Ectomycorrhizal mats alter forest soil biogeochemistry

Dense hyphal mats formed by ectomycorrhizal (EcM) fungi are prominent features in Douglas-fir forest ecosystems, and have been estimated to cover up to 40% of the soil surface in some forest stands. Two morphotypes of EcM mats have been previously described: rhizomorphic mats, which have thick hyphal rhizomorphs and are found primarily in the organic horizon, and hydrophobic mats, which occur in the mineral horizon and have an ashy appearance. This study surveyed EcM mat and non-mat soils from eight early and late seral conifer forest stands at the H.J. Andrews Experimental Forest in western Oregon. EcM mats were classified by morphology and taxonomic identities were determined by DNA sequencing. A variety of chemical and biochemical properties, including enzymes involved in C, N, and P cycling were measured. Analysis was confined to a comparison of rhizomorphic mats colonizing the organic horizon with non-mat organic soils, and hydrophobic mats with non-mat mineral soils. Both the organic and mineral horizons showed differences between mat and non-mat enzyme profiles when compared on a dry weight basis. In the organic horizon, rhizomorphic mats had greater chitinase activity than non-mat soils; and in the mineral horizon, hydrophobic mats had increased chitinase, phosphatase, and phenoloxidase activity compared to the non-mat soil. The rhizomorphic mats had 2.7 times more oxalate than the non-mats and significantly lower pH. In the mineral horizon, hydrophobic mats had 40 times more oxalate and significantly lower pH than non-mat mineral soils. Microbial biomass C was not significantly different between the rhizomorphic mat and non-mat organic soils. In the mineral horizon, however, the hydrophobic mats had greater microbial biomass C than the non-mat soils. These data demonstrate that soils densely colonized by EcM fungi create a unique soil environment with distinct microbial activities when compared to non-mat forest soils.     

Influence of ectomycorrhizal mat soils on lignin and cellulose degradation

Summary The ectomycorrhizal fungus Hysterangium setchellii (Fisher) forms extensive hyphal mats at the soillitter interface with the roots of the host tree Douglas fir Pseudotsuga menziesii [(Mirb.) Franco]. Microbial biomass, and lignin and cellulose decomposition rates were measured seasonally for 1 year, using ¹⁴C techniques in ectomycorrhizal mat soils and adjacent non-mat soils in a second-growth Douglas fir forest. The microbial biomass and cellulose degradation rates were 3–6 times higher in ectomycorrhizal mat soils than in adjacent nonmat soils. Lignin degradation rates were higher in ectomycorrhizal mat soils than adjacent non-mat soils. Our data suggest that the ectomycorrhizal fungus H. setchellii provides a microenvironment with increased microbiological activity which results in faster lignin and cellulose decomposition.     

Can ectomycorrhizal fungi circumvent the nitrogen mineralization for plant nutrition in temperate forest ecosystems?

Nitrogen (N) limits plant growth in many forest ecosystems. The largest N pool in the plant-soil system is typically organic, contained primarily within the living plants and in the humus and litter layers of the soil. Understanding the pathways by which plants obtain N is a priority for clarifying N cycling processes in forest ecosystems. In this review, the interactions between saprotrophic microorganisms and ectomycorrhizal fungi in N nutrition with a focus on the ability of ectomycorrhizal fungi to circumvent N mineralization for the nutrition of plants in forest ecosystems will be discussed. Traditionally, it is believed that in order for plants to fulfill their N requirements, they primarily utilize ammonium (NH₄⁺) and nitrate (NO₃⁻). In temperate forest ecosystems, many woody plants form ectomycorrhizas which significantly improves phosphorus (P) and N acquisition by plants. Under laboratory conditions, ectomycorrhizal fungi have also been proven to be able to obtain N from organic sources such as protein. It was thus proposed that ectomycorrhizal fungi potentially circumvent the standard N cycle involving N mineralization by saprotrophic microorganisms. However, in many forest ecosystems the majority of the proteins in the forest floor form complexes with polyphenols. Direct access of N by ectomycorrhizal fungi from a polyphenol-protein complex may be limited. Ectomycorrhizal fungi may depend on saprotrophic microorganisms to liberate organic N sources from polyphenol complexes. Thus, interactions between saprotrophic microorganisms and ectomycorrhizal fungi are likely to be essential in the cycling of N within temperate forest ecosystems.     

