Consumption of mycorrhizal and saprophytic fungi by Collembola in grassland soils

Although soil-dwelling Collembola can influence plant growth and nutrient cycling, their specific role in soil food webs is poorly understood. Soil-free microcosm studies suggest that Collembola are primarily fungivores where they feed preferentially on saprophytic fungi (SF) over other fungal types. We directly assessed collembolan consumption of arbuscular mycorrhizal fungi (AMF) and SF using plant-soil mesocosms and natural abundance stable carbon isotope techniques. Mycorrhizal Andropogon gerardii (C₄ grass) seedlings were placed in pots containing Collembola and soil from a C₃ plant dominated site, while mycorrhizal Pascopyrum smithii (C₃ grass) seedlings were placed in pots with Collembola and soil collected at a C₄ plant dominated site. After 6 weeks, collembolans assimilated carbon derived from C₃ and C₄ sources in both A. gerardii and P. smithii treatments. Comparing Collembola isotope values in AMF vs. AMF-suppressed treatments, our data show that both AMF and SF were consumed in these experimental soil environments.     

Arbuscular mycorrhizal fungi are directly and indirectly affected by glyphosate application

Glyphosate is a systemic non-selective herbicide, the most widely used in the world. Alongside with its use in agricultural and forestry systems, this herbicide is used in grasslands in late summer with the aim of promoting winter species with the consequent increase in stocking rate. However, its effects on non-target organisms, such as arbuscular mycorrhizal fungi (AMF), are unclear. Arbuscular mycorrhizal fungi (AMF) colonize the root of more than 80% of terrestrial plants, improving their growth and survival, and therefore playing a key role in ecosystem structure and function. The aim of this work was to investigate the possible pathways through which glyphosate application affects AMF spores viability and root colonization in grassland communities. Our hypothesis is that glyphosate application can damage AMF directly (through contact with spores and external hyphae) or indirectly through the changes it generates on host plants. The experiment had a factorial array with three factors: (1) plant species, at two levels (Paspalum dilatatum and Lotus tenuis), (2) doses of glyphosate, at three levels (0 l ha⁻¹, 0.8 l ha⁻¹ and 3 l ha ⁻¹), and (3) application site, at two levels: soil (direct pathway) and plant foliage (indirect pathway). Spore viability was reduced even under the lowest glyphosate rate, but only when it was applied on the soil. Total root colonization for both species was similarly decreased when glyphosate was applied to plant foliage or on soil, with no difference between 0.8 and 3 l ha⁻¹. The number of arbuscules was 20% lower when glyphosate was applied on plant foliage, than when it was applied on the soil. Our findings illustrate that glyphosate application negatively affects AMF functionality in grasslands, due to different causes depending on the herbicide application site. While, under field conditions, the occurrence of direct and/or indirect pathways will depend on the plant cover at the time of glyphosate application, the consequences of this practice on the plant community structure will vary with the mycorrhizal dependence of the species composition regardless of the pathway involved.     

Interaction between legume and arbuscular mycorrhizal fungi identity alters the competitive ability of warm-season grass species in a grassland community

Arbuscular mycorrhizal fungi (AMF) and N₂-fixing legumes can alter the community structure of grasses. However, the effect of AMF, N₂-fixing legumes, and their interaction on the dynamics of prairie grass communities remains unclear. The aim of this study was to clarify the influence of two AMF (Glomus cubense and Glomus sp.) and two legumes (Medicago sativa and Dalea purpurea) on the competitive relationship between three native cool-season (Elymus canadensis, Elymus trachycaulus ssp. subsecundus, and Elymus lanceolatus ssp. lanceolatus) and two native warm-season species of grasses (Schizachyrium scoparium and Bouteloua gracilis). Results show that AMF and legumes altered the community structure of the grasses. G. cubense favoured the productivity of warm-season B. gracilis when growing with M. sativa. This might be related to a negative impact of G. cubense on the nitrogen-fixing activity of M. sativa and to a lower N-use efficiency of E. canadensis and E. lanceolatus ssp. lanceolatus under competition. This suggested an increased ability of B. gracilis to use the available N resource as affected by more competitive species, whereas Glomus sp. reduced the competitive ability of this grass when associated with M. sativa. The decrease in B. gracilis biomass was thus likely caused by enhancement of P uptake by M. sativa over this grass. Glomus sp. was beneficial to S. scoparium, another warm-season species, in the absence of legumes, and this may be attributed to improved P-use efficiency of this grass under competition with cool season-grasses. In contrast, AMF and legumes were not beneficial for the cool season grasses. G. cubense depressed the growth of E. trachycaulus ssp. subsecundus, and M. sativa decreased nutrient uptake by cool-season native grasses. This study shows that beneficial effect of the arbuscular mycorrhizal symbiosis on the coexistence of warm-season grasses with more competitive cool-season grasses depends on the identity of the AMF symbiont, the presence of legume species, and nitrogen resource availability that was affected by the most competitive species or P-use efficiency of warm season species.     

