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Soil pH is probably the most important variable explaining bacterial richness and community composition locally as well as globally. In contrast, pH effects on fungi appear to be less pronounced, but also less studied. Here we analyze the community responses of bacteria and fungi in parallel over a local extreme pH gradient ranging from 4 to 8. We established the pH gradient by applying strongly alkaline wood ash in dosages of 0, 3, 9, 15, 30, and 90 t ha–1 to replicated plots in a Picea abies plantation and assessed bacterial and fungal community composition using high throughput amplicon sequencing 1 year after ash application. At the same time, the experiment investigated if returning wood ash to plantation forests pose any immediate threats for the microbial communities. Among the measured environmental parameters, pH was by far the major driver of the microbial communities, however, bacterial and fungal communities responded differently to the pH increment. Whereas both bacterial and fungal communities showed directional changes correlated with the wood ash-induced increase in pH, the bacterial community displayed large changes at wood ash dosages of 9 and 15 t ha–1 while only higher dosages (>30 t ha–1) significantly changed the fungal community. The results confirm that fungi are less sensitive to pH changes than bacteria but also that fertilizing plantation forests with wood ash, viewed through the lens of microbial community changes, is a safe management at standard dosages (typically 3 t ha–1).

Arbuscular mycorrhizal fungi (AMF) are important in plant nutrient uptake, but their function is prone to environmental constraints including soil factors that may suppress AMF transfer of phosphorus (P) from the soil to the plant. The objective of this study was to disentangle the biotic and abiotic components of AMF-suppressive soils. Suppression was measured in terms of AMF-mediated plant uptake of 33P mixed into a patch of soil and treatments included soil sterilization, soil mixing, pH manipulation and inoculation with isolated soil fungi. The degree of suppression was compared to volatile organic compound (VOC) production by isolated fungi and to multi-element analysis of soils. For a selected suppressive soil, sterilization and soil mixing experiments confirmed a biotic component of suppression. A Fusarium isolate from that soil suppressed the AMF activity and produced greater amounts than other fungal isolates of the antimicrobial VOC trichodiene (a trichothecene toxin precursor), beta-chamigrene, alpha-cuprenene and p-xylene. These metabolites deserve further attention when unravelling the chemical background behind the suppression of AMF activity by soil microorganisms. For the abiotic component of suppression, soil liming and acidification experiments confirmed that suppression was strongest at low pH. The pH effect might be associated with changed availability of specific suppressive elements. Indeed 33P uptake from the soil patches correlated negatively to Al levels and Al toxicity seems to play a major role in the AMF suppressiveness at pH below 5.0–5.2. However, the documentation of a biotic component of suppression for both low and high pH soils leads to the conclusion that biotic and abiotic components of suppression may act in parallel in some soils. The current insight into the components of soil suppressiveness of the AMF activity aids to develop management practices that allow for optimization of AMF functionality.

Wood ash is a by-product from energy production that can be recycled to forests to regain nutrients and prevent acidification. However, low concentrations of nitrogen (N) in wood ash may reduce its potential positive effect on plant growth. In addition, wood ash can have a high content of toxic heavy metals such as Cd, thus there are concerns that it may increase Cd accumulation in plants. We grew Deschampsia flexuosa (Wavy hair-grass) in pots in acidic nutrient poor forest soil fertilized with different concentrations of wood ash (corresponding to field application of 0, 1.1, 3.3, 11 and 33 t ha−1). Additionally, to disentangle the pH and nutrient effects of wood ash application, we included treatments with either CaO, to simulate pH effects of wood ash, or potassium (K) + phosphorus (P) fertilizer to mimic the nutrient effects. After 4.5 months of growth, we measured soil pH, plant biomass, Cd accumulation in shoots and N concentration in the various compartments of the system. Wood ash addition stimulated plant growth, whereas CaO and K + P addition resulted in more moderate increases in biomass. Despite the low concentration of N in the wood ash, plant uptake of N increased in wood ash amendments, probably because wood ash stimulated mineralization of soil organic N. Plant Cd content significantly increased at the highest dose of wood ash. Our results suggest that addition of wood ash significantly stimulates plant growth due to the combined effect of increased pH, elevated nutrient levels and increased N mineralization. Furthermore, despite the rather high Cd content in used wood ash (16.3 mg kg−1), wood ash amendments up to 11 t ha−1 did not result in significantly increased plant uptake of Cd.

Isotope labeling enables the detection and quantification of nutrient fluxes between soil and plants through arbuscular mycorrhizal (AM) fungi. Here we describe the use of radioactive isotopes, 33P and 32P, to study the uptake of P from soil by AM fungal mycelium and its transfer to the host plant through the mycorrhizal pathway.

