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Phosphorus (P) loss from paddy fields is a significant issue in sustainable rice production by threatening water environments. We aimed to examine the suitability of mycorrhiza-defective rice (non-mycorrhizal) and its mycorrhizal progenitor to evaluate P loss control via arbuscular mycorrhizal (AM) fungi. We also aimed to investigate the AM effect on P loss via runoff and leaching. We grew the two rice lines in microcosms with and without AM fungi, measured P loss via runoff and leaching before and after nitrogen–phosphorus–potassium fertilization, and quantified plant P content and soil P concentration after the final harvest. Mycorrhizal and non-mycorrhizal rice pair systems in the absence of AM fungi had similar plant, soil, runoff, and leachate P contents (except PO43−). In the presence of AM fungi, the concentrations of all P forms in runoff water and leachate in mycorrhizal rice were lower than those in nonmycorrhizal rice regardless of their solubility in water and availability to plants. The cumulative P loss from mycorrhizal systems was 10% less than that from their nonmycorrhizal counterparts. This mycorrhizal/non-mycorrhizal rice pair is an efficient experimental tool for research on the control of P loss from paddy fields with AM fungi. AM colonization contributes to the sustainability of rice production by decreasing P loss from paddy fields.

Little is known about how N and P co-limited ecosystems respond to single nutrient enrichment. This work assesses the susceptibility of above- and belowground ecosystem components and of their linkages in an N and P co-limited pasture to N- and P-enrichment. We tested if the plants’ responses can be explained by the concept of serially linked nutrients introduced by Agren (Ecol Lett 7:185–191, 2004). In this concept, the control of the growth rate by one nutrient is assumed to depend on the control of a different cellular process by another nutrient. We investigated the responses of shoot and root biomass and C:N:P stoichiometry of the grass Setaria sphacelata (Schumach.) to moderate N, P, and N + P application over 5 years. In addition, the effects of nutrient enrichment on soil nutrient pools, on arbuscular mycorrhizal fungi (AMF) as well as on microbial biomass, activity, and community structure (phospholipid fatty acids: PLFA) were tested. In order to evaluate the importance of different factors explaining microbial responses, we applied a likelihood-based information-theoretic approach. The application of N + P increased aboveground grass biomass (+61 %). Root biomass was stimulated by P-treatment (+45 %). Grass C:N:P stoichiometry responded by altering the P-uptake (P-treatment) or by translocating P from shoot to root (N-treatment). In particular, root C:N and C:P stoichiometry decreased in P- and in N-treatment. Extractable fractions of soil C, N, and P were significantly affected by nutrient enrichment. P application increased the biomass of Gram-positive bacteria (+22 %) and the abundance of AMF (+46 %), however, results of the IT-approach suggested indirect effects of nutrient enrichment on microbes. The responses of the N and P co-limited pasture to particular nutrient enrichment support the concept of serially linked nutrients. The present study provides evidence for the fundamental importance of P for controlling resource allocation of plants in responses to nutrient enrichment. Resource allocation of the grass rather than direct effects of nutrient additions drives changes in AMF, microbial biomass, community structure, and activity.

Aims The arbuscular mycorrhizal symbiosis is a widespread symbiosis in terrestrial biomes with functional implications for the ecology of both plants and soil organisms. We here asked whether phylogenetic host specificity (PHS) in arbuscular mycorrhizal (AM) fungal communities exists.

