Potential for monoterpenes to affect ectomycorrhizal and saprotrophic fungal activity in coniferous forests is revealed by novel experimental system

The fungal community in coniferous forest soils plays a pivotal role in ecosystem processes such as decomposition and carbon and nutrient cycling. Both saprotrophic (SP) and ectomycorrhizal (ECM) fungi occur throughout the upper soil horizons in coniferous forests and could therefore be exposed to high concentrations of monoterpenes occurring in the needle litter and roots of some tree species. Previous work has noted the differential effects of monoterpenes on the mycelial growth of a range of both SP and ECM fungi when grown in artificial nutrient media. This study used a novel experimental system to assess the effect of environmentally relevant concentrations of monoterpenes on the activity of ECM and SP fungi grown on more natural substrata. Exposure of the ECM fungus Paxillus involutus (Batsch) Fr. to vapours of either α-pinene or β-pinene resulted in a significantly greater proportion of root tips being colonised by the fungus when it was grown with seedlings of Picea abies (L.) Karst. Exposure to monoterpenes resulted in a significant decrease in respiration rate of two species of litter degrading SP fungi, Mycena galopus var. candida J. E. Lange and Collybia butyracea (Bull.) P. Kumm. There was no difference in response between the two SP species, despite the fact that previous tests in liquid nutrient media, with monoterpenes at higher concentrations, indicated that one species was sensitive and one was not. The high volatility and low solubility of monoterpenes in water make them challenging to work with. The experimental system developed here, although still artificial, provides a bridge between pure culture studies in defined media and all the complexities of forest soils in the field, by allowing the exposure of fungi to environmentally relevant monoterpene concentrations in more natural substrata.     

Distribution of monoterpenes between organic resources in upper soil horizons under monocultures of Picea abies, Picea sitchensis and Pinus sylvestris

The aim of this study was to compare the monoterpene content and distribution in litters and roots of three conifer species: Picea abies (L.) Karst, Picea sitchensis (Bong.) Carr. and Pinus sylvestris (L.). We analysed the monoterpene content of green needles, needle litter, F (fermentation) layer material and roots collected from monoculture plots. The rate of loss of monoterpenes from freshly fallen litter in the field was also studied at two monthly intervals over 10 months, to assess the length of time that monoterpenes entering the litter layer remain. Monoterpene analysis was carried out by extracting homogenised samples in hexane and identifying and quantifying the resulting monoterpenes using gas chromatography with flame ionisation detection (GC-FID) and gas chromatography-mass spectrometry (GC-MS). Mean total monoterpene concentrations varied significantly between the three species examined (e.g. in freshly fallen litter 1531 ± 96, 100 ± 5 and 1175 ± 122 μg g⁻¹ d. wt for P. abies, P. sitchensis and P. sylvestris); each species had distinctive and consistent monoterpene profiles associated with each type of tissue, and total monoterpene concentrations in green needles varied between individual trees of the same species, particularly for P. sitchensis. A substantial proportion of the monoterpene content of green needles remained in the needles after litter fall for P. abies (42%), P. sitchensis (11%) and P. sylvestris (30%). Although rates of monoterpene loss from needle litters varied initially (P. sylvestris > P. abies > P. sitchensis), the majority of the monoterpene content was lost after 4-6 months. Maximum monoterpene emission rates from decaying litter were calculated of 39 (P. abies), 1.7 (P. sitchensis) and 39 μg m⁻² h⁻¹ (P. sylvestris). Monoterpene concentrations in F layer material were very low (<10 μg g⁻¹ d. wt). Roots, particularly in P. sylvestris, represented a significant pool of monoterpenes (185 ± 16, P. abies; 258 ± 54, P. sitchensis; 2133 ± 200 μg g⁻¹ d. wt, P. sylvestris). The monoterpene profile was similar between roots and litter of P. sylvestris (α-pinene most abundant), and for P. sitchensis, (limonene and α-pinene most abundant), although a different pattern was observed between needle litter (most abundant β-pinene) and roots (most abundant myrcene) of P. abies. The relatively high concentrations and different profiles of monoterpenes characterised in upper organic soil horizons here emphasise the need for their influence on soil ecological processes to be assessed.     