Interaction between earthworms and arbuscular mycorrhizal fungi on the degradation of oxytetracycline in soils

The interactive impact of earthworms (Eisenia fetida) and arbuscular mycorrhizal fungi (Rhizophagus intraradices, AM fungi) on the degradation of oxytetracycline (OTC) in soils was studied under greenhouse conditions. Treatments included maize plants inoculated vs. not inoculated with AM fungi and treated with or without earthworms at low (1 mg kg⁻¹ soil DM) or high (100 mg kg⁻¹ soil DM) OTC rates. The root colonization rate, the hyphal density of mycorrhizae, the residual OTC concentration in soils, catalase, dehydrogenase, urease, soil microbial biomass C, Shannon–Wiener index (H) for microbial communities from T-RFLP profiles were measured at harvest. The results indicated that earthworms and AM fungi would individually or interactively enhance OTC decomposition and significantly decreased the residual OTC concentration at both high and low OTC rates. Both earthworms and AM fungi could promote the degradation of OTC by increasing soil microbial biomass C at both high and low OTC rates. The effect of soil enzyme activity and soil microbial diversity on OTC decomposition was different between high and low OTC rates. Hyphomicrobium and Bacillus cereus were dominant bacteria, and Thielavia and Chaetomium were dominant phyla of fungi at all occasions. Earthworm activity stimulated the growth of Hyphomicrobium and Thielavia, while AM fungi may stimulate B. cereus, Thielavia and Chaetomium, resulting in greater OTC decomposition. The interaction between earthworms and AM fungi in affecting the degradation of OTC may be attributed to different mechanisms, depending on soil microbial biomass, function (enzyme activity) and communities (the abundance of Hyphomicrobium, B. cereus, Thielavia and Chaetomium) in the soil.     

Belowground responses by AM fungi and animal trophic groups to repeated defoliation in an experimental grassland community

We tested a hypothesis that the effects of defoliation on plants and soil organisms vary with the number of successive defoliations. We established a 23-week greenhouse experiment using replicated grassland microcosms that were composed of three plant species, Trifolium repens, Plantago lanceolata and Phleum pratense, growing together in grassland soil with a diverse soil community. The experiment consisted of two treatment factors-defoliation and harvest time-in a fully factorial design. The defoliation treatment had two levels, i.e. no trimming and trimming of plants every 2 weeks, and the harvest time five levels, i.e. harvests after 1-3, 5 and 7 trimmings. Shoot production (trimmed plus harvested shoot mass), harvested shoot and root mass and root N and C concentrations increased with time but were reduced by defoliation. Colonization rates of arbuscular mycorrhizal (AM) fungi decreased with time in T. repens roots but were enhanced by defoliation, whereas AM colonization rates in P. pratense roots were not affected by harvest time or defoliation. The abundance of bacterivorous and fungivorous nematodes decreased and that of herbivorous and predatory nematodes increased with time, while the abundance of omnivorous nematodes and detritivorous enchytraeids varied in time without a linear trend. Defoliation had no effect on fungivores and predators but increased the abundance of bacterivores. Defoliation also increased the abundance of herbivores, omnivores and detritivores after 2 trimmings and that of omnivores and detritivores after 5 trimmings, but had a negative effect on omnivores after 3 trimmings and on herbivores after 7 trimmings. Among nematode genera, some deviation from the trophic group responses existed: for instance, defoliation reduced the abundance of bacterivorous Acrobeloides spp. and did not affect the abundance of herbivorous Filenchus spp. and Paratylenchus spp. Our results show that the effects of defoliation on plants, AM fungi and some soil animal trophic groups may remain constant all the way through several defoliations, whereas other animal trophic groups may have different and even opposite responses to defoliation depending on the length of the defoliation period before monitoring. This shows how separate studies with defoliation periods of different length can produce contradictory results of the effects of defoliation on the abundance of soil animals.     