埃塞俄比亚罗毕高原含铬雏形土地区凋落物饲喂的白蚁(Macrotermes spp)的化学性质及其生化活动的研究

Termite(Macrotermes spp.) mounds are complex biological habitats originated by the termite activity and possessing peculiar physical, chemical and biochemical properties. In this study we examined the concentration of nutrients and the biochemical activity of abandoned soil and mounds colonized by termites of the genera Macrotermes located in the Borana District, Ethiopia. To elucidate the magnitude and persistence of the termite-induced effects, we also studied an abandoned mound, previously colonized by termites of the same genera formed on the same soil. Results confirmed that termite-colonized mounds are ‘hot spots' of nutrient concentration and microbial activity in tropical soils. This is due to the termite driven litter input and decomposition. The abandoned mounds showed higher microbial biomass and activity and displayed a nutrient redistribution and a greater microbial activity than the adjacent soils. These findings allowed us to hypothesize a model of nutrient cycling in colonized soils and a partition of the relative roles of termites and soil microorganisms in nutrient location and turnover in tropical soils. These results may be also useful for the optimal management of termite-colonized soils.     

An ecosystem-scale radiocarbon tracer to test use of litter carbon by ectomycorrhizal fungi

The degree to which ectomycorrhizal fungi rely on decomposing litter as a carbon source in natural ecosystems is unknown. We used a radiocarbon (¹⁴C) tracer to test for uptake of litter carbon by ectomycorrhizal fungi as part of the Enriched Background Isotope Study (EBIS) in Oak Ridge Reservation, Tennessee. In EBIS, leaf litter from a highly ¹⁴C-labeled Quercus alba (white oak) forest was reciprocally transplanted with litter from a nearby low-labeled forest that had not been as strongly exposed to ¹⁴C. These litter transplants were conducted yearly. We measured Δ¹⁴C signatures of ectomycorrhizal fungi collected from each forest four months and 2.25 years after the first litter transplant. The ectomycorrhizas were associated with white oak trees. We found no significant differences in ¹⁴C signatures of ectomycorrhizal fungi exposed to low-labeled versus high-labeled litter, indicating that less than 2% of the carbon in ectomycorrhizal biomass originated from transplanted litter. In contrast, ectomycorrhizal Δ¹⁴C signatures from the high-labeled forest were 117-140‰ higher than those from the low-labeled forest. This pattern suggests that ectomycorrhizal fungi acquired most (or all) of their carbon from their host plants, probably via direct transfer of photosynthate through the roots.     

Functional guild classification predicts the enzymatic role of fungi in litter and soil biogeochemistry

Linking community composition to ecosystem function is a challenge in complex microbial communities. We tested the hypothesis that key biological features of fungi - evolutionary history, functional guild, and abundance of functional genes – can predict the biogeochemical activity of fungal species during decay. We measured the activity of 10 different enzymes produced by 48 model fungal species on leaf litter in laboratory microcosms. Taxa included closely related species with different ecologies (i.e. species in different “functional guilds”) and species with publicly available genomes. Decomposition capabilities differed less among phylogenetic lineages of fungi than among different functional guilds. Activity of carbohydrases and acid phosphatase was significantly higher in litter colonized by saprotrophs compared to ectomycorrhizal species. By contrast, oxidoreductase activities per unit fungal biomass were statistically similar across functional guilds, with white rot fungi having highest polyphenol oxidase activity and ectomycorrhizal fungi having highest peroxidase activity. On the ecosystem level, polyphenol oxidase activity in soil correlated with the abundance of white rot fungi, while soil peroxidase activity correlated with the abundance of ectomycorrhizal fungi in soil. Copy numbers of genes coding for different enzymes explained the activity of some carbohydrases and polyphenol oxidase produced by fungi in culture, but were not significantly better predictors of activity than specific functional guild. Collectively, our data suggest that quantifying the specific functional guilds of fungi in soil, potentially through environmental sequencing approaches, allows us to predict activity of enzymes that drive soil biogeochemical cycles.     