Volatile monoterpenes in soil atmosphere under birch and conifers: Effects on soil N transformations

The aim of this study was to examine the occurrence and concentrations of volatile organic compounds (VOCs), in particular, volatile monoterpenes, in soil atmosphere under silver birch (Betula pendula L.) and two conifers, Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.), and to determine the effects of the most relevant monoterpenes on transformations of soil N. The study site was a 70-year-old tree species experiment in Kivalo, northern Finland. VOCs were collected using two methods, passive air samplers and a chamber method. In soil atmosphere under spruce and especially under pine, the concentrations of monoterpenes were high, α- and β-pinene, Δ-3-carene and myrcene being the most abundant compounds, whereas concentrations of monoterpenes in soil atmosphere under birch were negligible. Samples of humus layer from the birch stand incubated in vitro and exposed to vapors from monoterpenes typical of coniferous forest soil showed decreased rates of net N mineralization but simultaneously increased rates of C mineralization. The response of soil microbial biomass C and N to different monoterpenes varied, but some monoterpenes considerably decreased soil microbial biomass. Altogether these results suggest that these compounds have negative effects on soil N transformations, but may serve as carbon and energy source for part of soil microbes.     

Effect of litter quality on its decomposition in broadleaf and coniferous forest

Recently there has been much interest in the effect of litter mixing as well as the effect of different forest habitats on the decomposition process. Our aim was to test two hypotheses: high quality litter promotes decomposition of poor quality litter, and litter decomposes faster in broadleaf than in coniferous forest. We conducted a litter mixing experiment using litterbags placed in two forest floors, in which treatments consisted of litter monocultures of each of two campy species (Castanopsis eyrei and Pinus massoniana), as well as mixtures of these two species. The results showed that C. eyrei leaves decomposed significantly faster in the coniferous habitat than in their native habitat. On the other hand, P. massoniana needles decomposed significantly faster in their native coniferous habitat than in the broadleaf habitat. In our experiment we found that the mixture had different effect on different quality litter. P. massoniana needles (poor quality) had a positive effect on the decomposition of C. eyrei leaves (high quality), while C. eyrei leaves had a negative effect on the decomposition of P. massoniana needles in the mixture case in both broadleaf and coniferous habitats. The diversity of the fungi identified from different litters varied among treatments and the mass loss was positively correlated with the Shannon–Weaver diversity index of fungi. It is suggested that fungi may be one of the major drivers to control the decomposition process.     

Indirect host effect on ectomycorrhizal fungi: Leaf fall and litter quality explain changes in fungal communities on the roots of co-occurring Mediterranean oaks

Host trees can modify their soil abiotic conditions through their leaf fall quality which in turn may influence the ectomycorrhizal (ECM) fungal community composition. We investigated this indirect interaction using a causal modelling approach. We identified ECM fungi on the roots of two coexisting oak species growing in two forests in southern Spain - Quercus suber (evergreen) and Quercus canariensis (winter deciduous)-using a PCR-based molecular method. We also analysed the leaf fall, litter and soil sampled beneath the tree canopies to determine the concentrations of key nutrients. The total mycorrhizal pool was comprised of 69 operational taxonomic units (OTUs). Tomentella and Russula were the most species-rich, frequent and abundant genera. ECM fungi with epigeous and resupinate fruiting bodies were found in 60% and 34% of the identified mycorrhizas, respectively. The calcium content of litter, which was significantly higher beneath the winter-deciduous oak species due to differences in leaf fall quality, was the most important variable for explaining ECM species distribution. The evaluation of alternative causal models by the d-sep method revealed that only those considering indirect leaf fall-mediated host effects statistically matched the observed covariation patterns between host, environment (litter, topsoil, subsoil) and fungal community variables.     