The significant contribution of mycorrhizal fungi and earthworms to maize protection and phytoremediation in Cd-polluted soils

Mycorrhizal fungi and earthworms can individually or interactively influence plant growth and heavy metal uptake. The influence of earthworms and arbuscular mycorrhizal (AM) fungi either alone or in combination on maize (Zea mays L.) growth and cadmium (Cd) uptake was investigated in a calcareous soil artificially spiked with Cd. Soils were contaminated with Cd (10 and 20 mg Cd kg⁻¹), inoculated or un-inoculated with the epigeic earthworm Lumbricus rubellus and two AM fungal species (Rhizophagus irregularis and Funneliformis mosseae) for two months of growth under greenhouse conditions. Generally, earthworms alone increased both shoot P uptake and biomass but decreased shoot Cd concentration and root Cd uptake. AM fungi individually often increased total maize P uptake, declined shoot Cd concentration, and consequently produced higher total biomass. However, R. irregularis enhanced shoot Cd uptake at low Cd level and root Cd uptake at high Cd level. In plants inoculated with F. mosseae species, earthworms increased shoot biomass and Cd uptake, decreased root biomass and Cd uptake at all Cd levels, and increased shoot Cd concentration at low Cd level. In plants colonized by R. irregularis species, however, earthworm addition decreased maize biomass only at high Cd level and root Cd concentration and total maize Cd uptake at both Cd levels. Earthworm activity decreased Cd transfer from the soil to maize roots at low Cd level, but this was counterbalanced in the presence of F. mosseae. Mycorrhizal symbiosis significantly reduced the transfer of Cd from roots to shoots, independence of earthworm effect. Overall, it is concluded that L. rubellus and AM fungi, in particular F. mosseae isolate, improved maize tolerance to Cd toxicity both individually and interactively by increasing plant growth and P nutrition, and restricting Cd transfer to the aboveground biomass. Consequently, the single and interactive effects of the two soil organisms might potentially be important not only in protecting maize plants against Cd toxicity, but also in Cd phytostabilization in soils polluted by this highly toxic metal.     

Soil microbial community composition as affected by restoration practices in California grassland

Agricultural practices have strong impacts on soil microbes including both the indices related to biomass and activity as well as those related to community composition. In a grassland restoration project in California, where native perennial bunchgrasses were introduced into non-native annual grassland after a period of intensive tillage, weeding, and herbicide use to reduce the annual seed bank, microbial community composition was investigated. Three treatments were compared: annual grassland, bare soil fallow, and restored perennial grassland. Soil profiles down to 80 cm in depth were investigated in four separate layers (0-15, 15-30, 30-60, and 60-80 cm) using both phospholipid ester-linked fatty acid (PLFAs) and ergosterol as biomarkers in addition to microbial biomass C by fumigation extraction. PLFA fingerprinting showed much stronger differences between the tilled bare fallow treatment vs. grasslands, compared to fewer differences between restored perennial grassland and annual grassland. The presence or absence of plants over several years clearly distinguished microbial communities. Microbial communities in lower soil layers were little affected by management practices. Regardless of treatment, soil depth caused a strong gradient of changing habitat conditions, which was reflected in Canonical Correspondence Analysis of PLFAs. Fungal organisms were associated with the presence of plants and/or litter since the total amount and the relative proportion of fungal markers were reduced in the tilled bare fallow and in lower layers of the grassland treatments. Total PLFA and soil microbial biomass were highly correlated, and fungal PLFA biomarkers showed strong correlations to ergosterol content. In conclusion, microbial communities are resilient to the grassland restoration process, but do not reflect the change in plant species composition that occurred after planting native bunchgrasses.     