Soil aggregation is a crucial aspect of ecosystem functioning in terrestrial ecosystems. Arbuscular mycorrhizal fungi (AMF) play a key role in soil aggregate formation and stabilization. Here we quantitatively analyzed the importance of experimental settings as well as biotic and abiotic factors for the effectiveness of AMF to stabilize soil macroaggregates. We gathered 35 studies on AMF and soil aggregation and tested 13 predictor variables for their relevance with a boosted regression tree analysis and performed a meta-analysis, fitting individual random effects models for each variable. The overall mean effect of inoculation with AMF on soil aggregation was positive and predictor variable means were all in the range of beneficial effects. Pot studies and studies with sterilized sandy soil, near neutral soil pH, a pot size smaller than 2.5 kg and a duration between 2.2 and 5 months were more likely to result in stronger effects of AMF on soil aggregation than experiments in the field, with non-sterilized or fine textured soil or an acidic pH. This is the first study to quantitatively show that the effect of AMF inoculation on soil aggregation is positive and context dependent. Our findings can help to improve the use of this important ecosystem process, e.g. for inoculum application in restoration sites.

Aims To assess whether the yew roots, which are able to provide a very constant environment due to their long life-span, can maintain the original arbuscular mycorrhizal (AM) fungal community during yew population decline.

Arbuscular mycorrhizal (AM) fungi, a group of obligate symbionts of terrestrial plants, have a global distribution range. Yet, we lack concrete synthetic and empirical evidence that could reveal whether distinct ecological niches are distributed across Glomeromycota through determining linkages between environmental factors and the distribution of these taxa. We have modelled the probability of occurrence of Gigasporaceae and Acaulosporaceae as a function of candidate environmental factors. These families are among the most common but non-ubiquitous taxa in AM-driven ecosystems. We have constructed our database using studies with a global scope and carried out our analysis through a logistic regression approach. The probability of occurrence of Acaulosporacae increased in acidic environments and soils with high bulk density. By contrast, a key factor that affected probability of occurrence of Gigasporaceae was precipitation. Through the analysis of an unprecedentedly large amount of data we could infer that niche processes mediate occurrence of a group of fungi at scales broader than the local scale of the individual studies gathered in the analysed dataset. Knowledge of well-supported niche features could enhance discovery of new taxa of AM fungi, and would facilitate development of study designs with greater ecological realism.

Year of release of a cultivar reflects the agricultural and breeding practices of its time; we hypothesize that there are differences in mycorrhizal responsiveness of new high yielding and old crop plants and landraces. We evaluated the importance of the year of release on mycorrhizal responsiveness, arbuscular mycorrhizal (AM) fungal root colonization and P efficiency. We also analyzed the effect of experimental treatments, P efficiency (P acquisition and P utilization efficiency) and AM fungal root colonization on a potential mycorrhizal responsiveness trend for year of release. We conducted a meta-analysis on 39 publications working on 320 different crop plant genotypes. New cultivars were less intensely colonized but were more mycorrhiza-responsive (and possibly dependent) compared to ancestral genotypes. This trend was potentially influenced by the moderator variables density, pre-germination, plant, plant type and AMF species. AM root colonization was also important for the mycorrhizal responsiveness trend for year of release, but P efficiency was not. With the data available we could find no evidence that new crop plant genotypes lost their ability to respond to mycorrhiza due to agricultural and breeding practices.

The relative importance of roots and AM-fungi on soil physical processes was investigated by controlling the presence of roots and AM fungi in pot experiments using a mycorrhiza-defective tomato mutant and a wild-type tomato (Solanum lycopersicum L.). Root-Zone and Bulk Soil sections were established by splitting pots into two lengthwise halves using a nylon mesh that contained roots whilst allowing the free movement of fungal hyphae. Post-incubation microbial populations and fungal biomass were measured and related to soil stability, pore structure and water repellency. Unplanted controls consistently had the least fungal biomass, fatty acids, water-stable aggregates (WSA) and water repellency. Wild-type-planted treatments had significantly more WSA than mycorrhiza-defective treatments (P < 0.01). Fluctuations in water content induced by transpiration caused significant changes in soil pore structure, measured using high-resolution X-Ray computer tomography. Porosity and mean pore size increased in soil aggregates from planted treatments, which had larger more heterogeneous pores than those in the unplanted soils. AM fungi accentuated soil stability. However, changes were not linked to repellency and fungal biomass. The presence of plants, regardless of AM fungi, appears to have the greatest impact on increasing soil stability.