Microfungal communities in soil,litter and casts of Lumbricus terrestris L. (Lumbricidae): a laboratory experiment

The anecic earthworm Lumbricus terrestris L. was kept in laboratory microcosms containing beech forest soil without litter, with beech leaf litter or with lime leaf litter. The structure of microfungal communities in soil, litter and fresh and aged (100 days) earthworm faeces was analysed using the washing and plating technique. The passage of mineral soil through the gut of L. terrestris affected the structure of the fungal community only little. In contrast, in the litter treatments the structure of the fungal community in fresh earthworm casts significantly differed from that in soil and litter. The majority of soil and litter inhabiting fungi survived passage through the gut of L. terrestris and the fungal community in casts consisted of a mixture of soil and litter inhabiting fungi. However, the frequency of Cladosporium spp., Alternaria spp., Absidia spp., and other taxa was strongly reduced in fresh casts. The degree of colonization of litter particles (number of isolates per number of plated particles) also decreased, but some fungi (mainly Trichoderma spp.) benefited from gut passage and flourished in fresh casts. During ageing of cast material the dominance structure of the fungal community changed. Both the degree of colonization of organic particles and the species diversity increased and approached that in soil. However, the structure of the fungal community in casts remained cast specific even after 100 days of incubation. It is concluded that the feeding and burrowing activity of L. terrestris accelerates the colonization of litter by the edaphic mycoflora but also extends the range of occurrence of litter-associated fungi into mineral soil layers.     

Saprotrophic fungi transform organic phosphorus from spruce needle litter

Fungal decomposition of and phosphorus transformation from spruce litter needles (Picea abies) were simulated in systems containing litter needles inoculated with individual saprotrophic fungal strains and their mixtures. Fungal strains of Setulipes androsaceus (L.) Antonín, Chalara longipes (Preus) Cooke, Ceuthospora pinastri (Fr.) Höhn., Mollisia minutella (Sacc.) Rehm, Scleroconidioma sphagnicola Tsuneda, Currah & Thormann and an unknown strain NK11 were used as representatives of autochthonous mycoflora. Systems were incubated for 5.5 months in laboratory conditions. Fungal colonization in systems and competition among strains were assessed using the reisolation of fungi from individual needles. After incubation, needles were extracted with NaOH and extracts were analysed using ³¹P nuclear magnetic resonance spectroscopy (NMR). Needle decomposition was determined based on the decrease in C:N ratio. Systems inoculated with the basidiomycete S. androsaceus revealed substantial decrease in C:N ratio (from 25.8 to 11.3) while the effect of ascomycetes on the C:N ratio was negligible. We suppose that tested strains of saprotrophic ascomycetes did not participate substantially in litter decomposition, but were directly involved in phosphorus transformation and together with S. androsaceus could transform orthophosphate monoesters and diesters from spruce litter needles into diphosphates, polyphosphates and phosphonates. These transformations seem to be typical for saprotrophic fungi involved in litter needle decomposition, although the proportion of individual phosphorus forms differed among studied fungal strains. Phosphonate presence in needles after fungal inoculation is of special interest because no previous investigation recorded phosphonate synthesis and accumulation by fungi. Our results confirmed that the ³¹P NMR spectroscopy is an excellent instrumental method for studying transformations of soil organic phosphorus during plant litter decomposition. We suggest that polyphosphate production by S. androsaceus may contribute to the phosphorus cycle in forest ecosystems because this fungus is a frequent litter colonizer that substantially participates in decomposition.     

Photoacceleration of plant litter decomposition in an arid environment

In arid ecosystems, abiotic processes facilitate the physical and chemical degradation of plant litter to the extent that decomposition models that use climatic and litter composition variables as surrogates for microbial activity are not predictive. The purpose of this study was to estimate the potential contribution of photodegradation to the decomposition of plant litters that varies in architecture and chemical composition. Litter of Pinus edulis, Juniperus monosperma and Populus deltoides were exposed to ambient and attenuated sunlight, with and without supplemental water additions, at a riparian forest site along the Middle Rio Grande (New Mexico, USA). Mass loss, elemental composition, and microbial extracellular enzyme activities (EEA) were measured over 639 days. The composition of the fungal communities associated with the decomposing litters was compared by analyses of fungal ITS nrDNA sequences. Litter exposed to ambient sunlight had greater mass loss rates than shaded litter, independent of the water treatment: Populus increased by 100%, Pinus by 86% and Juniperus by 46%. The increases were proportional to exposed litter surface area per g dry mass. EEA potentials, particularly oxidative activities, were low in comparison to those measured in mesic ecosystems. For Populus litter, the principal driver of photoacceleration appeared to be photodegradation of cellulose; for Pinus, it was photodegradation of polyphenols; for Juniperus accelerated mass loss was associated with photodegradation of both polysaccharides and polyphenols. Fungal community composition varied by litter type, but the dominant colonizers were yeasts and dark-septate hyphal taxa; a finding consistent with the low enzymatic oxidation potential. This study shows that photochemical oxidation can supplement enzymatic oxidation and increase decomposition rates. As a result, organic matter decomposition in arid ecosystems is not restricted to periods of high moisture availability as is plant production. This decoupling may partly account for the low soil organic matter content of these ecosystems.     

