Stable isotope signatures confirm carbon and nitrogen gain through ectomycorrhizas in the ghost orchid Epipogium aphyllum Swartz

Epipogium aphyllum is a rare Eurasian achlorophyllous forest orchid known to associate with fungi that form ectomycorrhizas, while closely related orchids of warm humid climates depend on wood- or litter-decomposer fungi. We conducted (13) C and (15) N stable isotope natural abundance analyses to identify the organic nutrient source of E. aphyllum from Central Norway. These data for orchid shoot tissues, in comparison to accompanying autotrophic plants, document C and N flow from ectomycorrhizal fungi to the orchid. DNA data from fungal pelotons in the orchid root cortex confirm the presence of Inocybe and Hebeloma, which are both fungi that form ectomycorrhizas. The enrichment factors for (13) C and (15) N of E. aphyllum are used to calculate a new overall average enrichment factor for mycoheterotrophic plants living in association with ectomycorrhizal fungi (ε(13) C ± 1 SD of 7.2 ± 1.6 ‰ and ε(15) N ± 1 SD of 12.8 ± 3.9 ‰). These can be used to estimate the fungal contribution to organic nutrient uptake by partially mycoheterotrophic plants where fully mycoheterotrophic plants are lacking. N concentrations in orchid tissue were unusually high and significantly higher than in accompanying autotrophic leaf samples. This may be caused by N gain of E. aphyllum from obligate ectomycorrhizal fungi. We show that E. aphyllum is an epiparasitic mycoheterotrophic orchid that depends on ectomycorrhizal Inocybe and Hebeloma to obtain C and N through a tripartite system linking mycoheterotrophic plants through fungi with forest trees.     

Dramatic changes in ectomycorrhizal community composition, root tip abundance and mycelial production along a stand-scale nitrogen deposition gradient

? Nitrogen (N) availability is known to influence ectomycorrhizal fungal components, such as fungal community composition, biomass of root tips and production of mycelia, but effects have never been demonstrated within the same forest. ? We measured concurrently the abundance of ectomycorrhizal root tips and the production of external mycelia, and explored the changes in the ectomycorrhizal community composition, across a stand-scale N deposition gradient (from 27 to 43 kg N ha?1 yr?1) at the edge of a spruce forest. The N status was affected along the gradient as shown by a range of N availability indices. ? Ectomycorrhizal root tip abundance and mycelial production decreased five and 10-fold, respectively, with increasing N deposition. In addition, the ectomycorrhizal fungal community changed and the species richness decreased. The changes were correlated with the measured indices of N status, in particular N deposition and N leaching. ? The relationship between the altered ectomycorrhizal community, root tip abundance and mycelial production is discussed in the context of the N parameters. We suggest that increased N deposition to forests will cause large changes in ectomycorrhizal fungal community structure and functioning, which, in turn, may result in reduced N uptake by roots and fungi, and increased losses of N by leaching.     

Solubilisation and colonisation of wood ash by ectomycorrhizal fungi isolated from a wood ash fertilised spruce forest