Potential of earthworms,ants, millipedes,and termites for dissemination of vesicular-arbuscular mycorrhizal fungi in soil

We studied the effects of earthworms, termites, ants, and millipedes on the dissemination of vesciulararbuscular mycorrhizal (VAM) propagules. Earthworm (Lumbricus terrestris L.) casts collected from the garden were air-dried, stored, and examined for the presence of VAM fungi by inoculating the cast material onto onion plants grown in sterilized soil. VAM propagules survived for a period of 12 months. The results showed that earth-worms can contribute to the dissemination of VAM propagules. Nests of ants (Camponotus compressus Fabr.) collected from different locations were examined for the presence of VAM propagules. They harboured up to 790 infective propagules g^-1 of nest, showing that ants can be potential vectors in disseminating VAM fungi. The faecal pellets of the millipede (Phyllogonostreptus nigrolabiatus Newport) collected from the rearing tank were tested for VAM colonization. Though VAM propagules were present in the fresh faecal pellets, they lost their viability after 4 days of storage. Termitaria samples of both mound-building and subterranean termites were also examined for VAM propagules but those found were non-viable.     

Bacteria and protozoa in soil microhabitats as affected by earthworms

The effects of incorporation of elm leaves (Ulmus glabra) into an agricultural sandy loam soil by earthworms (Lumbricus festivus) on the bacterial and protozoan populations were investigated. Three model systems consisting of soil, soil with leaves, and soil with leaves and earthworms, respectively, were compared. The total, viable, and culturable number of bacteria, the metabolic potentials of bacterial populations, and the number of protozoa and nematodes were determined in soil size fractions. Significant differences between soil fractions were shown by all assays. The highest number of microorganisms was found in microaggregates of 2–53 μm and the lowest in the <0.2μm fraction. A major part of the bacteria in the latter fraction was viable, but non-culturable, while a relatively higher number of culturable bacteria was found in the macroaggregates. The number of colony-forming units and 5-cyano-2,3-ditolyl tetrazolim chloride (CTC)-reducing bacteria explained a major part of the variation in the number of protozoa. High protozoan activity and predation thus coincided with high bacterial activity. In soil with elm leaves, fungal growth is assumed to inhibit bacterial and protozoan activity. In soil with elm leaves and earthworms, earthworm activity led to increased culturability of bacteria, activity of protozoa, number of nematodes, changed metabolic potentials of the bacteria, and decreased differences in metabolic potentials between bacterial populations in the soil fractions. The effects of earthworms can be mediated by mechanical mixing of the soil constituents and incorporation of organic matter into the soil, but as the earthworms have only consumed a minor part of the soil, priming effects are believed partly to explain the increased microbial activity. Received: 7 January 1996     

长期不同施肥对棕壤AM真菌群落结构的影响

【目的】 探明长期不同施肥条件下玉米–大豆轮作棕壤丛枝菌根 (Arbuscular mycorrhizal, AM) 真菌群落结构的变化及其影响因素。 【方法】 以沈阳农业大学棕壤肥料长期 (38年) 定位试验耕层土壤 (0—20 cm) 为材料,于2016年4月选取其中6个施肥处理:1) 不施肥 (CK);2) 单施化学氮肥 (N);3) 施用化学氮磷肥 (NP);4) 施用化学氮磷钾肥 (NPK);5) 单施有机肥 (M);6) 有机肥和化学氮磷肥配施 (MNP)。采用PCR-DGGE和克隆测序,分析了棕壤AM真菌群落结构,并结合环境因素进行冗余分析和典型对应分析。 【结果】 施用有机肥处理土壤有机碳 (TOC)、全氮 (TN)、全磷 (TP)、全钾 (TK)、碱解氮 (AHN)、有效磷 (AP)、速效钾 (AK)、可溶性有机碳 (DOC) 含量显著高于单施化肥和不施肥处理,且趋势表现为有机肥处理 > 化肥处理 > CK;与CK相比,单施化肥处理显著降低了土壤pH值,施用有机肥处理显著提高了土壤pH值。长期施肥显著改变了土壤中的AM真菌孢子密度,施用有机肥处理的孢子密度显著高于单施化肥处理和不施肥处理。冗余分析结果表明,土壤pH与AM真菌多样性指数和均匀度均呈显著负相关,而孢子密度与AP、TK、AHN呈显著正相关,AM真菌多样性指数与孢子密度则不相关。聚类分析表明长期不同施肥将棕壤中AM真菌分为两大类群,分别为不施肥区和施肥区,其相似度仅为42%。通过割胶克隆测序得出从土壤样品中分离的AM真菌种群主要为球囊霉菌,典型对应分析表明AP、AK、TOC、NH ₄+-N显著影响AM真菌的群落组成。 【结论】 长期定位施肥改变了棕壤的理化性质,从而对AM真菌的群落结构产生了显著影响。      

