing between the 200 cellulose crystal planes was significantly decreased in wood degraded by brown rot, whereas changes observed in wood degraded by the two white rot fungi examined varied according to the selectivity for lignin. The conclusions were supported by a quantitative analysis of the structural components in the wood before and during decay confirming the distinct differences observed for brown and white rot fungi. The results from this study were consistent with differences in degradation methods previously reported among fungal species, specifically more non-enzymatic degradation in brown rot versus more enzymatic degradation in white rot.     

古田山国家级自然保护区木腐真菌物种多样性及分布

木腐真菌是微生物的一个重要类群, 主要以倒木为生长基质, 通过产生各种水解酶将倒木的纤维素、木质素和半纤维素分解为小分子物质, 对促进森林生态系统中的营养物质循环发挥着重要的生态功能。于2016年8月在浙江古田山国家级自然保护区开展的木腐真菌野外调查, 利用形态学和DNA序列分析对采集的标本进行了物种鉴定, 并分析了木腐真菌的物种组成和地理成分。在采集的158份标本中鉴定木腐真菌45属92种, 其中白腐真菌78种, 褐腐真菌14种。古田山的木腐真菌物种区系组成中, 热带-亚热带成分比例最高。在158份木腐真菌标本中, 97份标本采自直径大于10 cm的倒木或树桩上, 分属于76个种, 是木腐真菌生长的主要基质大小类型; 48份标本采自直径为2-10 cm的枝干上, 分属38个种; 13份标本采自直径小于2 cm的枝干上, 分属12种。不同腐烂等级倒木上生长的真菌数量和种类差异明显, 其中一级腐烂倒木上采集到9份标本(7种), 二级腐烂倒木上采集到86份标本(45种), 三级腐烂倒木上49份标本(29种), 四级腐烂倒木上14份标本(14种)。结果表明, 林分中倒木直径大小和腐烂程度是影响木腐真菌生长与分布的重要因子。     

大兴安岭林区多孔菌的区系组成与种群结构

本文在过去10 a野外调查和室内鉴定及分析研究的基础上,对大兴安岭林区多孔菌的区系组成和种群结构进行了分析,发现大兴安岭林区的多孔菌具有较高的多样性,共有5目11科56属129种,占中国多孔菌区系的21.36%,优势科是多孔菌科。种的区系地理成分分为7类,以北温带成分和世界广布成分为主,具有明显的北温带成分的区系特征。大兴安岭的多孔菌常见种较多,种群结构中共生菌3种,寄生菌27种,腐生菌占大多数,有99种。在能够引起木材腐朽的126种真菌中,白腐菌93种,占多数,褐腐菌33种,占少数,但该地区褐腐菌所占比例明显高于全国范围内褐腐菌在多孔菌中的比例。通过对大兴安岭主要树种上的种群结构进行比较,表明阔叶树上的木材腐朽菌绝大部分是白色腐朽菌,而针叶树上的白腐菌与褐腐菌数量相差不大,褐腐菌对于针叶林特别是落叶松的更新具有非常重要的作用。     

Signature Wood Modifications Reveal Decomposer Community History

Correlating plant litter decay rates with initial tissue traits (e.g. C, N contents) is common practice, but in woody litter, predictive relationships are often weak. Variability in predicting wood decomposition is partially due to territorial competition among fungal decomposers that, in turn, have a range of nutritional strategies (rot types) and consequences on residues. Given this biotic influence, researchers are increasingly using culture-independent tools in an attempt to link variability more directly to decomposer groups. Our goal was to complement these tools by using certain wood modifications as ‘signatures’ that provide more functional information about decomposer dominance than density loss. Specifically, we used dilute alkali solubility (DAS; higher for brown rot) and lignin:density loss (L:D; higher for white rot) to infer rot type (binary) and fungal nutritional mode (gradient), respectively. We first determined strength of pattern among 29 fungi of known rot type by correlating DAS and L:D with mass loss in birch and pine. Having shown robust relationships for both techniques above a density loss threshold, we then demonstrated and resolved two issues relevant to species consortia and field trials, 1) spatial patchiness creating gravimetric bias (density bias), and 2) brown rot imprints prior or subsequent to white rot replacement (legacy effects). Finally, we field-tested our methods in a New Zealand Pinus radiata plantation in a paired-plot comparison. Overall, results validate these low-cost techniques that measure the collective histories of decomposer dominance in wood. The L:D measure also showed clear potential in classifying ‘rot type’ along a spectrum rather than as a traditional binary type (brown versus white rot), as it places the nutritional strategies of wood-degrading fungi on a scale (L:D=0-5, in this case). These information-rich measures of consequence can provide insight into their biological causes, strengthening the links between traits, structure, and function during wood decomposition.     