In Sweden application of granulated wood ash has been suggested as a method to supplement nutrient loss resulting from harvesting of forest residues for bioenergy production. Mycelia of two ectomycorrhizal fungi Piloderma sp. 1 and Ha-96-3, were commonly found to colonise ash granules in a wood ash fertilised spruce forest. Thirty-eight fungal isolates were selected from 10 taxa to investigate the possible role of different ectomycorrhizal fungi in nutrient mobilisation from ash. The taxa were Cenococcum geophilum Fr., Piloderma croceum Erikss. and Hjortst., Piloderma sp. 1, Thelephora terrestris (Ehrenb.) Fr., Tylospora fibrillosa Donk, and five unidentified species, all originating from a wood ash fertilised spruce forest. The isolates were tested for their ability to solubilise tricalcium phosphate (TCP) or hardened wood ash (HWA) in vitro. Ha-96-3, P. croceum and Piloderma sp. 1 were the only taxa which solubilised TCP. Abundant calcium oxalate crystals were formed in TCP and HWA plates with Piloderma sp. 1. Ha-96-3 and two isolates of P. croceum produced intermediate amounts of crystals. Ha-96-1 and T. fibrillosa produced low amounts of crystal but no crystal formation was observed by any of the other isolates. Piloderma sp. 1 from HWA plates had significantly higher concentrations of P, compared to P. croceum or Ha-96-3. Piloderma sp. 1 and P. croceum were further tested for their ability to colonise wood ash in microcosms containing intact mycorrhizal associations. After 7 months Piloderma sp. 1 colonised ash amended patches with a dense, mat like mycelium, whereas P. croceum mycelia avoided the ash patches. Possible differences between these fungi in patterns of carbon allocation were investigated by labelling seedlings with 14CO(2). Piloderma sp. 1 mycelia allocated significantly more 14C to ash patches than P. croceum. P. croceum allocated relatively more 14C to control patches than to the ash patches. The possible role of ectomycorrhizal fungi in mobilisation of nutrients from wood ash is discussed.     

Ectomycorrhizal weathering of the soil minerals muscovite and hornblende

Ectomycorrhizal fungi are hypothesized to enhance mineral weathering in forest soils. Several studies have shown an increased uptake of mineral-derived nutrients by trees when in symbiosis with ectomycorrhizal fungi. However, it is difficult to determine from these studies if the improved nutrient uptake is the result of increased weathering or better exploitation of the substrate by the ectomycorrhizal fungi. In a pot experiment, Pinus sylvestris (Scots pine) seedlings were grown with or without ectomycorrhizal fungi, and with or without the mineral muscovite as the only potassium (K) source or the mineral hornblende as the only magnesium (Mg) source. After 27 wk, all pools of non-mineral-bound K or Mg were determined. The ectomycorrhizal fungus Paxillus involutus increased weathering of muscovite but not hornblende. The other ectomycorrhizal fungi tested, Piloderma croceum and Suillus bovinus, did not increase weathering of either muscovite or hornblende compared with the nonmycorrhizal trees. The P. involutus-mediated mobilization of K from muscovite resulted in increased K content of root plus adhering hyphae, but not of shoots. In conclusion, ectomycorrhizal fungi may increase weathering of minerals in response to nutrient deficiencies, but this response is species specific.     

氮沉降对外生菌根真菌的影响

综述了国外氮沉降对外生菌根真菌的影响研究现状 ,主要从菌根形成、形态 (菌丝体、菌根 )变化、子实体生产力和外生菌根真菌群落结构等方面对氮沉降的响应进行了综述 ,并初步探讨了氮饱和的临界负荷。研究表明 ,过量氮沉降会给外生菌根真菌在以下几个方面带来负影响 :(1)影响外生菌根真菌与寄主植物之间的养分分配和循环 ;(2 )降低子实体生产力 ;(3)减少菌丝 ;(4 )降低菌根量及其活力 ;(5 )降低外生菌根真菌丰富度 ;(6 )改变外生菌根真菌群落结构组成 ;(7)降低外生菌根真菌群落功能。还指出了未来该方面研究重点和方向     

Organic anion exudation by ectomycorrhizal fungi and Pinus sylvestris in response to nutrient deficiencies