Suppression of the activity of arbuscular mycorrhizal fungi (AMF) in unsterile soils, as determined by transfer of 33P from soils to plants, has been recently demonstrated, and soils differ in their degree of suppressiveness. The previous experiments included only one R. irregularis isolate, BEG87. The aim of this study was to analyze the sensitivity to suppression of different cultivated AMF isolates as well of native AMF communities using shoot P-33 content as a proxy of AMF activity in mesh-enclosed patches of unsterile soil. We found that the suppression of a panel of Rhizophagus isolates differed from the suppression of R. irregularis BEG87. Suppression overall correlated with low soil pH and high soil phosphorus, but there were large differences between the isolates. Some Rhizophagus isolates were less sensitive to soil suppression than BEG87, and tolerance to suppression may be a relevant property to consider when choosing future AMF inoculants. For native AMF communities showing high root colonization, we found one example of a community that was not suppressed by its 'home' soil, while BEG87 activity in that soil was completely suppressed. Hence, some AMF communities appear to be reservoirs for inoculants with high tolerance towards suppression.

In the face of global climate change there is an increasing demand for biofuel, which exerts pressure on production and thus management of biofuel plantations. The intensification of whole-tree harvest from biofuel plantations increases export of nutrients. Returning ash from biofuel combustion to the forest plantations can amend the soil nutrient status and thus facilitate sustainable forest management. However, ash affects the forest floor decomposer food web, potentially changing organic matter turnover, carbon sequestration and nitrogen availability. Our aim was to examine the response of decomposer organisms, food web structure and nitrogen mineralization function after ash application. In a coniferous forest plantation amended with 0, 3, 4.5 or 6 t ash ha− 1, we sampled in several depths of the forest floor for key organisms of the decomposer food web (fungal biomass, 0–12 cm; bacteria, protozoa, nematodes and enchytraeids, 0–3 cm and 3–6 cm; microarthropods and earthworms, 0–5 cm), 2, 14 and 26 months after ash application. We used structural equation modelling (SEM) to detangle the direct and indirect effects of ash application on organisms in the decomposer food web and on nitrogen availability. We found that ash increased the abundance of bacteria and protozoa, as well as the inorganic nitrogen pool at 0–3 cm depth, whereas the effect of ash was negligible at 3–6 cm depth. Earthworm abundance increased, whereas enchytraeid abundance decreased 2 years after ash application. The structural equation modelling showed that ash application stimulated the bacterial feeding pathway and increased nitrogen mineralization. Contrary, ash had a negative effect on fungal biomass at the first sampling, however, this effect subdued over time. Our results suggest that as the soil decomposer food web is resilient to ash application, this is a viable option for sustainable management of biofuel plantations.

Ectomycorrhizal (ECM) fungi are key players in N cycling in coniferous forests, and forest management such as application of wood ash can affect their functionality. The aim of this study was to determine the effects of wood ash application on ECM fungal mycelial production, capacity to retain N, diversity and community composition. In-growth mesh bags were installed in control and treated plots. After 6 months, 15N labeled ammonium and nitrate were applied into the mesh bags, and 24 h later extramatrical mycelium (EMM) was extracted and analyzed. Wood ash had no effects on EMM in-growth, N retention capacity, diversity or community composition. In contrast, there were significant seasonal differences in the amount of EMM produced. These results demonstrate that applying up to 6 t ha−1 of wood ash in this type of plantation forest is a safe management practice that does not increase N leaching or negatively affect ECM fungi.

Most plants form symbiotic associations with arbuscular mycorrhizal fungi (AMF). AMF increase the uptake of plant nutrients by extending their extra-radical mycelium (ERM) in the soil where other groups of microorganisms may suppress the activity of the ERM. However, little is known about such suppression in natural soils. This work aimed to investigate the incidence of AMF suppression among soils sampled from highly variable natural ecosystems, and used 33 P uptake by the ERM to evaluate AMF activity. A second aim was to identify factors behind the observed AMF-suppression. We found that AMF-suppressiveness varied markedly among natural soils and occurred more frequently in low pH than in high pH soils. A previous study for cultivated soils revealed a strong biological component of suppressiveness against AMF, and in accordance we found that the composition of both fungal and bacterial communities differed significantly between AMF-suppressive and non-suppressive natural soils. Acidobacteria, Acidothermus, Xanthomonadaceae, Archaeorhizomyces sp., Mortierella humilis and some Mycena spp. were significantly more abundant in AMF-suppressive soils and may therefore be direct antagonists of AMF. This implies that the functioning of AMF in natural ecosystems is strongly modulated by specific soil microbes.

Ash from combustion of biofuels, for example wood chips, is often deposited as waste, but due to its high content of essential plant nutrients and alkalinity, it has been proposed to recycle ash as a fertilizer and liming agent in biofuel production forest. However, current legislation sets rather strict limitations for wood-ash application in biofuel production systems. The soil microfood web, that is microorganisms and their microfaunal grazers, protozoa and nematodes, is pivotal for essential ecosystem processes such as decomposition and plant nutrient release. Therefore, a thorough assessment of the impacts on microfood web structure and functioning must precede actions towards raising the currently allowed application rates. In a Danish Norway spruce plantation, we evaluate the impact of wood ash applied at dosages from 0 to the extreme case of 90 t ash ha −1 on the microfood web, the bacterial community structure, soil content of inorganic nitrogen, organic matter, dissolved organic carbon and nitrogen. Using structural equation modelling (SEM), we disentangled the direct effect of the disturbance imposed by ash per se, the associated pH increase and changes in prey abundance on individual organism groups in the microfood web. The SEM showed that the pH rise was the main driver of increasing abundances of culturable heterotrophic bacteria with increasing ash doses, and via trophical transfer, this also manifested as higher abundances of bacterial grazers. Fungal-feeding nematodes were unaffected by ash, whereas carnivorous/omnivorous nematodes decreased due to the direct effect of ash. Increasing ash doses enhanced the difference between bacterial communities of control plots and ash-amended plots. The ash-induced stimulation of culturable heterotrophic bacteria and bacterial grazers increased inorganic nitrogen availability at ash doses of 9 t ha −1 and above. Hence, raised limits for ash application may potentially benefit tree growt