Do oribatid mites (Acari: Oribatida) show a higher preference for ubiquitous vs. specialized saprotrophic fungi from pine litter?

Previous studies of oribatid mite feeding preferences for different saprotrophic fungi were limited to ubiquitous fungal species, whereas saprophytes specialized to decompose particular substrates have been neglected. We examined the preference of seven oribatid mite species (Adoristes ovatus, Eniochthonius minutissimus, Eueremaeus silvestris, Nothrus silvestris, Oppiella subpectinata, Porobelba spinosa and Spatiodamaeus verticillipes) for nine autochthonous saprotrophic fungi from Scots pine litter (Pinus sylvestris). Among the fungal species offered were specific coniferous litter colonizers (Allantophomopsis lycopodina, Ceuthospora pinastri, Hormonema dematioides, Scleroconidioma sphagnicola, Verticicladium trifidum, Marasmius androsaceus and Sympodiella acicola) and two ubiquitous species (Cladosporium herbarum and Oidiodendron griseum). The fungi were inoculated on fragments of pine needles and offered simultaneously and separately to the mites. Our main hypothesis, that oribatid mites (usually occurring in more than one type of ecosystems) would prefer the ubiquitous fungal species rather than those specific to pine litter, was supported only partly. The ubiquitous C. herbarum was highly preferred by all studied mites, but most of them preferred one or more of the specialized fungi with similar intensity. The basidiomycete M. androsaceus along with sterile needles were consistently avoided by all mites in all experiments. Our results do not support the hypothesis, that the “true” fungivorous oribatid mites in traditional sense are more selective fungal feeders than are the “unspecialized” panphytophagous ones. We observed no gradation in preference of fungi for oribatid mites as a group, but rather a discontinuous and dynamic mosaic with particular mites preferring particular fungal species. This heterogeneous mosaic shapes the feeding niches occupied by particular oribatid mite species and probably reduces competition for food source among numerous species coexisting in a given habitat and time.     

西藏原始暗针叶林凋落物有机碳释放特征与土壤有机碳库关系研究

结合野外凋落物分解袋法和室内分析试验,对藏东南2种典型暗针叶林—急尖长苞冷杉(Abies georgei var.Smithii)和林芝云杉(Picea likiangensis var.linzhiensis)凋落物的分解和有机碳释放特征进行研究,分析了2种亚高山暗针叶林凋落物有机碳释放速率与土壤有机碳及其组分之间的关系。结果表明:藏东南2种原始暗针叶林凋落物分解均呈现出雨季分解快(4—9月)、旱季分解慢(10—翌年3月),前期分解快(3—9月)、后期分解慢(10—翌年2月)的特征,且冷杉(PLLF)分解速率大于云杉(AGSF),Olson指数衰减模型能够较好地模拟2种暗针叶林凋落物的分解,冷杉(PLLF)和云杉(AGSF)凋落物半分解时间为2.11,2.52年;分解95%时间为8.96,10.84年;2种暗针叶林凋落物中有机碳含量表现出先上升后下降,再平稳降低的趋势,而2种暗针叶林凋落物中有机碳释放速率表现出先短暂富集再释放的模式;2种暗针叶林土壤总有机碳(TOC)及其活性组分(MBC、POC、LOC)含量都具有明显的表聚性(p<0.01),且同一土壤层次内TOC、MBC、POC、LOC互相之间均呈极显著正相关关系(p<0.01);2种暗针叶林凋落物分解进程中有机碳的释放速率与表层土(0—10 cm)中TOC、MBC、POC、LOC含量、10—20 cm土层中的TOC、MBC含量以及20—40 cm土层中MBC含量之间呈现显著的正相关(p<0.05)。     