Low molecular weight organic anions (LMWOA) can enhance weathering of mineral grains. We tested the hypothesis that ectomycorrhizal (EcM) fungi and tree seedlings increase their exudation of LMWOA when supply of magnesium, potassium and phosphorus is low to enhance the mobilization of Mg, K and P from mineral grains. Ectomycorrhizal fungi and Pinus sylvestris seedlings were cultured in symbiosis and in isolation on glass beads with nutrient solution or with sand as a rooting medium, with a complete nutrient supply or with Mg, K, P or N in low supply. Concentrations of all dicarboxylic LMWOA in the rooting medium were measured. Nonmycorrhizal seedlings released predominantly malonate. Colonization with Hebeloma longicaudum decreased the amount of organic anions exuded, whereas Paxillus involutus and Piloderma croceum increased the concentration of oxalate but not the total amount of LMWOA. Phosphorus deficiency increased the concentration of LMWOA by nonmycorrhizal and EcM seedlings. Magnesium deficiency increased the concentration of oxalate by nonmycorrhizal and EcM seedlings, but not the concentration of total LMWOA. Paxillus involutus grown in pure culture responded differently to low nutrient supply compared with symbiotic growth. Ectomycorrhizal fungi did not increase the total concentration of LMWOA compared with nonmycorrhizal seedlings but, depending on the fungal species, they affected the type of LMWOA found.     

钾与信号抑制剂对外生菌根真菌分泌乙酸的调控作用

外生菌根真菌是森林生态系统中的重要成分,参与树木养分的吸收利用。试验液体培养外生菌根真菌,设置不同供钾水平,添加钙信号抑制剂,研究了它们的有机酸和氢离子分泌,以及乙酸分泌对供钾和信号抑制剂的响应。结果表明,供试菌株的生长,氮、磷、钾含量和吸收量因菌株不同而异,生物量变化于52.91—121.72 mg/瓶之间,相差1倍以上。外生菌根真菌吸收养分的差异可能与它们对土壤养分环境的长期适应、进化、选择有关。在外生菌根真菌的培养液中,分别检测出草酸、乙酸、苹果酸、柠檬酸和丁二酸等,以及大量的氢离子,说明菌根真菌能分解土壤含钾矿物,释放钾离子,改善寄主植物的钾营养。其中,乙酸分泌量较大,具有普遍性,低钾刺激分泌乙酸,高钾时分泌减少,其分泌速率与供钾浓度和菌丝吸钾量之间呈显著负相关(r=-0.734,r=-0.617,n=60)。钾对菌根真菌分泌乙酸的调控作用具有改善森林钾素营养,防止土壤养分淋失的生理和生态意义。此外,在低钾条件下,阴离子通道和钙信号抑制剂抑制外生菌根真菌分泌乙酸。说明钙信号和阴离子通道参与了乙酸分泌,缺钾可能是刺激乙酸分泌的原初信号,通过信号转导和一系列级联反应促进乙酸分泌。     

Long-term dynamics of aboveground fungal communities in a subalpine Norway spruce forest under elevated nitrogen input

As anthropogenic N deposition has been suspected to be the main reason for the decline of macromycetous sporocarp production in forest ecosystems, various N-fertilization experiments were started in the mid 1990s. The dynamics of ectomycorrhizal (root-inhabiting) and terricolous saprobic (litter-inhabiting) fungal communities were studied by exhaustive sporocarp inventories in a substitution Norway spruce (Picea abies) forest in two 256-m² plots sampled for periods of 1 week at 1-m² resolution between 1994 and 2007. N was added to the soil twice per year in one plot from the fourth year onwards. The effects of N input and time on aboveground fungal communities were assessed using redundancy analysis, principal response curves and non-parametric multivariate ANOVA. Results of this long-term experiment revealed that both ectomycorrhizal and saprobic fungal communities responded to an increase in soil N input. The ectomycorrhizal community reacted by a fast decrease in sporocarp production and in species richness, whereas the saprobic community was less affected. The response was highly species specific, especially for the saprobic community. The difference in species composition between control and fertilized plots was significant after 1 year of N addition for ectomycorrhizal fungi and only after 3 years for saprobic fungi. An aging effect affected sporocarp production in the whole area. For both communities, this unidirectional drift in species composition was as important as the treatment effect. This result highlights the importance of considering the respective role of treatment and year effects in long-term field experiments on fungal communities.     

