Root fungal symbionts interact with mammalian herbivory, soil nutrient availability and specific habitat conditions

Herbivory, competition and soil fertility interactively shape plant communities and exhibit an important role in modifying conditions for host-dependent fungal symbionts. However, field studies on the combined impacts of natural herbivory, competition and soil fertility on root fungal symbionts are rare. We asked how mammalian herbivory, fertilization, liming and plant–plant competition affect the root colonization of arbuscular mycorrhizal fungi (AMF) and dark septate endophytic (DSE) fungi of the dicot herb, Solidago virgaurea. The 2-year full-factorial experiment was conducted in two contrasting habitats: non-acidic and acidic mountain tundra. We found that herbivory increased arbuscular colonization (i.e. the site of resource exchange) at fertile non-acidic sites, where vegetation was rich in species having AMF symbionts, whereas at infertile acidic sites, where plants having AMF symbiont are scarce, the response was the opposite. Herbivory of the host plant negatively affected DSE hyphal and sclerotial colonization in unfertilized plots, possibly due to reduced carbon flow from the host plant while there was no effect of herbivory in fertilized plots. DSE colonization was highest in unfertilized exclosures where soil nutrient concentrations were also lowest. Liming had a negative effect on DSE hyphal colonization, and its effect also interacted with herbivory and the habitat. Biomass removal of the neighboring plants did not affect the root colonization percent of either arbuscules or DSE. Our results show that the impacts of aboveground mammalian herbivory, soil nutrient availability and specific habitat conditions on belowground root fungal symbionts are highly dependent on each other. Arbuscule response to herbivory appeared to be regulated by specific habitat conditions possibly caused by differences in the AMF availability in the soil while DSE response was associated with availability of host-derived carbon. Our result of the relationship between herbivory and soil nutrients suggests an important role of DSE in ecosystem processes.     

陆地农业生态系统丛枝菌根真菌物种多样性研究进展

丛枝菌根真菌(AMF)是一种古老的、在自然界中普遍存在的土壤微生物,能与大部分陆生植物形成互惠互利的菌根共生体.在这种共生关系中,AMF从植物获取自身生长所需碳源的同时,帮助宿主吸收氮、磷等营养物质.AMF在农业生态系统中具有重要作用,能够促进植物生长、改善作物品质、提高植物抗逆性、稳定土壤结构、维护生态平衡和维持农业可持续发展.本文总结了近几年来陆地农业生态系统AMF的研究进展,着重从我国陆地农业生态系统AMF物种多样性、AMF生物多样性时空分布特征及影响AMF多样性的因素等几个方面,综述了陆地农业生态系统AMF的物种多样性,并对以后的研究进行了展望.     

Phylogenetic relatedness explains highly interconnected and nested symbiotic networks of woody plants and arbuscular mycorrhizal fungi in a Chinese subtropical forest

Elucidating symbiotic relationships between arbuscular mycorrhizal fungi (AMF) and plants contributes to a better understanding of their reciprocally dependent coexistence and community assembly. However, the main drivers of plant and AMF community assembly remain unclear. In this study, we examined AMF communities from 166 root samples of 17 woody plant species from 10 quadrats in a Chinese subtropical forest using 454 pyrosequencing of 18S rRNA gene to describe symbiotic AMF–plant association. Our results show the woody plant–AMF networks to be highly interconnected and nested, but in antimodular and antispecialized manners. The nonrandom pattern in the woody plant–AMF network was explained by plant and AMF phylogenies, with a tendency for a stronger phylogenetic signal by plant than AMF phylogeny. This study suggests that the phylogenetic niche conservatism in woody plants and their AMF symbionts could contribute to interdependent AMF and plant community assembly in this subtropical forest ecosystem.     

