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.     

Initial organic matter transformation of soil amended with composted sewage sludge

 An incubation experiment with composted sewage sludge (CSS) just added to the soil was conducted to determine its initial effects on C decomposition, N nitrification and the transformation of organic matter. CSS was mixed with a sandy loam soil from uncultivated ochric epipedon of a Typic Haploxeralf at rates of 0, 40 and 80 t ha^–1 (dry weight). The data obtained showed that with regard to the unamended soil, both the 40 and the 80 t ha^–1 treatments produced the same result in decreasing respiratory activity, but the addition of increasing amounts of CSS progressively delayed C decomposition. The nitrification index (NI), defined as the relation between nitrate-N and nitrate-N + ammonium-N, increased in correlation with the C mineralization coefficient. Total organic matter decreased after incubation whereas the humic substances increased in relation to the total C mineralized. Received: 28 October 1999     

Involvement of soluble organic matter in increased plant growth in solarized soils

 Although soil solarization is used to control soil-borne pests, it also results in increased growth response (IGR) of plants, beyond the effect of pest control. IGR is attributed to various abiotic factors (e.g. increased mineral nutrient concentrations) and biotic factors. In this work, we studied the role played by dissolved organic matter (DOM) in soil extracts in the IGR. DOM concentrations were about twice as high in solarized soil than in untreated soil. In two out of three soils, solarization appeared to increase amino acid synthesis, indicating that it had a favorable effect on microbial activity. Elemental composition, carbohydrate levels, E₄ : E₆ ratios and FTIR spectra did not differentiate between DOM extracted from solarized soils and DOM extracted from untreated soils. Growth of corn plants increased with increasing concentrations of DOM. Addition to the soil of DOM extracted from leonardite increased populations of fluorescent pseudomonads, known as beneficial bacteria, and reduced fungal populations. We conclude that the increase in DOM concentration following soil solarization is a potentially positive plant-growth-enhancement factor. Received: 21 June 1999     

Fitting plants to soil through mycorrhizal fungi: Mycorrhiza effects on plant growth and soil organic matter

Summary Vesicular-arbuscular mycorrhizal (VAM) fungi affect diverse aspects of plant form and function. Since mycorrhiza-mediated changes in host-plant responses to root colonization by different VAM fungi vary widely, it is important to assess each endophyte for each specific effect it can elicit from its host as part of the screening process for effectiveness. Three species of VAM fungi and a mixture of species were compared with non-VAM controls for their effects on soil organic matter contents and on nutrition and morphology in two varieties (native and hybrid) of corn (Zea mays L.) and one of sunflower (Helianthus annuus L.) in P-sufficient and N-deficient soil in pot cultures. Differences in soil organic matter due to the fungal applications were highly significant with all host plants. Native corn responded more to VAM colonization than the hybrid did; differences in treatments were significant in leaf area, plant biomass, and root: shoot ratio in the former, but not in the latter. Responses in the sunflower were similar to those in the native corn. Significant VAM treatment-related differences in shoot N and P contents were not reflected in shoot biomass, which was invariant. Correlations between plant or soil parameters and the intensity of VAM colonization were found only in soil organic matter with the native corn, in specific leaf area in the hybrid corn, and in plant biomass in the sunflower. The presence of the different endophytes and not the intensity of colonization apparently elicited different host responses.     

Composition of the organic phosphorus fraction in basidiocarps of saprotrophic and mycorrhizal fungi

In our screening, we aimed to detect phosphonates and other forms of organic phosphorus in basidiocarps and vegetative mycelia of six common basidiomycetes. Organic phosphorus-containing compounds were extracted in alkali and analysed using ³¹P NMR. Monoesters, diesters, pyrophosphates and polyphosphates detected in high amounts reflected the high metabolic activity in basidiocarps (growth, production of basidiospores). Phosphonates were present in all samples, in concentrations ranging from 14 mg kg⁻¹ of the extracted phosphorus in Boletus badius basidiocarp to 140 mg kg⁻¹ in Amanita muscaria vegetative mycelium. Detection of phosphonates in basidiocarps together with our previous evidence from laboratory experiments support the fungal production of natural phosphonates in forest ecosystems.     