The effect of pH on the growth of saprotrophic and ectomycorrhizal ammonia fungi in vitro

Some saprotrophic and ectomycorrhizal fungi produce reproductive structures, preferably in slightly alkaline to neutral forest soil. This research examines the growth of these "ammonia fungi" in liquid medium at various pH values. In the first experiment, the capacity of six buffers was examined to select appropriate buffers for stabilizing pH in the neutral-to-alkaline range by culture of three species of the ammonia fungi in media initially adjusted to pH 7, 8 or 9. The highest buffering capacity was shown in 2-(N-morpholino) ethanesulfonic acid (MES) at pH 7, and N, N-bis (2-hydroxyethyl) glycine (Bicine) and N-2-hydroxyethylpiperazine-N'-2-ethanesulfonic acid (HEPES) at pH 8 and 9. In the second experiment, the growth of 15 strains of both saprotrophic and ectomycorrhizal ammonia fungi was tested on the medium initially adjusted to pH 3, 4, 5, 6 or 7 with MES, or to pH 8 or 9 buffered with Bicine. Many of the saprotrophic species grew well at pH 7 or 8; the ectomycorrhizal species showed optimum growth at pH 5 or 6. The pH suitable for the in vitro growth of these fungi was correlated with the pH of forest soil where these fungi occur.     

Measuring external mycelia production of ectomycorrhizal fungi in the field: the soil matrix matters

Assessing mycorrhizal fungi production in field settings has been hindered by the inability to measure external mycelia. Recently, external mycelia production was measured in the field using a novel in-growth core technique with acid-washed sand as the in-growth matrix. Here, we tested the assumption that external mycelia production in acid-washed sand is representative of that in native soil. External mycelia production was estimated as the difference in fungal growth between closed (allowing only saprotrophic fungal production) and open (allowing mycorrhizal and saprotrophic fungal production) cores using a factorial design of soil matrices (acid-washed sand vs native) and fertilization treatments (control vs nitrogen (N)) in a longleaf pine (Pinus palustris) plantation. In native soils, the ectomycorrhizal to saprotrophic fungal biomass signal was strong and consistent facilitating the assessment of external mycelia production, which was 300% higher than corresponding rates in acid-washed sand and inversely correlated with soil N. These results demonstrate the efficacy and importance of using native soil as the in-growth matrix to measure ectomycorrhizal fungi external mycelia production in field settings.     

Ectomycorrhizal Fungal Protein Degradation Ability Predicted by Soil Organic Nitrogen Availability

In temperate and boreal forest ecosystems, nitrogen (N) limitation of tree metabolism is alleviated by ectomycorrhizal (ECM) fungi. As forest soils age, the primary source of N in soil switches from inorganic (NH₄⁺ and NO₃⁻) to organic (mostly proteins). It has been hypothesized that ECM fungi adapt to the most common N source in their environment, which implies that fungi growing in older forests would have greater protein degradation abilities. Moreover, recent results for a model ECM fungal species suggest that organic N uptake requires a glucose supply. To test the generality of these hypotheses, we screened 55 strains of 13 Suillus species with different ecological preferences for their in vitro protein degradation abilities. Suillus species preferentially occurring in mature forests, where soil contains more organic matter, had significantly higher protease activity than those from young forests with low-organic-matter soils or species indifferent to forest age. Within species, the protease activities of ecotypes from soils with high or low soil organic N content did not differ significantly, suggesting resource partitioning between mineral and organic soil layers. The secreted protease mixtures were strongly dominated by aspartic peptidases. Glucose addition had variable effects on secreted protease activity; in some species, it triggered activity, but in others, activity was repressed at high concentrations. Collectively, our results indicate that protease activity, a key ectomycorrhizal functional trait, is positively related to environmental N source availability but is also influenced by additional factors, such as carbon availability.     