Production of lignocellulose-degrading enzymes and degradation of leaf litter by saprotrophic basidiomycetes isolated from a Quercus petraea forest

Due to the production of lignocellulose-degrading enzymes, saprotrophic basidiomycetes can significantly contribute to the turnover of soil organic matter. The production of lignin- and polysaccharide-degrading enzymes and changes of the chemical composition of litter were studied with three isolates from a Quercus petraea forest. These isolates were capable of fresh litter degradation and were identified as Gymnopus sp., Hypholoma fasciculare and Rhodocollybia butyracea. Within 12 weeks of incubation, H. fasciculare decomposed 23%, R. butyracea 32% and Gymnopus sp. 38% of the substrate dry mass. All fungi produced laccase and Mn-peroxidase (MnP) and none of them produced lignin peroxidase or other Mn-independent peroxidases. There was a clear distinction in the enzyme production pattern between R. butyracea or H. fasciculare compared to Gymnopus sp. The two former species caused the fastest mass loss during the initial phase of litter degradation, accompanied by the temporary production of laccase (and MnP in H. fasciculare) and also high production of hydrolytic enzymes that later decreased. In contrast, Gymnopus sp. showed a stable rate of litter mass loss over the whole incubation period with a later onset of ligninolytic enzyme production and a longer lasting production of both lignin and cellulose-degrading enzymes. The activity of endo-cleaving polysaccharide hydrolases in this fungus was relatively low but it produced the most cellobiose hydrolase. All fungi decreased the C/N ratio of the litter from 24 to 15-19 and Gymnopus sp. also caused a substantial decrease in the lignin content. Analytical pyrolysis mass spectrometry of litter decomposed by this fungus showed changes in the litter composition similar to those caused by white-rot fungi during wood decay. These changes were less pronounced in the case of H. fasciculare and R. butyracea. All fungi also changed the mean masses of humic acid and fulvic acid fractions isolated from degraded litter. The humic acid fraction after degradation by all three fungi contained more lignin and less carbohydrates. Compared to the decomposition by saprotrophic basidiomycetes, litter degradation in situ on the site of fungal isolation resulted in the relative enrichment of lignin and differences in lignin composition revealed by analytical pyrolysis. It can most probably be explained by the participation of non-basidiomycetous fungi and bacteria during natural litter decomposition.     

Decomposition and substrate quality of leaf litters and fine roots from three exotic plantations and a native forest in the southwestern highlands of Ethiopia

Substrate quality and decomposition (measured as CO₂ release in laboratory microcosms) of fresh leaf litter and fine roots of Cupressus lusitanica, Pinus patula, Eucalyptus grandis and native forest trees were studied. Changes in litter chemistry in each forest stand were analysed by comparing fresh leaf litter (collected from trees) and decomposed litter from the forest floor. Elemental concentrations, proximate fractions including monomeric sugars, and cross polarisation magic-angle spinning (CPMAS) ¹³C NMR spectra were analysed in leaf litters, decomposed litter and fine roots. Leaf litters and fine roots varied in their initial substrate chemistry with Ca concentration in leaf litters being higher than that in fine roots. In each stand, fine roots had a higher acid unhydrolysable residue (AUR) (except for the Pinus stand), higher holocellulose concentration and lower concentration of water-soluble extractives (WSE) and dichloromethane extractives (NPE) than fresh leaf litter. Likewise, ¹³C NMR spectra of fine roots showed lower alkyl and carboxyl C, and higher phenolic (except P. patula), aromatic and O-alkyl C proportions than leaf litters. Compared with fresh leaf litter, decomposed litter had lower concentrations of potassium, holocellulose, WSE, NPE, arabinose and galactose, similar or higher concentrations of Mg, Ca, S and P, and higher concentrations of N and AUR. CPMAS ¹³C NMR spectra of decomposed litter showed a higher relative increase in signal intensity due to methoxyl C, aromatic C, phenolic C and carboxylic C compared with alkyl C. In a microcosm decomposition study, the proportion of initial C remaining in leaf litter and fine roots significantly fitted an exponential regression model. The decomposition constants (k) ranged between 0.0013 and 0.0030 d⁻¹ for leaf litters and 0.0010-0.0017 d⁻¹ for fine roots. In leaf litters there was a positive correlation between the k value and the initial Ca concentration, and in fine roots there was an analogous positive correlation with initial WSE. Leaf litters decomposed in the order Cupressus>native forest>Eucalyptus∼Pinus, and fine roots in the order Pinus>native forest>Cupressus∼Eucalyptus. In each stand the fine root decomposition was significantly lower than the leaf litter decomposition, except for the P. patula stand where the order was reversed.     