The identification of causal links between microbial community structure and ecosystem functions are required for a mechanistic understanding of ecosystem responses to environmental change. One of the most influential factors affecting plants and microbial communities in soil in managed ecosystems is the current land-use. In forestry, wood ash has been proposed as a liming agent and a fertilizer, but has been questioned due to the risk associated with its Cd content. The aim of this study was to determine the effects of wood ash on the structure and function of decomposer microbial communities in forest soils and to assign them to causal mechanisms. To do this, we assessed the responses to wood ash application of (i) the microbial community size and structure, (ii) microbial community trait-distributions, including bacterial pH relationships and Cd-tolerance, to assign the microbial responses to pH and Cd, and (iii) consequences for proxies of the function soil organic matter (SOM) turnover including respiration and microbial growth rates. Two sets of field-experiments in temperate conifer forest plantations were combined with laboratory microcosm experiments where wood ash additions were compared to additions of lime and Cd. Wood ash induced structural changes in the microbial community in both field experiments, and striking similarities were observed between the application of ash and that of lime in the microcosm experiments. Wood ash increased pH, and led to a shift toward faster SOM decomposition and a reduced importance of fungi. This coincided with shifts in bacterial community trait distributions for pH, with pH optima closely tracking the new soil pH. A Cd solution could induce Cd-tolerance in the microcosm experiments, but the ash did not affect the microbial tolerance to Cd in field or microcosm experiments. We demonstrate that the microbial community responded strongly to the application of wood ash to forest soils with consequences for its functiona

Reutilizing biomass ash on soil has been proposed to counteract soil acidity and to save fertilizer inputs by recycling valuable nutrients contained in biomass ash such as calcium (Ca), magnesium (Mg) and phosphorus (P). However, the heavy metal content of biomass ashes, such as cadmium (Cd), copper (Cu) and nickel (Ni), is a major environmental concern. This work is part of the project ASHBACK (www.ashback.dk) which addresses the potentials and possible problems in re-distributing wood ash to the forest. The aim of this thesis was to determine the effects of biomass ash application in agricultural and forest soils focusing on soil microbial communities’ composition and function, particularly mycorrhizal fungi. Two study sites were used for this study, one in an agricultural field where different biomass ashes were evaluated as replacements for P fertilizers in barley, and a second one in a Norway spruce forest where different amounts of wood ash were spread on the soil to study the effects on ectomycorrhizal (ECM) fungi, bioaccumulation of metals in sporocarps, and microbial communities. Laboratory microcosm experiments were run in parallel to the field studies, to compare the effects of wood ash with factorial additions of lime and Cd to disentangle the pH and Cd effects of wood ash amendments using community trait distributions. Barley yield, P content, and Cd content were not affected by biomass ashes. Some arbuscular mycorrhizal (AM) fungal species were reduced when biomass ashes were used. However, the AM colonization was as abundant and active as in the untreated plots. In the Norway spruce experiment, ECM mycelial production and nitrogen (N) retention capacity remained unaffected. Moreover, ECM fungal richness, diversity and community composition did not change with wood ash additions. Sporocarps collected from wood ash treated plots did not bioaccumulate more heavy metals than control plots. When analysing the soil microbial community st

Reutilizing biomass ashes in agriculture can substitute inputs of P from finite primary sources. However, recycling of ashes is disputed due to their content of toxic substances such as heavy metals. This study evaluates the potential risk of replacing easily soluble inorganic P fertilizer with P in biomass ashes in a barley crop grown on soil with adequate P status. Two contrasting doses of three different types of ashes were applied to an agricultural field with spring barley and compared to similar doses of triple-superphosphate fertilizer. In the second growing season after biomass ash application, grain, straw and root dry matter yield, and P and Cd uptake were determined. Resin-extractable P was measured in soil and the symbiotic arbuscular mycorrhizal fungal activity, colonization, and community composition were assessed. Crop yield was not affected by ash application, while P-uptake and mycorrhizal status were slightly enhanced with high ash applications. Changes to the mycorrhizal community composition were evident with high ash doses. Cadmium uptake in aboveground plant tissue was unaffected by ash treatments, but increased in roots with increasing doses. Consequently, we conclude that fertilization with biomass ashes can replace conventional fertilizers without risk to barley crops in the short term.