Despite the importance of arbuscular mycorrhizal fungi (AMF) to ecosystem processes, few experimental tools are available to quantify AMF contributions to process rates. In this study we examine the efficacy of an experimental system consisting of wildtype (WT) and different non-mycorrhizal (Myc−) genotype pairs of tomato (Solanum lycopersicum L.), specifically focusing on cv Micro-Tom. Two conditions necessary to make such a system useful were examined; (1) that the Myc− genotype(s) do not get colonized in a full soil AMF community background, while the WT does, and B) that there are no non-target effects of the Myc− phenotype on soil microbes. We assessed the second condition by growing Myc− genotypes and WT in non-mycorrhizal soil, monitoring plant growth (root, shoot biomass; root length; root diameter size distribution) and soil microbial community structure (PLFA analysis) as indicators of any changes in root tissue quality or rhizodeposition. All tested Myc− genotypes showed a drastically reduced colonization in mycorrhizal soil. However, in non-mycorrhizal soil, M161 had greater root biomass and M20 greater microbial biomass compared to WT. Only one of the Myc− mutants examined fully met the criteria. We conclude that the BC1/WT pair is a powerful experimental system and recommend caution when using Myc− mutants in mycorrhizal ecology.

Experiments suggest that biomass-derived black carbon (biochar) affects microbial populations and soil biogeochemistry. Both biochar and mycor- rhizal associations, ubiquitous symbioses in terrestrial ecosystems, are potentially important in various ecosystem services provided by soils, contributing to sustainable plant production, ecosystem restoration, and soil carbon sequestration and hence mitigation of global climate change. As both biochar and mycor- rhizal associations are subject to management, under- standing and exploiting interactions between them could be advantageous. Here we focus on biochar effects on mycorrhizal associations. After reviewing the experimental evidence for such effects, we critically examine hypotheses pertaining to four mechanisms by which biochar could influence mycorrhizal abundance and/or functioning. These mechanisms are (in decreas- ing order of currently available evidence supporting them): (a) alteration of soil physico-chemical proper- ties; (b) indirect effects on mycorrhizae through effects on other soil microbes; (c) plant-fungus signaling interference and detoxification of allelochemicals on biochar; and (d) provision of refugia from fungal grazers. We provide a roadmap for research aimed at testing these mechanistic hypotheses.

While several recent studies have described changes in microbial communities associated with exotic plant invasion, how arbuscular mycorrhizal fungi (AMF) communities respond to exotic plant invasion is not well known, despite the salient role of this group in plant interactions. Here, we use molecular methods (terminal restriction fragment length polymorphism analyses based on the large subunit of the rRNA gene) to examine AMF community structure in sites dominated by the invasive mycorrhizal forb, Centaurea maculosa Lam. (spotted knapweed), and in adjacent native grassland sites. Our results indicate that significant AMF community alteration occurs following C. maculosa invasion. Moreover, a significant reduction in the number of restriction fragment sizes was found for samples collected in C. maculosa-dominated areas, suggesting reduced AMF diversity. Extraradical hyphal lengths exhibited a significant, on average 24%, reduction in C. maculosa-versus native grass-dominated sites. As both AMF community composition and abundance were altered by C.maculosa invasion, these data are strongly suggestive of potential impacts on AMF-mediated ecosystem processes. Given that the composition of AMF communities has the potential to differentially influence different plant species, our results may have important implications for site restoration after weed invasion.