Direct and indirect effects of enhanced UV-B radiation on the decomposing and competitive abilities of saprobic fungi

Increases in UV-B radiation have been shown to slow the rate of litter decomposition in ecosystems. However, it is unclear if this is a result of direct UV-B effects on saprobic microorganisms, or a result of UV-B-induced changes in litter quality that indirectly affect decay by saprobes. In this study, we evaluated the magnitude of direct and indirect effects on litter decomposition of Brassica napus by soil fungi, under growth chamber conditions. We found that, both, direct and indirect UV-B negatively influenced litter decomposition, however, direct effects were much more pronounced. We then tested whether UV-B radiation would have species-specific effects on fungal colonization and competitive ability, rather than influencing all fungal species equally. We predicted that darkly pigmented fungi would increase their relative competitive ability under high UV-B. The test fungi were all isolated from field soil under Brassica napus. Two fungi were hyaline (Aspergillus terreus, Trichoderma koningii), two were darkly-pigmented (Cladosporium sphaerospermum, Epicoccum purpurascens) and one had a hyaline mycelium but darkly-pigmented conidia (Aspergillus niger). Elevated UV-B radiation had differential direct and indirect effects on fungal growth, and caused shifts in the competitive balances between pigmented and non-pigmented fungi. However, in only two of six pair-wise challenges did the pigmented species increase their relative competitive ability under UV-B conditions. It is clear that UV-B profoundly influence fungal community structure in soil, but the direction of such effects remains unpredictable.     

Dynamics and stratification of bacteria and fungi in the organic layers of a scots pine forest soil

The abundance and micro-stratification of bacteria and fungi inhabiting the organic layers of a Scots pine forest (Pinus sylvestris L.) were investigated. An experiment using stratified litterbags, containing organic material of four degradation stages (fresh litter, litter, fragmented litter and humus) was performed over a period of 2.5 years. Dynamics and stratification of fluorescent stained bacteria and fungi, ratios between bacterial and fungal biomass, and relationships with moisture and temperature are described. Average bacterial counts in litter and fragmented litter were similar, i.e., approximately 5×10⁹ bacteriag^–1 (dry weight) organic matter, and significantly exceeded those in humus. The mean bacterial biomass ranged from 0.338 to 0.252mg carbon (C) g^–1 (dry weight) organic matter. Lengths of mycelia were significantly below the usually recorded amounts for comparable temperate coniferous forests. The highest average hyphal length, 53mg^–1 (dry weight) organic matter, was recorded in litter and decreased significantly with depth. The corresponding mean fungal biomass ranged from 0.050 to 0.009mg Cg^–1 (dry weight). The abundance of bacteria and fungi was influenced by water content, that of fungi also by temperature. A litterbag series with freshly fallen litter of standard quality, renewed bimonthly, revealed a clear seasonal pattern with microbial biomass peaks in winter. The mean hyphal length was 104mg^–1 (dry weight) and mean number of bacteria, 2.40×10⁹ bacteria g^–1 (dry weight). Comparable bacterial and fungal biomass C were found in the freshly fallen litter [0.154 and 0.132mgCg^–1 (dry weight) organic material, respectively]. The ratio of bacterial-to-fungal biomass C increased from 1.2 in fresh litter to 28.0 in humus. The results indicate the existence of an environmental stress factor affecting the abundance of fungi in the second phase of decomposition. High atmospheric nitrogen deposition is discussed as a prime factor to explain low fungal biomass and the relatively short lengths of fungal hyphae in some of the forest soil layers under study. Received: 26 June 1997     