Relationships between fungal uptake of ammonium, fungal growth and nitrogen availability in ectomycorrhizal Pinus sylvestris seedlings

 Nitrogen deposition and intentional forest fertilisation with nitrogen are known to affect the species composition of ectomycorrhizal fungal communities. To learn more about the mechanisms responsible for these effects, the relations between fungal growth, nitrogen uptake and nitrogen availability were studied in ectomycorrhizal fungi in axenic cultures and in symbiosis with pine seedlings. Effects of different levels of inorganic nitrogen (NH₄) on the mycelial growth of four isolates of Paxillus involutus and two isolates of Suillus bovinus were assessed. With pine seedlings, fungal uptake of ¹⁵N-labelled NH₄ was studied in short-term incubation experiments (72 h) in microcosms and in long-term incubation experiments (3 months) in pot cultures. For P. involutus growing in symbiosis with pine seedlings, isolates with higher NH₄ uptake were affected more negatively at high levels of nitrogen availability than isolates with lower uptake. More NH₄ was allocated to shoots of seedlings colonised by a high-uptake isolate, indicating transfer of a larger fraction of assimilated NH₄ to the host than with isolates showing lower NH₄ uptake rates. Thus low rates of N uptake and N transfer to the host may enable EM fungi avoid stress induced by elevated levels of nitrogen. Seedlings colonised by S. bovinus transferred a larger fraction of the ¹⁵N label to the shoots than seedlings colonised by P. involutus. Seedling shoot growth probably constituted a greater carbon sink in pot cultures than in microcosms, since the mycelial growth of P. involutus was more sensitive to high NH₄ in pots. There was no homology in mycelial growth rate between pure culture and growth in symbiosis, but N uptake in pure culture corresponded to that during growth in symbiosis. No relationship was found between deposition of antropogenic nitrogen at the sites of origin of the P. involutus isolates and their mycelial growth or uptake of inorganic nitrogen. Accepted: 18 September 1998     

Mycorrhizal growth in pure cultures in the presence of pesticides

The effects of pesticides on 64 ectomycorrhizal fungi of boreal forest trees were studied in vitro. The pesticides (fungicides: benomyl, chlorothalonil, copper oxychloride, maneb and propiconazole; herbicides: chlorthiamid, glyphosate, hexazinone, linuron and terbuthylazine; insecticide: cypermethrin) were selected as those commonly used in Nordic forest nurseries and afforestation sites. In general, the fungicides proved to be more toxic to ectomycorrhizal fungi than the herbicides and cypermethrin. The fungicides, chlorothalonil and propiconazole, had the clearest inhibitory effect on growth of mycorrhizal fungi. Conversely, maneb, glyphosate and terbuthylazine stimulated the growth of some mycorrhizal fungi. Leccinum versipelle and L. scabrum, Paxillus involutus and Cenococcum geophilum were the most sensitive ectomycorrhizal fungi to the various pesticides.     

Detoxification of ferulic acid by ectomycorrhizal fungi

The ectomycorrhizal fungi Laccaria amethystina and Lactarius deterrimus grown in liquid culture were used to study the fate of added ferulic acid. Laccaria amethystina degraded ferulic acid to the major metabolite vanillic acid. The intermediate vanillin was not detected. Lactarius deterrimus showed a completely different detoxification pattern. Two dimers and one trimer of ferulic acid could be identified as polymerization products of this fungus. A bioassay of the possible biological activities of ferulic acid and vanillic acid on these fungi revealed that vanillic acid was less toxic than ferulic acid for Laccaria amethystina but that both phenolic acids were toxic for Lactarius deterrimus. The results are discussed with respect to ectomycorrhizal fungal growth in the organic layer of forest soils and between living root cells of ectomycorrhizas.     