Direct estimates of C:N ratios of ectomycorrhizal mycelia collected from Norway spruce forest soils

Direct estimates of C:N ratios of ectomycorrhizal (EM) mycelia growing in situ in forest soils have been obtained for the first time. The mycelial samples were collected from sand-filled mesh bags that were buried in the soil and incubated for 12-18 months in two Norway spruce forests in southern Sweden. At harvest the mesh bags were heavily colonized and the mycelia were extracted from the sand with water. The collected mycelia had earlier been identified as belonging to EM fungi based on their C isotopic composition. The mean value of the C:N ratio for mycelia was 20.2±0.8 (n=25). EM mycelia collected at different soil depths (5, 15 and 30 cm) had similar C:N ratios. C:N ratios of microbial biomass obtained by fumigation-extraction of similar soils have usually been lower (6-13) so possible differences in the extraction efficiency of C and N from bacteria and fungi are discussed.     

Litter decomposition reduces either N2O or NO production in strongly acidic coniferous and broad-leaved forest soils under anaerobic conditions

Purpose

The beneficial effect to the environment of nitrate (NO₃ ^?) removal by denitrification depends on the partitioning of its end products into nitrous oxide (N₂O), nitric oxide (NO), and dinitrogen (N₂). However, in subtropical China, acidic forest mineral soils are characterized by negligible denitrification capacity and thus reactive forms of N could not be effectively converted to inert N₂, resulting in a negative environmental consequence. In this study, the influences of C input from litter decomposition on denitrification rate and its gaseous products under anoxic conditions in the acidic coniferous and broad-leaved forest soils in subtropical China were investigated using the acetylene (C₂H₂) blockage technique in the laboratory.

Materials and methods

The coniferous and broad-leaved forest soils with and without litter addition were incubated under anaerobic conditions for 244 h. There were three treatments for each forest soil including addition of 0.5 and 1% corresponding litter (gram of litter per gram of soil) and the control without addition of litter.

Results and discussion

The results showed that litter addition into the broad-leaved forest soil had no effect on average rates of denitrification (calculated as the sum of NO, N₂O, and N₂), whereas in the coniferous forest soil, the addition resulted in a significant increase in average denitrification rate. In the broad-leaved forest soil, both rates of litter addition decreased the production of NO but increased the production of N₂, and high rates of litter addition into the coniferous forest soil promoted the reduction of N₂O to N₂.

Conclusions

Increased decomposition of litter in the forest soils could effectively reduce N₂O and NO production through denitrification under anaerobic conditions.     

Soil niche effect on species diversity and catabolic activities in an ectomycorrhizal fungal community

The species of an ectomycorrhizal (ECM) community were investigated in a temperate oak forest by morphotyping and ITS rDNA sequencing. Thirty-six ECM morphotypes were found at the site. The niche effect (as organic soil, mineral soil or dead woody debris artificially introduced in the soil) on the ECM community structure and on the potential catabolic activities of the most abundant morphotypes was studied. The morphotypes in each niche were subjected to enzymatic tests developed for hydrolytic and oxidative enzymes involved in the decomposition of organic compounds. The ECM community structure varied widely depending on the soil horizon or habitat patch. The species richness was higher in the A1 horizon than in the other niches. Different ECM species had different activity patterns for the eight enzymatic tests while co-occurring in the same niche. Catabolic activities also changed within species between niches. Dead woody debris were extensively colonized by two saprotrophic fungi (Megacollybia platyphylla and Armillaria sp.) and, in this particular niche, ECM morphotypes predominantly belonged to the genera Lactarius and Tomentella. These morphotypes showed high chitinase activities. This study suggested also that some ECM fungi could obtain nutrients via the chitin degradation of dead or live saprobes.     

Uptake of an amino acid by ectomycorrhizal fungi in a boreal forest

We assessed the degree to which ectomycorrhizal fungi exploit organic nitrogen in situ. In an Alaskan boreal forest, we identified pairs of sporocarps from five taxa of ectomycorrhizal fungi. We added ¹³C-labeled alanine to the soil surrounding one sporocarp within each pair; the second served as an unlabeled control. Peak rates of ¹³C-respiration from alanine were higher in the labeled sporocarp plots than the controls, indicating that the ¹³C-alanine was detectably respired from the soil. “Reference” plots adjacent to the sporocarps served as an indication of background ¹³C-respiration rates released by the soil community as a whole. Ectomycorrhizal sporocarps displayed higher ¹³C-respiration rates than their reference plots. Thus, the sporocarps and associated mycorrhizal mycelium appeared to contribute significantly to the release of alanine-derived ¹³CO₂, confirming the hypothesis that ectomycorrhizal fungi may access soil amino acid pools under natural conditions.     

