Marcel G. A. van der Heijden

The arbuscular mycorrhizal (AM) symbiosis is widespread throughout the plant kingdom but most terrestrial ecosystems also contain a considerable number of non‐mycorrhizal plants. The interaction of such non‐host plants with AM fungi is still poorly understood. In this study, we investigated whether the non‐mycorrhizal plant Arabidopsis thaliana, the model organism for plant molecular biology and genetics, interacts with AM fungi. We demonstrate, for the first time, that the presence of fungal networks formed by the AM fungus Rhizophagus irregularis reduces A. thaliana growth by 50% or more. In addition, by using bright field, confocal and electronic microscopy we show that this fungus can colonize roots of A. thaliana, although arbuscules were never observed. These results reveal high susceptibility of A. thaliana to R. irregularis, suggesting that A. thaliana is a suitable model plant to study non‐host/AM fungi interactions and the biological basis of AM incompatibility.Commentary: Interactions between arbuscular mycorrhizal and non‐mycorrhizal plants: do non‐mycorrhizal species at both extremes of nutrient‐availability play the same game?

There is a great interest in ecology in understanding the role of soil microbial diversity for plant productivity and coexistence. Recent research has shown increases in species richness of mutualistic soil fungi, the arbuscular mycorrhizal fungi (AMF), to be related to increases in aboveground productivity of plant communities. However, the impact of AMF richness on plant–plant interactions has not been determined. Moreover, it is unknown whether species‐rich AMF communities can act as insurance to maintain productivity in a fluctuating environment (e.g., upon changing soil conditions).We tested the impact of four different AMF taxa and of AMF diversity (no AMF, single AMF taxa, and all four together) on competitive interactions between the legumeTrifolium pratenseand the grassLolium multiflorumgrown under two different soil conditions of low and high sand content. We hypothesized that more diverse mutualistic interactions (e.g., when four AMF taxa are present) can ease competitive effects between plants, increase plant growth, and maintain plant productivity across different soil environments. We used quantitative PCR to verify that AMF taxa inoculated at the beginning of the experiment were still present at the end.The presence of AMF reduced the competitive inequality between the two plant species by reducing the growth suppression of the legume by the grass. High AMF richness enhanced the combined biomass production of the two plant species and the yield of the legume, particularly in the more productive soil with low sand content. In the less productive (high sand content) soil, the single most effective AMF had an equally beneficial effect on plant productivity as the mixture of four AMF. Since contributions of single AMF to plant productivity varied between both soils, higher AMF richness would be required to maintain plant productivity in heterogeneous environments. Overall this work shows that AMF diversity promotes plant productivity and that AMF diversity can act as insurance to sustain plant productivity under changing environmental conditions.

Nutrient loss from ecosystems is among the top environmental threats to ecosystems worldwide, leading to reduced plant productivity in nutrient‐poor ecosystems and eutrophication of surface water near nutrient‐rich ecosystems. Hence, it is of pivotal importance to understand which factors influence nutrient loss. Here it is demonstrated that arbuscular mycorrhizal (AM) fungi, widespread soil fungi that form mutualistic relationships with the majority of land plants, reduce nutrient loss from grassland microcosms during rain‐induced leaching events.Grassland microcosms with AM fungi lost 60% less phosphorus and 7.5% less ammonium compared to control microcosms without AM fungi. Similar results were obtained for microcosms planted with each of three different grass species. In contrast, nitrate leaching was not affected by AM fungi but depended on the amount of nutrients supplied to the microcosms. Moreover, fertilization of the microcosms reduced the abundance of AM fungi and their ability to reduce phosphorus leaching losses. Extrapolation of these results suggests that the disruption of the mycorrhizal symbiosis is one of the reasons for enhanced phosphorus loss from fertilized ecosystems.The microcosms contained a sandy soil, a soil type vulnerable to leaching losses. The reduction of phosphorus leaching by AM fungi may, therefore, represent an upper limit. Advantages and limitations of the experimental setup for assessing the impact of AM fungi on nutrient cycling are discussed.The results indicate that AM fungi contribute to ecosystem sustainability by promoting a closed phosphorus cycle and reducing phosphorus leaching losses.

*The impact of various agricultural practices on soil biodiversity and, in particular, on arbuscular mycorrhizal fungi (AMF), is still poorly understood, although AMF can provide benefit to plants and ecosystems. Here, we tested whether organic farming enhances AMF diversity and whether AMF communities from organically managed fields are more similar to those of species-rich grasslands or conventionally managed fields. *To address this issue, the AMF community composition was assessed in 26 arable fields (13 pairs of organically and conventionally managed fields) and five semi-natural grasslands, all on sandy soil. Terminal restriction fragment length polymorphism community fingerprinting was used to characterize AMF community composition. *The average number of AMF taxa was highest in grasslands (8.8), intermediate in organically managed fields (6.4) and significantly lower in conventionally managed fields (3.9). Moreover, AMF richness increased significantly with the time since conversion to organic agriculture. AMF communities of organically managed fields were also more similar to those of natural grasslands when compared with those under conventional management, and were less uniform than their conventional counterparts, as expressed by higher beta-diversity (between-site diversity). *We suggest that organic management in agro-ecosystems contributes to the restoration and maintenance of these important below-ground mutualists.