Natural Abundance of <Superscript>15</Superscript>N in Nitrogen-Limited Forests and Tundra Can Estimate Nitrogen Cycling Through Mycorrhizal Fungi: A Review

The hyphae of ectomycorrhizal and ericoid mycorrhizal fungi proliferate in nitrogen (N)-limited forests and tundra where the availability of inorganic N is low; under these conditions the most common fungal species are those capable of protein degradation that can supply their host plants with organic N. Although it is widely understood that these symbiotic fungi supply N to their host plants, the transfer is difficult to quantify in the field. A novel approach uses the natural ¹⁵N:¹⁴N ratios (expressed as δ¹⁵N values) in plants, soils, and mycorrhizal fungi to estimate the fraction of N in symbiotic trees and shrubs that enters through mycorrhizal fungi. This calculation is possible because mycorrhizal fungi discriminate against ¹⁵N when they create compounds for transfer to plants; host plants are depleted in ¹⁵N, whereas mycorrhizal fungi are enriched in ¹⁵N. The amount of carbon (C) supplied to these fungi can be stoichiometrically calculated from the fraction of plant N derived from the symbiosis, the N demand of the plants, the fungal C:N ratio, and the fraction of N retained in the fungi. Up to a third of C allocated belowground, or 20% of net primary production, is used to support ectomycorrhizal fungi. As anthropogenic N inputs increase, the C allocation to fungi decreases and plant δ¹⁵N increases. Careful analyses of δ¹⁵N patterns in systems dominated by ectomycorrhizal and ericoid mycorrhizal symbioses may reveal the ecosystem-scale effects of alterations in the plant–mycorrhizal symbioses caused by shifts in climate and N deposition. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Diversity and Spatial Structure of Belowground Plant–Fungal Symbiosis in a Mixed Subtropical Forest of Ectomycorrhizal and Arbuscular Mycorrhizal Plants

Plant–mycorrhizal fungal interactions are ubiquitous in forest ecosystems. While ectomycorrhizal plants and their fungi generally dominate temperate forests, arbuscular mycorrhizal symbiosis is common in the tropics. In subtropical regions, however, ectomycorrhizal and arbuscular mycorrhizal plants co-occur at comparable abundances in single forests, presumably generating complex community structures of root-associated fungi. To reveal root-associated fungal community structure in a mixed forest of ectomycorrhizal and arbuscular mycorrhizal plants, we conducted a massively-parallel pyrosequencing analysis, targeting fungi in the roots of 36 plant species that co-occur in a subtropical forest. In total, 580 fungal operational taxonomic units were detected, of which 132 and 58 were probably ectomycorrhizal and arbuscular mycorrhizal, respectively. As expected, the composition of fungal symbionts differed between fagaceous (ectomycorrhizal) and non-fagaceous (possibly arbuscular mycorrhizal) plants. However, non-fagaceous plants were associated with not only arbuscular mycorrhizal fungi but also several clades of ectomycorrhizal (e.g., Russula) and root-endophytic ascomycete fungi. Many of the ectomycorrhizal and root-endophytic fungi were detected from both fagaceous and non-fagaceous plants in the community. Interestingly, ectomycorrhizal and arbuscular mycorrhizal fungi were concurrently detected from tiny root fragments of non-fagaceous plants. The plant–fungal associations in the forest were spatially structured, and non-fagaceous plant roots hosted ectomycorrhizal fungi more often in the proximity of ectomycorrhizal plant roots. Overall, this study suggests that belowground plant–fungal symbiosis in subtropical forests is complex in that it includes “non-typical” plant–fungal combinations (e.g., ectomycorrhizal fungi on possibly arbuscular mycorrhizal plants) that do not fall within the conventional classification of mycorrhizal symbioses, and in that associations with multiple functional (or phylogenetic) groups of fungi are ubiquitous among plants. Moreover, ectomycorrhizal fungal symbionts of fagaceous plants may “invade” the roots of neighboring non-fagaceous plants, potentially influencing the interactions between non-fagaceous plants and their arbuscular-mycorrhizal fungal symbionts at a fine spatial scale.     