Distinctive fungal and bacterial communities are associated with mats formed by ectomycorrhizal fungi

The distinct rhizomorphic mats formed by ectomycorrhizal Piloderma fungi are common features of the organic soil horizons of coniferous forests of the Pacific Northwest. These mats have been found to cover 25-40% of the forest floor in some Douglas-fir stands, and are associated with physical and biochemical properties that distinguish them from the surrounding non-mat soils. In this study, we examined the fungal and bacterial communities associated with Piloderma mat and non-mat soils. Each mat and non-mat area was repeatedly sampled at four times throughout the year. Characterization of the mat activity and community was achieved using a combination of N-acetylglucosaminidase (NAGase) enzyme assays, and molecular analysis of fungal and bacterial communities using T-RFLP profiles, clone libraries, and quantitative PCR. Piloderma mats had consistently greater NAGase activity across all dates, although the magnitude of the difference varied by season. Furthermore, we found distinct fungal and bacterial communities associated with the Piloderma mats, yet the size of the microbial populations differed little between the mat and non-mat soils. Significant temporal variation was seen in the NAGase activity and in the sizes of the fungal and bacterial populations, but the community composition remained stable through time. Our results demonstrate the presence of two distinct microbial communities occupying the forest floor of Douglas-fir stands, whose populations and activities fluctuate seasonally but with little change in composition, which appears to be related to the physiochemical nature of mat and non-mat habitats.     

Interactions between bacteria and ectomycorrhizal fungi

Interactions between eight ectomycorrhizal fungi and eight bacteria were tested on five laboratory media and in the rhizoplane of Pinus radiata. Depression of growth of the fungi by the bacteria in laboratory media was dependent on the medium and bore little relation to effects in the rhizoplane. In the rhizoplane, different bacteria could depress, have no effect or even stimulate growth of mycorrhizal fungi. Competition and antagonism are suggested as mechanisms for depression of the fungi. Some bacteria gave protection against the depressive effects of other bacteria. Considerable differences occurred between ectomycorrhizal fungi in their colonization of the rhizoplane in the absence of bacteria and also in their presence. The common mycorrhizal fungi Rhizopogon luteolus and Thelephora terrestris generally colonized roots well but the strain of Pisolithus tinctorius studied colonized poorly. Direct microscopy showed the percentage cover of the root by microorganisms was usually only 10–20%.It is proposed that interactions of ectomycorrhizal fungi with soil organisms are important in determining the successful introduction and persistence of inoculated ectomycorrhizal fungi. Fungi should be selected for compatibility with a wide range of soil microflora as well as efficiency in plant stimulation.     

The mycoflora of Eucalyptus maculata leaf litter

The mycoflora of the litter of Eucalyptus maculata was studied using three isolation methods. A total of 45 species, representing 22 genera were isolated from the various litter horizons. A comparison between the fungal flora of the litter (L) and that of the underlying mineral soil (A₁) was also made. It was found that 39 per cent of the species were common to both the forest litter and the soil proper. The composition of the litter fungal flora, however, differed from that of the underlying mineralized soil.     

粘土矿物中钾的生物活化: Ⅰ. 外生菌根

A pot experiment was Carried out to in vestigate effect of ectomycorrhizal fungi on eucalyptus growth and K bio-mobilization from soils and clay minerals,In the experiment,sands mixed with soil,KCl-satureated vermiculate and mica,respectively,were used to nurse eucalypus seedings which were nonectomycorrhized or ectomycorrhized by an ectomycorrhizal fungus Pisolithus tinctorius strain XL1(Pt XC1) isolated from a forest soil from Xichang,Sichuan Province,China,and a worldwide well-know ectomycorrhial fungus Pioslithus tinctorius strain 2144(Pt 2144) obtained in Australia.More depletion of HCl-souluble K by mycorrhizas from the soil and minerals than nomycorrhizas suggested that mycorrhizas had a great ability to mobilize K present in the interlayer and feldspar.Mycorrhizal seedlings depressed greatly K digested with HF-HClO4 from substrates after consecutive extractions of soils and minerals by water,ammonium acetate and boiling HCl,while nonmycorrhizal seedlings reduced it little if any,shwing that the mycorrhizal seedlings could mobilize and then utilize the structureal K in mineral lattice,Ectomycorrhizal fungi played a very important role not only in promoting the growth of eucalyptus seedlings but also in mobilizing K in soils and minerals.The infection of Pt XC1 led to a better growth of eucalyptus seedlings and more K accumulation in the seedlings than that of Pt 2144.The large differeces in K accumulation by the seedlings might be due to different abilityes of the two ectomycorrhizal fungi to mobilize K in intelayer and lattice pools in the clay minerals.     