Effect of cover crop management on soil organic matter

Characterization of soil organic matter (SOM) is important for determining the overall quality of soils, and cover crop system may change SOM characteristics. The purpose of this study was to examine the effect of cover crops on the chemical and structural composition of SOM. We isolated humic substances (HS) from soils with the following cover crop treatments: (a) vetch (Vicia Villosa Roth.)/rye (Sesale cereale L.), (b) rye alone, and (c) check (no cover crops) that were treated with various nitrogen (N) fertilizer rates. CPMAS-TOSS (cross-polarization magic-angle-spinning and total sideband suppression) ¹³C NMR results indicated that humic acids (HA) from soils under rye only were more aromatic and less aliphatic in character than the other two cover crop systems without fertilizer N treatment. Based on the DRIFT (diffuse reflectance Fourier transform infrared) spectra peak O/R ratios, the intensities of oxygen-containing functional groups to aliphatic and aromatic (referred to as recalcitrant) groups, the highest ratio was found in the HA from the vetch/rye system with fertilizer N. The lowest ratio occurred at the vetch/rye system without fertilizer N treatment. The O/R ratio of fulvic acids (FA) can be ranked as: vetch/rye without fertilizer>vetch/rye with fertilizer>no cover crop without fertilizer>rye alone (with or without fertilizer) soils. Both organic carbon (OC) and light fraction (LF) contents were higher in soils under cover crop treatments with and without fertilizer N than soils with no cover crop. These chemical and spectroscopic data show that cover crops had a profound influence on the SOM and LF characteristics.     

Assessment of biogeochemical trends in soil organic matter sequestration in Mediterranean calcimorphic mountain soils (Almería, Southern Spain)

Total soil organic matter levels and humic acid formation processes in mountain calcimorphic soils from Sierra María-Los Vélez Natural Park (Almería, Southern Spain) were found to differ depending on soil use (pine and oak forests, and cleared areas either cultivated or affected by bush encroachment). Biogeochemical indicators such as the concentration of exchangeable cations, or the concentration of the different types of humic substances were neither influenced by the type of vegetation nor soil use. In fact, multidimensional scaling and multiple correlations suggest that soil carbon sequestration processes are controlled by small-scale topographical features and their impact on water holding capacity. From a qualitative viewpoint, there were two more or less defined sets of soils: one set consisted of soils with humic acids with marked aliphatic character, displayed intense 2920 cm⁻¹ infrared band, and had low optical density. The resolution-enhanced infrared spectra suggested typical lignin patterns and well-defined amide bands, which point to a selective preservation of comparatively young organic matter. This situation contrasts with that in other set of soils with low C levels (<20 g kg⁻¹) where humic acids with featureless infrared spectra showed high aromaticity and were associated with perylenequinonic chromophors of fungal origin: this is considered the consequence of overlapping biogeochemical mechanisms involving both microbial synthesis and condensation processes. The results from visible and infrared derivative spectroscopies suggest that the reliability of statistically assessing the biogeochemical performance of the different uses on the site studied in terms of the intensity of the prevailing humic acid formation mechanisms, i.e., accumulation of inherited macromolecular substances in the former set, vs. microbial synthesis including the condensation of precursors of low molecular weight substances in the latter.     

The impact of trees, ectomycorrhiza and potassium availability on simple organic compounds and dissolved organic carbon in soil

The presence of tree roots and symbiotic mycorrhizal fungi is recognized to have a substantial impact on carbon dynamics in soils. In this study the effect of Pinus sylvestris seedlings and the ectomycorrhizal fungus Hebeloma crustuliniforme on a number of biogeochemical variables, mainly related to labile carbon pools was investigated. The impact of K limitation as a potential regulatory factor was also examined. Columns filled with E horizon ±plants and ±mycorrhizal fungi were incubated for 18.5 months. The results demonstrate that plants, as well as mycorrhizal fungi, significantly increased the concentrations of some simple organic acids, including oxalate, in soil solution. Observations for dissolved organic carbon were slightly contradictory but the cumulative amount found in drainage water was ∼20% higher in planted versus non-planted columns. Soil from planted treatments also showed more rapid mineralisation kinetics for oxalate. However carbon utilization (mineralisation vs. biomass) of oxalate and glucose by the soil microbial biomass was less influenced by plants. At harvest a component integration study of soil autotrophic and heterotrophic respiration was performed which revealed that both plant and mycorrhiza had a positive effect on the heterotrophic respiration. Potassium omission had little effect on the variables studied with the exception of the maximum mineralisation rate for oxalate, which increased when K was withdrawn. The results are discussed in the context of the dynamics of labile soil carbon pools and ecosystem C fluxes.     