Elevated CO2 impacts ectomycorrhiza-mediated forest soil carbon flow: fungal biomass production,respiration and exudation

Large quantities of carbon are exchanged between terrestrial ecosystems and the atmosphere, and extensive research efforts are made to understand carbon cycling and the impact of elevated atmospheric CO₂ levels. The response of soils to increased carbon availability is largely driven by root associated ectomycorrhizal fungi in forest ecosystems, since they partition host derived carbon belowground. In this review I examine how CO₂ enrichment affects ectomycorrhizal fungal biomass production, exudation, respiration, soil carbon fluxes, and other soil microbes, and the importance of the fungal species in these responses. I briefly discuss the significance of CO₂ alterations in the mycorrhizal symbiosis in the context of consequences for carbon sequestration, and present research priorities.     

Ectomycorrhizae and landfill site reclamations: fungal selection criteria

The ectomycorrhizal fungi Laccaria proxima and Hebeloma crustuliniforme , but not Paxillus involutus , tolerated reduced oxygen tensions characteristic of landfill site covering materials. Compared with conventional methods of ectomycorrhizal seedling production, significant time savings were made by use of fermenter cultured L. proxima inoculum. Further time savings could not, however, be made by medium supplementation with Betula pendula (Silver Birch) extract.
Since vertical leachate migration is common in most sites, fungal choice in relation to pH tolerance must be an important selection criterion. With pH regimes between acid and neutral, L. proxima should be chosen in contrast with basophilic regimes where H. crustuliniforme should be used.     

Ectomycorrhizal fungi in culture respond differently to increased carbon availability

Carbon (C) availability to ectomycorrhizal fungi is likely to increase at elevated atmospheric CO(2). To determine whether there are any broad patterns in species' responses that relate to their ecology, we compared growth, respiration, N uptake and C exudation of 17 fungal isolates in liquid culture. As a surrogate for increased C availability we used three different C:N ratios (10:1, 20:1 and 40:1), moving from conditions of C limitation to conditions of N limitation. Responses were species-specific, and suilloid fungi were the most responsive in terms of growth and respiration. In contrast, a group of eight isolates showed no growth increase above C:N 20:1. This inability to respond was not due to N limitation, although there were marked differences in N uptake between isolates. At higher C availability isolates generally became more efficient in converting C into biomass. Six isolates showed net release of exudates into the culture medium (up to 40% of the C in biomass and respiration). We conclude that the findings were in agreement with field observations, and suggest that pure culture observations can yield ecologically relevant information on how ectomycorrhizal fungi may respond under conditions of elevated CO(2).     

Changes in microbial activities and biomasses over a forest floor gradient in C-to-N ratio

Background and aims

Under chronically elevated N deposition, N retention mainly occur at high soil C-to-N ratio. This may be mediated through soil microbes, such as ectomycorrhizal (EM) fungi, saprotrophic fungi and bacteria, and the aim of this study was to evaluate the relationship between soil microbes and forest floor C-to-N ratios.

Methods

Soil samples from 33 Norway spruce (Picea abies (L.) H. Karst) forests in Denmark and southern Sweden in a forest floor C-to-N ratio gradient (ranging from 14 to 35) were analysed regarding the content of phospholipid fatty acids (PLFAs) to estimate their soil microbial community composition and the relative biomasses of different microbial groups. The relation of EM biomass to total fungal biomass was estimated as the loss of the fungal PLFA 18:2ω6,9 during incubation of soils and the production of EM mycelia was estimated using fungal in-growth mesh bags. The soil microbial variables were correlated to forest floor C-to-N ratio, NO ₃ ^- leaching, soil pH and stand age.

Results

Fungal proportions of microbial biomass, EM to total fungi and EM mycelial production were all positively related to C-to-N ratio, while NO ₃ ^- leaching was negatively related to C-to-N ratio.

Conclusions

Both EM and saprotrophic fungi change with forest floor C-to-N ratios and appear to play a central role in N retention in forest soil. A better understanding of the mechanisms behind this process may be revealed if the role of recalcitrant fungal metabolites for N retention (and soil C sequestration) can be identified. Research along this line deserves further studies.