武夷山不同类型森林土壤细菌、丝状真菌优势菌属的初步研究

本文对武夷山不同类型森林土壤优势细菌丝状真菌属的生态分布进行了初步研究。结果表明：不同类型森林土壤或同一种类型森林土壤剖面的不同层次,均有其特殊和优势的异养细菌丝状真菌。大致是“肥沃”的生境即适宜土壤异养微生物繁殖的环境,优势菌属种类多,密度也大。     

Methane oxidation kinetics differ in European beech and Norway spruce soils

Coniferous forest soils often consume less of the greenhouse gas methane (CH₄) than deciduous forest soils. The reasons for this phenomenon have not been resolved. It might be caused by differences in the diffusive flux of CH₄ through the organic layer, pH or different concentrations of potentially inhibitory compounds. Soil samples were investigated from three adjacent European beech ( Fagus sylvatica ) and Norway spruce ( Picea abies ) stands in Germany. Maximal CH₄ oxidation velocities (V_max(app)) and Michaelis Menten constants (K_M(app)), retrieved from intact soil cores at constant CH₄ concentrations, temperature and matric potential, were twice as great in beech as in spruce soils. Also atmospheric CH₄ oxidation rates measured in homogenized soil samples displayed the same trend. Greatest atmospheric CH₄ oxidation rates were detected in the O_a horizon or in the upper 5 cm of the mineral soil. In contrast to the beech soils, the O_a horizon of the spruce soils consumed no CH₄. A differential effect due to divergent diffusive flux through the litter layer was not found. pH and ammonium concentration were similar in samples from both forest soil types. Ethylene accumulation in all soils was negligible under oxic conditions. These collective results suggest that the different atmospheric CH₄ uptake by beech and spruce soils is caused by different CH₄ oxidizing capacities of methanotrophic communities in the O_a horizon and top mineral soil.     

Microbial function in adjacent subtropical forest and agricultural soil

Soil microbial communities and their activities are altered by land use change; however impacts and extent of these alterations are often unclear. We investigated the functional responses of soil microbes in agricultural soil under sugarcane and corresponding native soil under Eucalyptus forest to additions of contrasting plant litter derived from soybean, sugarcane and Eucalyptus in a microcosm system, using a suite of complimentary techniques including enzyme assays and community level physiological profiles (CLPP). Initially agricultural soil had 50% less microbial biomass and lower enzyme activities than forest soil, but significantly higher nitrification rates. In response to litter addition, microbial biomass increased up to 11-fold in agricultural soil, but only 1.8-fold in forest soil, suggesting a prevalence of rapidly proliferating ‘r’ and slower growing ‘K’ strategists in the respective soils. Litter-driven change in microbial biomass and activities were short lived, largely returning to pre-litter addition levels by day 150. Decomposition rates of sugarcane and soybean litter as estimated via CO₂ production were lower in agricultural than in forest soil, but decomposition of more recalcitrant Eucalyptus litter was similar in both soils, contradicting the notion that microbial communities specialise in decomposing litter of the dominant local plant species. Enzyme activities and community level physiological profiles (CLPP) were closely correlated to microbial biomass and overall CO₂ production in the agricultural soil but not the forest soil, suggesting contrasting relationships between microbial population dynamics and activity in the two soils. Activities of enzymes that break down complex biopolymers, such as protease, cellulase and phenol oxidase were similar or higher in the agricultural soil, which suggests that the production of extracellular biopolymer-degrading enzymes was not a factor limiting litter decomposition. Enzyme and CLPP analyses produced contrasting profiles of microbial activity in the two soils; however the combination of both analyses offers additional insights into the changes in microbial function and community dynamics that occur after conversion of forest to agricultural land.     