水稻秸秆燃烧对土壤有机质组成的影响研究

研究生物质(水稻秸秆)燃烧行为对土壤有机质含量和组成的影响,探讨不同燃烧次数下土壤中胡敏酸(HA)、富里酸(FA)、水溶性有机质(DOM)的含量和结构特征的变化情况。结果表明:土壤有机碳的变化仅在土壤表层几厘米以内,随燃烧频次增加土壤有机碳含量减少量增加,而水溶性有机质及组分则呈现不断增加的趋势。不同来源土壤的腐殖物质对生物质燃烧作用的响应不同,3种供试土样中腐殖物质随不同次数同强度秸秆燃烧并未呈现较一致的变化趋势,但每一种土壤的HA和FA含量及HA/FA、E4/E6值均随燃烧频次的增加呈现规律性的升高或者降低,而且FA含量的变化是影响HA/FA值大小的主导因素。此外,通过红外光谱图可知,随燃烧频次增加,FA中芳香性成分和含脂肪族C—OH、C—O物质增加,而HA由于本身芳香缩和度高而无明显变化。     

Modifications of degradation-resistant soil organic matter by soil saprobic microfungi

Modifications of humic (HA) and fulvic (FA) acids in their solutions and in sterile soil by microfungal species and two well-known HA degraders were studied by measurement of total oxidizable carbon (OC), absorbances, enzyme activities and CO₂ release. The effect of glucose on FA and HA, and also minerals on FA utilization was also observed. Microfungi affected HA more than FA. Common microfungal species decolorized HA and decreased their molecular size (evaluated in terms of A₄/A₆ ratio). Some of them decreased aromaticity of HA and FA as the only carbon sources. They did not affect OC, although released CO₂ from FA. Under higher availability of mineral nutrients, the FA aromaticity increased and FA decolorization decreased. The molecular size of HA decreased in the presence of glucose. In the FA medium complemented by minerals, the known basidiomycete HA degrader, Trametes versicolor, decreased the amount of aromatic compounds in contrast to microfungal species Alternaria alternata, Clonostachys rosea, Exophiala cf. salmonis, Fusarium coeruleum, F. redolens, Penicillium canescens, Phoma sp. and another basidiomycete Phanerochaete chrysosporium. No microfungal species exhibited lignin peroxidase activity. On the other hand, activities of manganese peroxidase (MnP) were recorded for all species incubated in FA. Carbon dioxide produced from soil inoculated by microfungi negatively correlated with the decolorization, aromaticity and OC of/in FA reisolated from the soil. The results support the hypothesis that soil microfungi can attack both HA and FA and can represent an important factor in their transformations in arable soils. The enzyme involved in FA modifications is probably fungal MnP. We enriched a group of known HA and FA degraders and showed some abilities of a few frequent soil microfungal species. This can be one of the first but important step towards learning the functioning of carbon release from the big reservoir represented by humic substances in arable soils.     

Arbuscular mycorrhizal fungi contribute to C and N enrichment of soil organic matter in forest soils

Increasing evidence suggests that accretion of microbial turnover products is an important driver for isotopic carbon (C) and nitrogen (N) enrichment of soil organic matter (SOM). However, the exact contribution of arbuscular mycorrhizal fungi (AMF) to soil isotopic patterns remains unknown. In this study, we compared ¹³C and ¹⁵N patterns of glomalin-related soil protein (GRSP), which includes a main fraction derived from AMF, litter, and bulk soil in four temperate rainforests. GRSP was an abundant C and N pool in these forest soils, showing significant ¹³C and ¹⁵N enrichment relative to litter and bulk soil. Hence, cumulative accumulation of recalcitrant AMF turnover products in the soil profile likely contributes to ¹³C and ¹⁵N enrichment in forest soils. Further research on the relationship between GRSP and AMF should clarify the exact extent of this process.     

Nitrogen availability mediates the effects of roots and mycorrhizal fungi on soil organic carbon decomposition: A meta-analysis

Plant roots and their associated mycorrhizal fungi critically mediate the decomposition of soil organic carbon (C), but the general patterns of their impacts over a broad geographical range and the primary mediating factors remain unclear. Based on a synthesis of 596 paired observations from both field and greenhouse experiments, we found that living roots and/or mycorrhizal fungi increased organic C decomposition by 30.9%, but low soil nitrogen (N) availability (i.e., high soil C:N ratio) critically mitigated this promotion effect. In addition, the positive effects of living roots and/or mycorrhizal fungi on organic C decomposition were higher under herbaceous and leguminous plants than under woody and non-leguminous plants, respectively. Surprisingly, there was no significant difference between arbuscular mycorrhizal fungi and ectomycorrhizal fungi in their effects on organic C decomposition. Furthermore, roots and/or mycorrhizal fungi significantly enhanced the decomposition of leaf litter but not root litter. These findings advance our understanding of how roots and their symbiotic fungi modulate soil C dynamics in the rhizosphere or mycorrhizosphere and may help improve predictions of soil global C balance under a changing climate.     