Effects of inoculation with ectomycorrhizal fungi on microbial biomass and bacterial functional diversity in the rhizosphere of Pinus tabulaeformis seedlings

This study investigated the effects of inoculation with three individual ectomycorrhizal (ECM) fungal species on soil microbial biomass carbon and indigenous bacterial community functional diversity in the rhizosphere of Chinese pine (Pinus tabulaeformis Carr.) seedlings under field experimental conditions. The results showed that ECM fungal inoculation significantly increased the ectomycorrhizal colonization compared with non-inoculated seedlings. ECM fungal inoculations have higher soil microbial biomass carbon than that of control, ranging from 49.6 μg C g^?1 dry soil in control to 134.02 μg C g^?1 dry soil in treatment inoculated with Boletus luridus Schaeff ex Fr. Multivariate analyses (PCA) of BIOLOG data revealed that the application of ECM fungi significantly influenced bacterial functional diversity in the rhizosphere of P. tabulaeformis seedlings. The highest average well-color development (AWCD) and functional diversity indices were also observed in treatment inoculated with B. luridus. A wider range of sole carbon sources were utilized by the bacterial community in the rhizosphere of inoculated seedlings. The data gathered from this study provides important information for utilization of ECM fungi in forest restoration project in the Northwestern China. The present study will also significantly broaden our understanding of practical importance in the application of ECM fungal inoculum to promote soil microbial community diversity of soil.     

Interactions between litter quality, decomposition and soil fertility: a laboratory study

Leaf litters from beech (Fagus sylvatica L.) and oak (Quercus robur L.) trees were collected from mixed, deciduous woodlands growing on three soil types that varied in mineral nutrient concentrations and N mineralisation potential. Litter quality, including %N, %Mn, %P, acid detergent fibre, cellulose, Klason lignin, phenylpropanoid constituents of lignin, hexose and pentose sugar (mainly from hemicelluloses) varied within species according to soil type. However, oak and beech showed the opposite responses to soil nutrient status for most of these variables. The litters were incubated in the laboratory for 12 months (at 18 °C and constant moisture) on beds of forest floor material from two soils of contrasting high nutrient material (HNM) or low nutrient material (LNM) nutrient status to investigate litter quality and substrate interactions. At 4, 8 and 12 months there were significant differences in mass losses from oak and beech litters from all sites, and for each litter type exposed to the HNM and LMN soils. At 12 months mean mass losses were higher for HNM treatment (38.7% oak, 27.8% beech) than for the LNM treatment (30.6% oak, 25.5% beech). However, the beech and oak litters from the different sites consistently responded in opposite ways on the same soil treatment reflecting site-related effects on litter quality. Initial concentration of Klason lignin was the best predictor for mass losses from litter species and litter types. Intra-specific variation in rates of litter decomposition of beech and oak litters from different sites, and differences in their interactions with the two forest floor materials, illustrate the complexities of proximate controls on decomposition that are often masked in system-level studies.     

Effects of nitrogen addition and litter properties on litter decomposition and enzyme activities of individual fungi

Litter decomposition is an important process of C and N cycling in the soil. Variation in the response of litter decomposition to nitrogen (N) addition (positive, negative or neutral) has been observed in many field studies. However, mechanism about variability in individual fungal species response to N addition has not yet been well demonstrated in the literature. Therefore, the objective of this study was to investigate the effects of N addition and litter chemistry properties on litter decomposition and enzyme activities of individual fungi. Three fungal species (Penicillium, Aspergillus, and Trichoderma) were isolated from a subtropical mixed forest soil. An incubation experiment was conducted using the individual fungi with two types of litter (leaf of Pinus massoniana and needle of Cryptocarya chinensis) and different N addition levels (0, 50 and 100 for N-deficient treatments, and 500 and 1000 μg N for N-excessive treatments). Cumulative CO₂-C, enzyme activities, and lignin and cellulose loss were measured during the incubation period of 60 days. Litter decomposition and enzyme activities significantly varied with the fungal species, while the N addition and litter types greatly affected fungal enzyme activities. The N treatments significantly increased lignin-rich needle decomposition by lignocellulose decomposers (Penicillium and Aspergillus) but did not affect their leaf decomposition. On the contrary, The N treatments stimulated leaf decomposition by cellulolytic species (Trichoderma) but did not affect its needle decomposition. Correlation analysis showed that lignin in the litter was the key component to affect litter decomposition. Activities of N-acetyl-β-glucosaminidase and phenol oxidase were both positively correlated to litter decomposition. The fungi (Penicillium and Aspergillus) with higher production of N-acetyl-β-glucosaminidase showed higher litter decomposition ability. The low N addition levels stimulated Penicillium and Aspergillus litter decomposition, but they still required more N source (e.g., litter N source) to support decomposition. Depressed fungal litter N uptake (lower N-acetyl-β-glucosaminidase activities) only occurred at the highest N addition level. Litter decomposition of Trichoderma depended more on external N and its litter decomposition capability was the lowest among the three species.     