The combined effects of earthworms and arbuscular mycorrhizal fungi on microbial biomass and enzyme activities in a calcareous soil spiked with cadmium

Earthworms and arbuscular mycorrhizal fungi (AMF) are known to independently affect soil microbial and biochemical properties, in particular soil microbial biomass (SMB) and enzymes. However, less information is available about their interactive effects, particularly in soils contaminated with heavy metals such as cadmium (Cd). The amount of soil microbial biomass C (MBC), the rate of soil respiration (SRR) and the activities of urease and alkaline phosphatase (ALP) were measured in a calcareous soil artificially spiked with Cd (10 and 20 mg Cd kg⁻¹), inoculated with earthworm (Lumbricus rubellus L.), and AMF (Glomus intraradices and Glomus mosseae species) under maize (Zea mays L.) crop for 60 days. Results showed that the quantity of MBC, SRR and enzyme activities decreased with increasing Cd levels as a result of the elevated exchangeable Cd concentration. Earthworm addition increased soil exchangeable Cd levels, while AMF and their interaction with earthworms had no influence on this fraction of Cd. Earthworm activity resulted in no change in soil MBC, while inoculation with both AMF species significantly enhanced soil MBC contents. However, the presence of earthworms lowered soil MBC when inoculated with G. mosseae fungi, showing an interaction between the two organisms. Soil enzyme activities and SRR values tended to increase considerably with the inoculation of both earthworms and AMF. Nevertheless, earthworm activity did not affect ALP activity when inoculated with G. mosseae fungi, while the presence of earthworm enhanced urease activity only with G. intraradices species. The increases in enzyme activities and SRR were better ascribed to changes in soil organic carbon (OC), MBC and dissolved organic carbon (DOC) contents. In summary, results demonstrated that the influence of earthworms alone on Cd availability is more important than that of AMF in Cd-polluted soils; and that the interaction effects between these organisms on soil microorganism are much more important than on Cd availability. Thus, the presence of both earthworms and AMF could alleviate Cd effects on soil microbial life.     

Decomposer community complexity affects plant competition in a model early successional grassland community

The effects of a simple decomposer community, consisting of collembola, enchytraeids and earthworms on the performance of plants in a model Mediterranean early successional grassland community were investigated. Interactions of plant functional groups and species within the soil decomposer community resulted in significant shifts of plant competitive strength and therefore are likely to affect the rate of the successional development of plant communities.Depending on the functional role of the decomposer taxa involved, either legumes, or non-leguminous forbs were more negatively affected. Therefore, increasing species numbers with in the decomposer community gradually reduced the relative contribution of forbs (legumes and non-leguminous forbs), although fundamentally different mechanisms were responsible for these effects. Surprisingly, not the biomass-dominant annelid taxa, but collembola had the strongest effects on plant performance. The grass-to-forb ratio, an indicator of successional change, shifted from 3 to 4 in presence of collembola, suggesting that indirect effects on microbial symbionts of plants were more important than classic decomposer effects via increased and more constant nutrient availability to plants. Total plant productivity, however, was not affected, since grasses gained competitive advantage and compensated with increased growth for the negative animal effects on forb performance. Our results highlight the importance of specific functional groups among decomposers for structuring grassland plant communities. However, great plasticity in the plant community through negative covariance between forbs and grasses, partly compensated for decomposer effects at the plant canopy level.     

Uptake and partitioning of 32P and 65Zn by white clover as affected by eleven isolates of mycorrhizal fungi

We compared the effect of eleven mycorrhizal fungi populations (Glomus spp.) from two different climatic regions (Iran vs. Switzerland) on the uptake and partitioning of ³²P and ⁶⁵Zn by white clover (Trifolium repens L. var. Milkanova). Fungi significantly differed in the colonization of roots with hyphae, vesicles or arbuscules and in their effect on the uptake of ³²P or ⁶⁵Zn by white clover. Fungi also differed in their effects on the percentages of ³²P or ⁶⁵Zn transported to the plant tops. There was no relationship between the uptake of ³²P and that of ⁶⁵Zn in plants colonized by different fungi. Isolates Z7 and Q12 (from Iran) were superior to Swiss isolates for their effect on the uptake of zinc (Zn) by white clover. Among the Swiss isolates, that from Langwiese and Hausweid resulted in the highest and lowest partitioning of both ³²P and ⁶⁵Zn to the plant tops, respectively. We conclude that the effectiveness of vesicular arbuscular mycorrhizae in uptake of phosphorus (P) and Zn may be nutrient specific and that some fungal isolates may also affect the relative partitioning of P and Zn between plant roots and tops.     