Controls on root colonization by arbuscular mycorrhizal fungi (AMF) include host nutrient status, identity of symbionts and soil physico-chemical properties. Here we show, in the field, that the subset of the AMF community colonizing the roots of a common grass species, Dactylis glomerata, was strongly controlled by neighboring roots of a different plant species, Centaurea maculosa, an invasive forb, thus adding a biological spatial component to controls on root colonization. Using an AMF-specific, 18s rDNA-based terminal restriction fragment length polymorphism (T-RFLP) analysis method, significant differences were found between AMF community fingerprints of samples derived from roots of grasses with (GCm) and without (G0) neighboring C. maculosa. There were also significant differences between samples derived from C. maculosa roots (Cmac) and both GCm and G0 roots. Sample ordination indicated three generally distinct groups consisting of Cmac, GCm and G0, with GCm samples being of intermediate distance between G0and Cmac. Our results indicate that, with the presence of C. maculosa, AMF communities of D. glomerata shift to reflect community composition associated with C. maculosa roots. These results highlight the importance of complex spatial distributions of AMF communities at the scale of a root system. An additional dimension to our study is that C. maculosa is an aggressively invasive plant in the intermountain West. Viewed in this light, these results suggest that pervasive influences of this plant on AMF communities, specifically in roots of its competitors, may represent a mechanism contributing to its invasive success. However, further work is clearly required to determine the extent to which AMF genotypic alteration by neighboring plants influences competitive relationships.

In order to more fully understand the basic biology of arbuscular mycorrhizal fungi (AMF), and their role in natural ecosystems, it is necessary to document seasonal changes of various aspects of the life history of these fungi. Due to their unique position at the root-soil interface, AMF have been described as `keystone mutualists' in ecosystems. Despite the importance of AMF in ecosystems, few studies exist that examine the seasonality of external hyphae and their exuded products (e.g. glomalin), the AMF variables directly related to ecosystem function through their contributions to soil aggregation. This study examined seasonal dynamics of several soil variables, with a specific interest in the seasonality of external hyphae and glomalin, a glycoprotein produced by AMF, which is correlated with soil aggregate stability. Here we measured glomalin concentrations and external AMF and non-AMF hyphal length, as well as soil moisture, percent fungal root colonization (AMF and non-AMF), and root length in soil in an intermountain grassland in western Montana over one growing season (13 time points). Of the glomalin pools and hyphal lengths measured, significant seasonal changes occurred for total glomalin (TG; 24.5% change), immunoreactive easily extractable glomalin (IREEG; 53.8% change), and AM hyphal length (107% change). Prior studies on glomalin in natural systems have not considered seasonal effects. The small seasonal change in glomalin pools lends further support to the hypothesis that glomalin is relatively stable in soils, and suggests that one-time sampling may be sufficient to satisfactorily capture this response variable. However, the generality of this observation has yet to be tested in a wider range of ecosystems.

Glomalin is a soil proteinaceous substance produced by arbuscular mycorrhizal fungi. Most of the information available concerning this protein has been collected in relation to its role in soil aggregation. In this study, we explored the distribution of glomalin across soil horizons, decomposition of glomalin, and relationship with soil C and N in an agricultural field, a native forest, and an afforested system. Glomalin was present in A, B, and C horizons in decreasing concentrations. Land-use type significantly affected glomalin concentrations (mg cm −3 ), with native forest soils having the highest concentrations of the three land-use types in both A and B horizons. In terms of glomalin stocks (Mg ha −1 ), calculated based on corrected horizon weights, the agricultural area was significantly lower than both afforested and native forest areas. As measured after a 413 day laboratory soil incubation, glomalin was least persistent in the A horizon of the afforested area.. In agricultural soils and native soils, ca. 50% of glomalin was still remaining after this incubation, indicating that some glomalin may be in the slow or recalcitrant soil C fraction. Comparison of glomalin decomposition with CO2-C respired during incubation indicates that glomalin makes a large contribution to active soil organic C pools. Soil C and N were highly correlated with glomalin across all soils and within each land-use type, indicating that glomalin may be under similar controls as soil C. Our results show that glomalin may be useful as an indicator of land-use change effects on deciduous forest soils.