Plant–mycorrhizal fungus co‐occurrence network lacks substantial structure

The interactions between plants and arbuscular mycorrhizal fungi (AMF) maintain a crucial link between macroscopic organisms and the soil microbial world. These interactions are of extreme importance for the diversity of plant communities and ecosystem functioning. Despite this importance, only recently has the structure of plant–AMF interaction networks been studied. These recent studies, which used genetic data, suggest that these networks are highly structured, very similar to plant–animal mutualistic networks. However, the assembly process of plant–AMF communities is still largely unknown, and an important feature of plant–AMF interactions has not been incorporated: they occur at an extremely localized scale. Studying plant–AMF networks in a spatial context seems therefore a crucial step. This paper studies a plant–AMF spatial co‐occurrence network using novel methodology based on information theory and a unique set of spatially explicit species‐level data. We apply three null models of which only one accounts for spatial effects. We find that the data show substantial departures from null expectations for the two non‐spatial null models. However, for the null model considering spatial effects, there are few significant co‐occurrences compared with the other two null models. Thus, plant–AMF spatial co‐occurrences seem to be mostly explained by stochasticity, with a small role for other factors related to plant–AMF specialization. Furthermore, we find that the network is not significantly nested or modular. We conclude that this plant–AMF spatial co‐occurrence network lacks substantial structure and, therefore, plants and AMF species do not track each other over space. Thus, random encounters seem more important in the first step of the assembly of plant–AMF communities. Synthesis The symbiotic interaction between plants and arbuscular mycorrhizal fungi (AMF) is crucial for ecosystem functioning. However, the factors affecting the assembly of plant‐AMF communities are poorly understood. An important factor of the assembly of plant‐AMF communities has been overlooked: plant‐AMF interactions occur at a localized spatial scale. Our study investigated the importance of space in the structure of plant‐AMF communities. We studied a plant‐AMF spatial co‐occurrence network using a unique set of spatially explicit data and applied three null models. We found that plant‐AMF spatial co‐occurrences seem to be mostly explained by stochasticity. In particular, our study shows that this plant‐AMF spatial co‐occurrence network lacks substantial structure and, therefore, plants and AMF species do not track each other over space. Thus, random encounters seem to drive the assembly of plant‐AMF communities.     

Soil,But Not Cultivar,Shapes the Structure of Arbuscular Mycorrhizal Fungal Assemblages Associated with Strawberry

Arbuscular mycorrhizal fungi are widespread plant symbionts occurring in most agricultural crops, where they can play key roles in the growth and health of their plant hosts. Plant benefits can depend on the identity of the associated arbuscular mycorrhizal fungi (AMF), but little is known about the identity of the fungal partners in most agricultural systems. In this study, we describe the AMF assemblages associated with four cultivars of strawberry in an outdoor experiment using two field soils with different origin and management history. Assemblages were characterised by clone library sequencing of 18S rRNA gene fragments. Soil dramatically influenced the degree of mycorrhizal colonisation and AMF assemblage structure in the roots. No differences were observed between cultivars. Fungi belonging to the genus Acaulospora dominated the AMF assemblages in one soil, but they were not detected in the other. These results suggest that physicochemical soil characteristics and management can play a role in determining the identity and structure of microbial communities associated with particular hosts in agricultural systems.     

Community assembly, species richness and nestedness of arbuscular mycorrhizal fungi in agricultural soils

Understanding how communities assemble is a central goal of ecology. This is particularly relevant for communities of arbuscular mycorrhizal fungi (AMF), because the community composition of these beneficial plant symbionts influences important ecosystem processes. Moreover, AMF may be used as sensitive indicators of ecological soil quality if they respond to environmental variation in a predictable way. Here, we use a molecular profiling technique (T-RFLP of 25S rRNA gene fragments) to test which factors determine AM fungal community composition in 40 agricultural soils in the Netherlands. In particular, we test whether species richness, dominance structure and community nestedness are influenced by management type (in pairs of organically and conventionally farmed fields), and we examine the contribution of crop species (maize vs. potato), soil type (sand vs. clay-textured soils) and habitat (plant root vs. bulk soil) on AMF community characteristics. AMF richness varied from 1 to 11 taxa per field. Communities from species-poor fields were found to be subsets of those in richer fields, indicating nestedness and a progressive 'loss' from the species pool. AMF taxa richness and occurrence in soil and plant roots were highly correlated, and richness was related to management intensity (phosphate availability and grass-cropping history together explained 32% and 50% of richness in roots and soils). Soil type together with soil chemical parameters explained only 17% of variance in AMF community structure. We synthesize these results by discussing the potential contribution of a 'bottleneck effect' on AMF communities through increased stochastic effects under environmental stress.     