Isolation and characterization of a mycorrhiza helper bacterium from rhizosphere soils of poplar stands

If mycorrhizal formation could be enhanced by co-inoculation with mycorrhiza helper bacteria (MHB) which promote rapid root colonization by specific ectomycorrhizal fungi, this would be of advantage to the poplar forest industry. A number of poplar rhizobacterial strains were isolated from 11 regions of the eastern China. Four of the isolates, SY15, DZ18, HLJ4, and PY10, were characterized as MHB potential strains based on their positive effect on growth of ectomycorrhizal fungi Pisolithus tinctorius (Pers.) and Lactarius insulsus (Fr.). Under greenhouse conditions, one of the bacterial isolate, DZ18, significantly promoted the poplar trees growth and ectomycorrhizal colonization of P. tinctorius and L. insulsus on Populus deltoides Marsh. In contrast, the other three isolates SY15, HLJ4, and PY10 promoted fungal growth in vitro experiments but did not enhance ectomycorrhizal (ECM) formation in the greenhouse experiment. Therefore, it was concluded that DZ18 can be considered as an MHB strain. DZ18 was identified as Bacillus sp. based on morphological, physiological, and biochemical analyses in combination with analysis of 16S rDNA gene sequences.     

Organic acid exudation by mycorrhizal Andropogon virginicus L. (broomsedge) roots in response to aluminum

Elevated aluminum (Al) availability limits plant growth on acidic soils. Although this element is found naturally in soils, acidic conditions create an environment where Al solubility increases and toxic forms of Al impact plant function. Plant resistance to Al is often attributed to organic acid exudation from plant roots and the chelation of cationic Al in the rhizosphere. The association of arbuscular mycorrhizal (AM) fungi with the roots of plants may alleviate Al toxicity by altering soil Al availability or plant exposure through the binding of Al to fungal structures or through the influence of fungi on exudation from roots. Diverse communities of AM fungi are found in soil ecosystems and research suggests that AM fungi exhibit functional diversity that may influence plant performance under varying edaphic environments. In the present study, we evaluated acidic isolates of six AM species in their responses to Al. Andropogon virginicus (broomsedge), a warm-season grass that commonly grows in a range of stressful environments including acidic soils, was used as a plant host for Acaulospora morrowiae, Glomus claroideum, Glomus clarum, Glomus etunicatum, Paraglomus brasilianum, and Scutellospora heterogama. Fungal spores were germinated and exposed to 0 or 100 μM Al on filter paper in sand culture or were grown and exposed to Al in sand culture in association with A. virginicus. Short- and long-term responses to Al were evaluated using direct measurements of fungal spore germination, hyphal elongation, and measurements of A. virginicus colonization and plant growth as a phytometer of AM function in symbio. Spore germination and hyphal elongation varied among AM species in response to Al, but patterns were not consistent with the influences of these AM species on A. virginicus under Al exposure. Exposure to Al did not influence colonization of roots, although large differences existed in colonization among fungal species. Plants colonized by G. clarum and S. heterogama exhibited the least reduction in growth when exposed to Al, produced the highest concentrations of Al-chelating organic acids, and had the lowest concentrations of free Al in their root zones. This pattern provides evidence that variation among AM fungi in Al resistance conferred to their plant hosts is associated with the exudation of Al-binding organic acids from roots and highlights the role that AM fungal diversity may play in plant performance in acidic soil environments.