Can the extent of degradation of soil fungal mycelium during soil incubation be used to estimate ectomycorrhizal biomass in soil?

The extent of degradation of the fungal biomass in forest soil during laboratory incubation was investigated as a measure of ectomycorrhizal (EM) biomass. The method simulates the disappearance of fungal mycelium after root trenching, where the EM fungi, deprived of its energy source (the tree), will start to die off. Incubating a forest humus soil at 25 °C resulted in a decrease in the relative proportion (mol%) of the phospholipid fatty acid 18:2ω6,9 (a fungal marker molecule) within 3-6 months, indicating that fungal biomass was disappearing. Incubation at 5 °C resulted in essentially no change in the amount of 18:2ω6,9. The measurement of ergosterol, another fungal marker molecule, gave similar results. Incubation of different forest soils (pine, spruce and spruce/oak), and assuming that the disappearance of fungal biomass during this period of time was entirely due to EM fungi, resulted in an estimation of EM biomass of between 47 and 84% of the total fungal biomass in these soils. The humus layer had more EM biomass than deeper mineral layers.     

The response of organic matter mineralisation to nutrient and substrate additions in sub-arctic soils

Global warming in the Arctic may alter decomposition rates in Arctic soils and therefore nutrient availability. In addition, changes in the length of the growing season may increase plant productivity and the rate of labile C input below ground. We carried out an experiment in which inorganic nutrients (NH₄NO₃ and NaPO₄) and organic substrates (glucose and glycine) were added to soils sampled from across the mountain birch forest-tundra heath ecotone in northern Sweden (organic and mineral soils from the forest, and organic soil only from the heath). Carbon dioxide production was then monitored continuously over the following 19 days. Neither inorganic N nor P additions substantially affected soil respiration rates when added separately. However, combined N and P additions stimulated microbial activity, with the response being greatest in the birch forest mineral soil (57% increase in CO₂ production compared with 26% in the heath soil and 8% in the birch forest organic soil). Therefore, mineralisation rates in these soils may be stimulated if the overall nutrient availability to microbes increases in response to global change, but N deposition alone is unlikely to enhance decomposition. Adding either, or both, glucose and glycine increased microbial respiration. Isotopic separation indicated that the mineralisation of native soil organic matter (SOM) was stimulated by glucose addition in the heath soil and the forest mineral soil, but not in the forest organic soil. These positive ‘priming’ effects were lost following N addition in forest mineral soil, and following both N and P additions in the heath soil. In order to meet enhanced microbial nutrient demand, increased inputs of labile C from plants could stimulate the mineralisation of SOM, with the soil C stocks in the tundra-heath potentially most vulnerable.     

Physical extent,frequency, and intensity of phosphatase activity varies on soil profiles across a Douglas-fir chronosequence

In forest soils, the availability of phosphate is largely dependent on phosphatase activity. We used soil imprinting to compare in situ activity and fine-scale distribution of phosphatase on soil profiles located across forest chronosequences of four age classes young (5–6 yrs), canopy closure (24–30 yrs), stem exclusion (61–71 yrs), and older (90–103 yrs) of mixed Douglas-fir/paper birch stands regenerated after fire or clearcutting in southern interior British Columbia, Canada. Chromatography paper treated with a mixture of substrate and colorimetric reagent was applied directly to vertical soil surfaces, accessed through root windows. Stands older than 61 years had both the highest level of in situ phosphatase activity and larger, more intense regions of activity. Bray-extractable phosphorus was negatively related to imprintable phosphatase activity. We compared the changes in phosphatase activity with differences in the ectomycorrhizal fungal (EMF) community that had been documented previously in the same stands. Of 84 ectomycorrhizal fungi found on roots in at least two of the stand-age classes, eight taxa were positively correlated and one taxon (Rhizopogon vinicolor/vesiculosus) negatively correlated with high phosphatase activity. The frequency of three taxa appeared to be positively correlated with larger areas of activity on the soil profiles. By using an imprinting approach, this study was able to demonstrate, for the first time, that in situ phosphatase activity and physical attributes of that activity (i.e., number, size, and relative rates of each area of activity) were related to concentrations of soil nutrients and with the frequency of individual ectomycorrhizal fungi.