Inoculating maize fields with earthworms (Aporrectodea trapezoides) and an arbuscular mycorrhizal fungus (Rhizophagus intraradices) improves mycorrhizal community structure and increases plant nutrient uptake

Earthworms and arbuscular mycorrhizal fungi (AMF) are important macrofauna and microorganisms of the rhizosphere. The effect of the inoculation of soil with earthworms (Aporrectodea trapezoides) and mycorrhiza (Rhizophagus intraradices) on the community structure of mycorrhizal fungi and plant nutrient uptake was determined with split plots in a maize field. Maize plants were inoculated or not inoculated with AMF, each treated with or without earthworms. Wheat straw was added as a feed source for earthworms. Inoculating AMF significantly increased maize yield (p?<?0.05), and the yield was further enhanced by the addition of earthworms. Alkaline phosphomonoesterase activities, soil microbial biomass carbon (SMBC) and nitrogen (SMBN) increased with the addition of both earthworms and AMF. Soil inorganic N and available K were positively affected by earthworms, while available P showed a negative relationship with AMF. Treatment with both AMF and earthworms increased shoot and root biomass as well as their N and P uptake by affecting soil phosphomonoesterase and urease activities, SMBC, SMBN, and the content of available nutrients in soil. The applied fungal inoculants were successfully traced by polymerase chain reaction with novel primers (AML1 and AML2) which target the small subunit rRNA gene. The amplicons were classified by restriction fragment length polymorphism and sequencing. Moreover, field inoculation with inocula of non-native isolates of R. intraradices appeared to have stimulated root colonization and yield of maize. Adding earthworms might influence native AMF community, and the corresponding abundance increased after earthworms were inoculated, which has positive effects on maize growth.     

Arbuscular mycorrhizal hyphae in grassland select for a diverse and abundant hyphospheric bacterial community involved in sulfonate desulfurization

Hyphae of symbiotic arbuscular mycorrhizal (AM) fungi extend into the soil, affecting the hyphosphere and interact with beneficial soil bacteria. This study aimed to elucidate differences in hyphosphere, hyphoplane and bulk soil bacterial communities and their role in mobilization of sulfonate-sulfur. Abundances of cultivable hyphosphere and hyphoplane bacteria were significantly increased over bulk soil. Cultivation independent fingerprinting revealed significantly different community structures of both hyphosphere and hyphoplane bacteria, fungi and AM fungi over bulk soil. However, cultivation dependent and independent analysis did not identify a difference between bacterial hyphoplane and hyphosphere (hyphospheric) communities. Isolated bacteria capable of aromatic sulfonate desulfurization were almost exclusively of hyphospheric origin. Members of the hyphospheric Gammaproteobacteria and Actinobacteria were found to possess marker gene asfA for aromatic sulfonate desulfurization and hrcR for attachment to fungal hyphae with a type III secretion system, that were not detected in bulk soil. These findings suggest that AM hyphae host a distinct population of sulfonate desulfurizing bacteria putatively capable of hyphal attachment with potential to increase plant sulfur supply.     

Contribution of exudates,arbuscular mycorrhizal fungi and litter depositions to the rhizosphere priming effect induced by grassland species