The importance of arbuscular mycorrhizae (AM) in plant and ecosystem responses to global changes, e.g. elevated atmospheric CO2, is widely acknowledged. Frequently, increases in AM root colonization occur in response to increased CO2, but also the lack of significant changes has been reported. The goal of this study was to test whether arbuscular mycorrhizae (root colonization and composition of root colonization) respond to plants grown in elevated CO2 as a function of soil depth. We grew Bromus hordeaceus L. and Lotus wrangelianus Fischer & C. Meyer monocultures in large pots with a synthetic serpentine soil profile for 4 yr in an experiment, in which CO2 concentration was crossed factorially with NPK fertilization. When analyzing root infection separately for topsoil (0–15 cm) and subsoil (15–45 cm), we found large (e.g., about 5-fold) increases of AM fungal root colonization in the subsoil in response to CO2, but no significant changes in the corresponding topsoil of Bromus. Only the coarse endophyte AM fungi, not the fine endophyte AM fungi, were responsible for the observed increase in the bottom soil layer, indicating a depth-dependent shift in the AM community colonizing the roots, even at this coarse morphological level. Other response variables also had significant soil layer * CO2 interaction terms. The subsoil response would have been hidden in an unstratified assessment of the total root system, since most of the root length was concentrated in the top soil layer. The increased presence of mycorrhizae in roots deeper in the soil should be considered in sampling protocols, as it may be indicative of changed patterns of nutrient acquisition and carbon sequestration.

Soil aggregation and soil structure are fundamental properties of natural and managed ecosystems. However, most of our knowledge on the role of plant species in soil aggregation is derived from work in agroecosystems or with agriculturally important plants. Here we examined the effects of five plant species on soil aggregate water stability. The five species (three grasses, one forb, and a legume) were from the same natural grassland, and were grown in monoculture plots in the field. Our first goal was to test if productivity-related or species-specific factors would prevail in determining soil aggregation. We also tested what the relative importance of the soil protein glomalin (produced by arbuscular mycorrhizal fungi, AMF) in soil aggregation is, compared to other factors, including AMF hyphal and root length and percent plant cover. We found significant differences in soil aggregate water stability (1–2 mm size class) for the five plant species examined, and corresponding differences in plant cover, root weight and length, AMF soil hyphal length, and glomalin concentrations. A structural equation modeling approach (path analysis) was used to distinguish direct from indirect effects of factors on soil aggregation based on covariance structures. Root length, soil glomalin, and percent cover contributed equally strong paths to water-stable aggregation. The direct effect of glomalin was much stronger than the direct effect of AMF hyphae themselves, suggesting that this protein is involved in a very important hypha-mediated mechanism of soil aggregate stabilization, at least for the 1–2-mm size class of aggregates.

The origins and composition of soil organic matter (SOM) are still largely uncertain. Arbuscular mycorrhizal fungi (AMF) are recognized as indirect contributors through their influence on soil aggregation, plant physiology, and plant community composition. Here we present evidence that AMF can also make large, direct contributions to SOM. Glomalin, a recently discovered glycoprotein produced by AMF hyphae, was detected in tropical soils in concentrations of over 60 mg cm−3. Along a chronosequence of soils spanning ages from 300 to 4.1 Mio years, a pattern of glomalin concentrations is consistent with the hypothesis that this protein accumulates in soil. Carbon dating of glomalin indicated turnover at time scales of several years to decades, much longer than the turnover of AMF hyphae (which is assumed to be on the order of days to weeks). This suggests that contributions of mycorrhizae to soil carbon storage based on hyphal biomass in soil and roots may be an underestimate. The amount of C and N in glomalin represented a sizeable amount (ca. 4–5%) of total soil C and N in the oldest soils. Our results thus indicate that microbial (fungal) carbon that is not derived from above- or below-ground litter can make a significant contribution to soil carbon and nitrogen pools and can far exceed the contributions of soil microbial biomass (ranging from 0.08 to 0.2% of total C for the oldest soils).