木材腐朽菌在森林生态系统中的功能

木材腐朽菌是森林生态系统的重要组成部分,在森林生态系统中起着极为重要的降解还原作用,主要包括担子菌门非褶菌目、子囊菌门盘菌纲和半知菌类的部分真菌,能全部或部分降解木材中的木质素、纤维素和半纤维素,其降解机制有3种：白色腐朽、褐色腐朽和软腐朽．木材腐朽菌与生态系统中其它生物关系密切,为很多昆虫、鸟类提供营养,有些昆虫也能使木腐菌得到传播．保护木材腐朽菌的生物多样性是保护森林生态系统、维护生态系统健康的重要因素．     

Use of electron microscopy for aiding our understanding of wood biodegradation

Abstract: The use of electron microscopy (EM) has proved to be an invaluable tool for studying structural aspects of lignocellulose degradation by fungi and bacteria and therefore improving our understanding of wood biodegradation. The present review details the application of conventional (SEM, TEM, STEM), analytical (EM X-ray microanalysis, (EDXA)), and immunogold cytochemical EM procedures in the field and gives specific examples of its use for each of the known important types of microbial wood decay including bacterial (tunnelling and erosion), soft rot (cavity and erosion), brown rot and white rot (simultaneous decay and preferential lignin degraders), and discusses important advances made by adopting the techniques. The more recent use of immunogold cytochemistry for studying microbe and fungal enzyme-wood cell wall interactions and its application for localization of specific wood-degrading (laccase, Mn(II) and lignin peroxidases, and cellulases) and H₂O₂ producing (pyranose oxidase) enzymes in situ during white rot decay are also reviewed, as is the application of EM for studying non-enzymatic wood decay. Methods for labelling and detecting wood components (lignin, hemi- and cellulose) in situ by EDXA and enzyme immunogold cytochemistry are also outlined. The use of EM in wood biodegradation research is rapidly expanding and is seen as an important compliment to biochemical and chemical approaches. The future should see even greater advances in our understanding of wood decay as more advanced and recently developed EM techniques are also exploited.     

The soil block test: Potential for improving our understanding of the role of soil source on performance

The soil block test is widely used in North America for evaluating the decay resistance of various wood-based materials. One drawback of this test is that soils from different sources may result in variable wood weight losses. Developing more definitive screening criteria for soils would help select soils that would be most appropriate for use in these procedures. Soils from seven different sources were characterized for soil texture, pH, water holding capacity, C:N ratio, bulk density and then used in soil block tests against two white rot and two brown rot fungi. While there were substantial differences in soil characteristics, none of the parameters were correlated with wood weight loss. The results illustrate the difficulty in predicting fungal behavior in laboratory tests based upon soil characteristics.     

Metabolic activities and ultrastructure imaging at late-stage of wood decomposition in interactive brown rot - white rot fungal combinations

Basidiomycota brown rot fungus (Fomitopsis pinicola) and two white rot fungi (Phlebia radiata, Trichaptum abietinum) were cultivated on thin slices of spruce wood individually and in interspecies combinations. Within 12 months, F. pinicola substantially decomposed spruce wood observed as mass loss, also in three-species combinations. However, white rot fungi through hyphal interactions negatively affected the brown-rot indicative iron reduction capacity of F. pinicola. Decay-signature gene expression in mycelial interaction zones indicated suppression of brown rot mechanism but stimulation of enzymatic white-rot lignin attack by P. radiata. Wood ultrastructure imaging showed white-rot dominance in the fungal combinations, whereas destructive brown-rot was evident with F. pinicola alone. Our results confirm the dynamic pattern of enzyme production in fungal combinations, and transition from brown to white rot decomposition metabolism during the late stage of wood decay after one year of interspecific interactions.     

Progressive changes in cell wall components of Pinus radiata during decay

Progressive changes in solubility characteristics and lignin content of Pinus radiata sapwood were assessed when small blocks were subjected to decay by brown (Gloeophyllum trabeum) and white (Perenniporia tephropora) rot fungi. The brown rot species removed lignin in approximate proportion to weight loss up to 10%; thereafter the amount of lignin altered little. In contrast, the decline in lignin content was near linear for P. tephropora. Increases in solubility (particularly with hot water and dilute alkali), in sugar content and in the acidity of aqueous extracts were recorded in wood blocks decayed to 15–20% weight loss. While these effects were more pronounced in samples decayed by G. trabeum, it appears that with both organisms structural components were degraded faster than the products could be utilised. In this case, cell wall chemistry may not have been a major determinant of weight losses.     

Wood Decomposition of Cyrilla racemiflora (Cyrillaceae) in Puerto Rican Dry and Wet Forests: A 13‐year Case Study1

We studied the decomposition of Cyrilla racemiflora logs over a 13‐yr period in tropical dry and wet forests in Puerto Rico. The mean mass loss, ratio of soft to hard wood, nutrient concentrations, and the diversity of wood‐inhabiting organisms were greater in logs decomposing in the dry forest than in the wet forest. Termites were also more abundant in the logs collected from the tropical dry forest than the tropical wet forest. High moisture content and a low animal diversity on the logs in the wet forest seem to retard wood decay in this habitat. Wood decay rates in the tropical dry forest can be related to the high diversity of species and functional groups of wood‐inhabiting organisms.     

Early soft rot colonization of Scots sapwood pine in above-ground exposure

The early colonization of Scots pine (Pinus sylvestris L.) sapwood exposed above ground (staple bed) was studied. Two different types of exposures were used, one in an open field and the other in a shaded field. Decay type and degree of degradation due to soft rot, and mass and strength loss of wood were correlated. Fungal species in Scots pine sapwood were identified by sequencing, using the fungal nuclear ribosomal DNA (nrDNA) after 24 months.The most abundant decay type found was soft rot, which also agreed with the mass loss (7–8%). Pine sapwood did not differ significantly between the two sites regarding the average mass loss during the time of exposure. The early colonization of wood by soft rot fungi together with mass loss indicates that this fungal type might be more common in above-ground conditions than recognized earlier.