The presence of plants induces strong accelerations in soil organic matter (SOM) mineralization by stimulating soil microbial activity – a phenomenon known as the rhizosphere priming effect (RPE). The RPE could be induced by several mechanisms including root exudates, arbuscular mycorrhizal fungi (AMF) and root litter. However the contribution of each of these to rhizosphere priming is unknown due to the complexity involved in studying rhizospheric processes. In order to determine the role of each of these mechanisms, we incubated soils enclosed in nylon meshes that were permeable to exudates, or exudates & AMF or exudates, AMF and roots under three grassland plant species grown on sand. Plants were continuously labeled with ¹³C depleted CO₂ that allowed distinguishing plant-derived CO₂ from soil-derived CO₂. We show that root exudation was the main way by which plants induced RPE (58–96% of total RPE) followed by root litter. AMF did not contribute to rhizosphere priming under the two species that were significantly colonized by them i.e. Poa trivialis and Trifolium repens. Root exudates and root litter differed with respect to their mechanism of inducing RPE. Exudates induced RPE without increasing microbial biomass whereas root litter increased microbial biomass and raised the RPE mediating saprophytic fungi. The RPE efficiency (RPE/unit plant-C assimilated into microbes) was 3–7 times higher for exudates than for root litter. This efficiency of exudates is explained by a microbial allocation of fresh carbon to mineralization activity rather than to growth. These results suggest that root exudation is the main way by which plants stimulated mineralization of soil organic matter. Moreover, the plants through their exudates not only provide energy to soil microorganisms but also seem to control the way the energy is used in order to maximize soil organic matter mineralization and drive their own nutrient supply.     

The community structure of arbuscular mycorrhizal fungi in roots of maize grown in a 50-year monoculture

We assessed the diversity of arbuscular mycorrhizal (AM) fungi colonizing maize roots grown in a long-term monoculture experiment established at Martonvásár, Hungary, 50 years ago to understand the effect of this extremely long monoculture on the community structure of these organisms. Mycorrhizal colonization of root samples was analyzed by PCR amplification and sequencing of partial ribosomal small subunit DNA fragments of fungal origin. Of the 257 sequences recovered, 203 belonged to Glomeromycota AM fungi. Phylogenetic analysis assigned the Glomeromycota sequences into 22 operational taxonomic units belonging to three families including Archaeosporaceae, Glomeraceae, and Paraglomeraceae. In agreement with previous reports, Glomus group A fungi dominated the arbuscular mycorrhizal fungi community of maize, but we found a relatively high richness of phylotypes within this group even after such an extreme and durable reduction of host plant diversity. Agricultural practices, including mineral fertilization and incorporating stalk residues, significantly affected the diversity within Glomus group A.     

Organic nitrogen uptake by arbuscular mycorrhizal fungi in a boreal forest

The breakdown of organic nitrogen in soil is a potential rate-limiting step in nitrogen cycling. Arbuscular mycorrhizal (AM) fungi are root symbionts that might improve the ability of plants to compete for organic nitrogen products against other decomposer microbes. However, AM uptake of organic nitrogen, especially in natural systems, has traditionally been difficult to test. We developed a novel quantitative nanotechnological technique to determine in situ that organic nitrogen uptake by AM fungi can occur to a greater extent than has previously been assumed. Specifically, we found that AM fungi acquired recalcitrant and labile forms of organic nitrogen. Moreover, N enrichment of soil reduced plot-scale uptake of these compounds. Since most plants host AM fungi, AM use of organic nitrogen could widely influence plant productivity, especially where N availability is relatively low.     

Beneficial effects of earthworms and arbuscular mycorrhizal fungi on establishment of leguminous trees on Pb/Zn mine tailings

Planting trees to stabilize metalliferous mine tailings is a widely used form of land reclamation although substantial soil amendment is invariably required, both to improve the physico-chemical status of the tailings and to ameliorate toxicity prior to planting. Here, we report a glasshouse study of the combined effects of burrowing earthworms (Pheretima guillelmi) and arbuscular mycorrhizal fungi (Glomus spp., AMF) on establishment of a naturally invasive, woody, nitrogen-fixing legume, Leucaena leucocephala, on topsoil-amended Pb/Zn mine tailings. AMF provided the most effective preliminary inoculant, improving N, P and K uptake, but earthworms had more influence improving N nutrition. In most cases, the combined effects of AMF and earthworms were additive and proved to be beneficial to plant growth, plant nutrition and for protection against uptake of toxic metals. AMF influenced metal uptake more than earthworms, but together they reduced mobility of Pb and Zn in soil by as much as 25%. Some minor but significant negative interactions were also evident; for example, earthworms enhanced soil microbial activity but inhibited the beneficial effects of AMF on N₂-fixation. We argue that increased attention to ecological interactions in soil could reduce costs and improve the efficacy of restoring a vegetation cover to land impacted by contaminated spoils.