Effect of mixing soil saprophytic fungi with organic residues on the response of Solanum lycopersicum to arbuscular mycorrhizal fungi

The effect of the dual inoculation with arbuscular mycorrhizal (AM) and saprophytic fungi and a combination of wheat straw and sewage sludge residues were studied by determining their effect on dry weight of tomato and on chemical and biochemical properties of soil. Incubation of organic residue (sewage sludge combined with wheat straw) with saprophytic fungi and plant inoculation with mycorrhizal fungi was essential to study plant growth promotion. Soil application of organic residues increased the dry weight of tomato inoculated with Rhizophagus irregularis. The greatest shoot dry mass was obtained when the organic residues were incubated with Trichoderma harzianum and applied to AM plants. However, the greatest percentage of root length colonized with AM in the presence of the organic residues was obtained with inoculation with Coriolopsis rigida. The relative chlorophyll was greatest in mycorrhizal plants regardless of the presence of either saprophytic fungus. The presence of the saprophytic fungi increased soil pH as the incubation time increased. Soil nitrogen and phosphorus contents and acid phosphatase were stimulated by the addition of organic residues, and contents of N and P. Total N and P content in soil increased when the organic residue was incubated with saprobe fungi, but this effect decreased as the incubation period of the residue with saprobe fungi increased. The same trend was observed for soil β‐glucosidase and fluorescein diacetate activities. The application of organic residues in the presence of AM and saprophytic fungi seems to be an interesting option as a biofertilizer to improve plant growth and biochemical parameters of soils.     

Soil organic matter composition as a factor affecting the accumulation of polycyclic aromatic hydrocarbons

Journal of Soils and Sediments - The objective of this study was to evaluate the mutual relations between the soil organic matter (SOM) fractions: fulvic acids (FA), humic acids (HA), humins (HN),...     

县域土壤有机质动态变化及其影响因素分析

本文以河北省曲周县为例, 采用 1980 年和 1999 年两次全县的土壤肥力监测以及农户调查数据和统计数据, 系统分析了过去 20 年中土壤有机质的动态及其与之相关的农作管理方式的变化。 结果显示, 在过去的近 20 年 间, 曲周县土壤表层的有机质含量呈现增长的趋势, 导致这种变化的农作管理方式有化肥施用量的大幅度提升、秸 秆还田量的增加、盐碱地的开垦利用、灌溉面积和复种指数的提高以及主要种植模式和种植作物的土壤有机质处 于正平衡状态。 然而当前的生产管理方式尽管有利于土壤有机质积累, 但是也带来了一系列的生态环境问题。 实 施保护性耕作、降低化肥用量、提高秸秆还田量和有机肥的用量成为今后农业生产管理方式调整的主要方向。     

耐盐碱细菌与有机物料对盐碱土团聚体形成的影响

从滨海盐碱土中分离筛选到分解秸秆、产胞外聚合物的耐盐碱细菌,通过土柱和盆栽试验研究其与有机物料相互作用对盐碱土团聚体形成以及植物生长的影响.结果表明,盐碱土中接种既分解秸秆又产胞外聚合物的耐盐碱细菌(菌株 M6),配施未腐熟秸秆,直接在盐碱土中腐熟分解,团聚体形成效果最好,大团聚体含量增加 60% 以上;施加玉米秸秆与施腐熟有机肥相比,前者更有利于盐碱土大团聚体的形成;盐碱土中接种既分解秸秆又产胞外聚合物细菌对团聚体形成的促进作用优于单功能分解秸秆细菌(菌株 J2)或者分泌胞外聚合物细菌(菌株 DF-2);菌株 M6 对盐碱环境下玉米的出苗、生长存活及干物质的积累有促进作用.     