Summary: 1. Almost all plants are engaged in symbiotic relationships with mycorrhizal fungi. These soil fungi can promote plant growth by supplying limiting nutrients to plant roots in return for plant assimilates. 2. Many mycorrhizal fungi are not host specific and one fungal individual can colonize and interconnect a considerable number of plants. The existence of these so‐called mycorrhizal networks implies that fungi have the potential to facilitate growth of other plants and distribute resources among plants irrespective of their size, status or identity. In this paper, we explore the significance of mycorrhizal fungal networks for individual plants and for plant communities. 3. We address the following questions: (i) are all plant species benefitting from mycorrhizal networks, (ii) is benefit dependent on the size or age of a plant, (iii) is fungal support related to the relative dominance of plants in a community, (iv) are there host dependent barriers and physiological constraints for support and (v) what is the impact of mycorrhizal networks on plant–plant interactions and plant community dynamics? Moreover, using a review of published studies, we test whether mycorrhizal networks facilitate growth of small seedlings that establish between or near larger plants. 4. We found 60 cases where seedling species were grown together with larger plants with or without mycorrhizal fungal networks. Mycorrhizal networks promoted seedling growth in 48% of the cases (for 21 seedling species), while negative effects (25%) and no effects (27%) were also common. Seedlings associating with ectomycorrhizal fungi benefitted in the majority of the cases while effects on seedlings associating with arbuscular mycorrhizal fungi were more variable. Thus, the facilitative effects of mycorrhizal fungal networks depend on seedling species identity, mycorrhizal identity, plant species combinations and study system. We present a number of hypothetical scenarios that can explain the results based on cost–benefit relationship of individual members in a network. 5. Synthesis. Overall, this review shows that mycorrhizal networks play a key role in plant communities by facilitating and influencing seedling establishment, by altering plant–plant interactions and by supplying and recycling nutrients.

Nitrogen (N) inputs to ecosystems have increased worldwide, often leading to large changes in plant community structure and reducing plant diversity. Yet, the interaction of increased N availability with other factors that determine plant community composition, are still poorly understood. Here, we test whether the impact of N addition on plant communities depends on the presence of arbuscular mycorrhizal fungi (AMF). AMF are widespread plant symbionts that facilitate growth of many plant species. We hypothesize that AM fungi reduce the negative impact of N addition on plant communities by supporting growth of species that are sensitive to N enrichment.We established experimental grassland microcosms consisting of 18 plant species. These microcosms were subjected to high and low N supply and were inoculated with AMF or remained nonmycorrhizal. Both N addition and AMF had a big impact on plant community composition, but with opposite effects. N addition induced a 2.8‐fold increase in grass biomass and reduced legume biomass. Grasses dominated the microcosms at high N supply, especially when AMF were absent. In contrast, AMF enhanced biomass of all legumes species (on average 6.8‐fold) and reduced the relative abundance of grasses. The proportion of legume biomass out of total shoot biomass at high N supply was 19% with AMF and only 3% without AMF. Our results show that responses of plant communities to N enrichment depend on AMF and that AMF can reduce the negative impact of increased N availability on plant community structure by reducing grass dominance.

Microbes are the unseen majority in soil and comprise a large portion of life’s genetic diversity. Despite their abundance, the impact of soil microbes on ecosystem processes is still poorly understood. Here we explore the various roles that soil microbes play in terrestrial ecosystems with special emphasis on their contribution to plant productivity and diversity. Soil microbes are important regulators of plant productivity, especially in nutrient poor ecosystems where plant symbionts are responsible for the acquisition of limiting nutrients. Mycorrhizal fungi and nitrogen‐fixing bacteria are responsible forc.5–20% (grassland and savannah) to 80% (temperate and boreal forests) of all nitrogen, and up to 75% of phosphorus, that is acquired by plants annually. Free‐living microbes also strongly regulate plant productivity, through the mineralization of, and competition for, nutrients that sustain plant productivity. Soil microbes, including microbial pathogens, are also important regulators of plant community dynamics and plant diversity, determining plant abundance and, in some cases, facilitating invasion by exotic plants. Conservative estimates suggest thatc.20 000 plant species are completely dependent on microbial symbionts for growth and survival pointing to the importance of soil microbes as regulators of plant species richness on Earth. Overall, this review shows that soil microbes must be considered as important drivers of plant diversity and productivity in terrestrial ecosystems.