Foraging strategies of the external mycelium of the arbuscular mycorrhizal fungi Glomus intraradices and Scutellospora calospora

The responsiveness of the external mycelium of Glomus intraradices and Scutellospora calospora was tested in a multiple-choice experimental system in which mycelium encountered patches amended with nitrogen (N) or phosphorus (P), either alone or in combination with a host plant. We hypothesised that only AMF mycelium with sufficient supply of photosynthate from an actively growing host would respond to the amendments provided. Mycelium was allowed to grow either 11 or 21 weeks before we analysed hyphal proliferation in amended patches introduced in mesh bags that were not reached by roots but by foraging mycelium only. Hyphal length, the AMF signature fatty acid 16:1w5, and root colonisation in new host plant seedlings were used to measure AMF growth and resource allocation in the patches. Mycelium from both fungal strains was able to colonise new host roots and sand in all patches but S. calospora was overall more responsive to the amendments than G. intraradices. G. intraradices grew equally into all patches, including the unamended control, whereas S. calospora produced significantly more hyphal length in the patch containing a host plant than in the rest of the patches. Both strains showed lower hyphal growth at the second harvest and mycelium of G. intraradices lost almost entirely its capacity to develop new mycelium in all choices presented. Lipid measurements showed this fungus did not use storage lipids to exploit the patches. S. calospora mycelium had reduced growth and colonisation ability but still showed some growth in the patches at the second harvest. A reduction in the content of NLFA 16:1w5 from the first to the second harvest suggested that S. calospora mycelium likely used storage lipids to sustain proliferation in the patches. The results indicated that S. calospora was more active and used more resources for foraging than G. intraradices and that external mycelium foraging was maintained mainly with recently acquired plant carbon (C). This supported in general our hypothesis but showed as well that the two AMF strains had different strategies and resource allocation to forage. The overall low response of both AMF to the choices presented suggested that the responsiveness of mycelium searching freely in the substrate is lower than that observed in experimental systems in which the amendments have been placed in close contact with actively growing mycelium fronts in close vicinity with host roots.     

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.     

Factors affecting soil organic matter turnover in a mediterranean ecosystems from sierra de gador (Spain): An analytical approach

Abstract

Humus formations from forest and brushwood ecosystems on calcareous substrates in representative sites from Sierra de Gador in Southern Spain were analyzed. The humus composition, as determinated by three independent fractionation procedures, suggests that soil organic matter turnover is not related to the vegetation biotypes, but to a series of local factors among which climatic and soil characteristics predominate. Of the parameters studied, the amount of fulvic acids, and to a lesser extent, the relative proportion of soil perylenequinonic pigments (P‐type humic acids), reflected differences in altitude. The percentage of nitrogen (N) in humic acids was the parameter that was most informative of the characteristics of humus. This finding contrasts with the poorly differentiated patterns of the particulate organic matter fractions, whose evolution is probably independent of environmental factors because of physical protection by soil carbonates. The carbon dioxide (CO₂) release curves reproduced under laboratory conditions showed that the intrinsic biodegradability of soil organic matter does not reflect the nature of the plant residues in the sites sampled (170–2240 m), but is weakly dependent on climatic factors. We postulate that the calcium saturation, together with the active participation of inorganic colloids, force the convergent evolution of soil humus by compensating most of the effects of the climatic and biotic factors, and the historic influence of human activity in the area studied.     

The quality of exogenous organic matter: short-term effects on soil physical properties and soil organic matter fractions

We examined the short-term effect of five organic amendments and compared them to plots fertilized with inorganic fertilizer and unfertilized plots on aggregate stability and hydraulic conductivity, and on the OC and ON distribution in physically separated SOM fractions. After less than 1 year, the addition of organic amendments significantly increased ( P  <   0.01) the aggregate stability and hydraulic conductivity. The stability index ranged between 0.97 and 1.76 and the hydraulic conductivity between 1.23 and 2.80 × 10⁻³ m/s for the plots receiving organic amendments, compared with 0.34–0.43, and 0.42–0.64 × 10⁻³ m/s, respectively, for the unamended plots. There were significant differences between the organic amendments (P <  0.01), although these results were not unequivocal for both soil physical parameters. The total OC and ON content were significantly increased ( P  <   0.05) by only two applications of organic fertilizers: between 1.10 and 1.51% OC for the amended plots versus 0.98–1.08% for the unamended and between 0.092 and 0.131% ON versus 0.092–0.098% respectively. The amount of OC and ON in the free particulate organic matter fraction was also significantly increased ( P  <   0.05), but there were no significant differences ( P  <   0.05) in the OC and ON content in the POM occluded in micro-aggregates and in the silt + clay-sized organic matter fraction. The results showed that even in less than 1 year pronounced effects on soil physical properties and on the distribution of OC and ON in the SOM fractions occurred.