Studies on Ectomycorrhiza: An Appraisal

Ectomycorrhizal (ECM) fungi are obligate symbionts of dominant vascular plants, liverworts and hornworts. There are reports of about 20,000 to 25,000 ECM fungi that promote plant growth by facilitating enhanced water and nutrient absorption, and provide tolerance to environmental stresses. These below-ground fungi play a key role in terrestrial ecosystems as they regulate plant diversity, nutrient and carbon cycles, and influence soil structure and ecosystem multifunctionality. Because ECM fungi are obligate root symbionts, host plant can have a strong effect on ECM species richness and community composition. The biogeographic pattern and detailed functioning and regulation of these mycorrhizosphere processes are still poorly understood and require detailed study. More recent researches have placed emphasis on a wider, multifunctional perspective, including the effects of ectomycorrhizal symbiosis on plant and microbial communities, and on ecosystem processes. Over the years the main focus in ECM research has been on the study of diversity and specificity of ECM strains, the role of ECM in regeneration of degraded ecosystem, the growth and establishment of seedlings through nutrient acquisition and the mediation of plant responses to various types of stress. In this review, recent progresses in ectomycorrhizal biology are presented, especially the potential role of ECM symbioses in resistance or tolerance to various biotic and abiotic stresses, and in maintinance of plant diversity for proper ecosystem functioning.     

Interaction between Earthworms and Arbuscular Mycorrhizal Fungi in Plants: A Review

Different kinds of soil animals and microorganisms inhabit the plant rhizosphere, which function closely to plant roots. Of them, arbuscular mycorrhizal fungi (AMF) and earthworms play a critical role in sustaining the soil-plant health. Earthworms and AMF belong to the soil community and are soil beneficial organisms at different trophic levels. Both of them improve soil fertility and structural development, collectively promoting plant growth and nutrient acquisition capacity. Earthworm activities redistribute mycorrhizal fungi spores and give diversified effects on root mycorrhizal fungal colonization. Dual inoculation with both earthworms and AMF strongly magnifies the response on plant growth through increased soil enzyme activities and changes in soil nutrient availability, collectively mitigating the negative effects of heavy metal pollution in plants and soils. This thus enhances phytoremediation and plant disease resistance. This review simply outlines the effects of earthworms and AMF on the soil-plant relationship. The effects of earthworms on root AMF colonization and activities are also analyzed. This paper also summarizes the interaction between earthworms and AMF on plants along with suggested future research.     

Plant species richness,identity and productivity differentially influence key groups of microbes in grassland soils of contrasting fertility

The abundance of microbes in soil is thought to be strongly influenced by plant productivity rather than by plant species richness per se. However, whether this holds true for different microbial groups and under different soil conditions is unresolved. We tested how plant species richness, identity and biomass influence the abundances of arbuscular mycorrhizal fungi (AMF), saprophytic bacteria and fungi, and actinomycetes, in model plant communities in soil of low and high fertility using phospholipid fatty acid analysis. Abundances of saprophytic fungi and bacteria were driven by larger plant biomass in high diversity treatments. In contrast, increased AMF abundance with larger plant species richness was not explained by plant biomass, but responded to plant species identity and was stimulated by Anthoxantum odoratum. Our results indicate that the abundance of saprophytic soil microbes is influenced more by resource quantity, as driven by plant production, while AMF respond more strongly to resource composition, driven by variation in plant species richness and identity. This suggests that AMF abundance in soil is more sensitive to changes in plant species diversity per se and plant species composition than are abundances of saprophytic microbes.     

Unraveling spatiotemporal variability of arbuscular mycorrhizal fungi in a temperate grassland plot

Soils provide a heterogeneous environment varying in space and time; consequently, the biodiversity of soil microorganisms also differs spatially and temporally. For soil microbes tightly associated with plant roots, such as arbuscular mycorrhizal fungi (AMF), the diversity of plant partners and seasonal variability in trophic exchanges between the symbionts introduce additional heterogeneity. To clarify the impact of such heterogeneity, we investigated spatiotemporal variation in AMF diversity on a plot scale (10 × 10 m) in a grassland managed at low intensity in southwest Germany. AMF diversity was determined using 18S rDNA pyrosequencing analysis of 360 soil samples taken at six time points within a year. We observed high AMF alpha- and beta-diversity across the plot and at all investigated time points. Relationships were detected between spatiotemporal variation in AMF OTU richness and plant species richness, root biomass, minimal changes in soil texture and pH. The plot was characterized by high AMF turnover rates with a positive spatiotemporal relationship for AMF beta-diversity. However, environmental variables explained only ≈20% of the variation in AMF communities. This indicates that the observed spatiotemporal richness and community variability of AMF was largely independent of the abiotic environment, but related to plant properties and the cooccurring microbiome.     

The influence of arbuscular mycorrhizae on the relationship between plant diversity and productivity

Ecological theory predicts a positive and asymptotic relationship between plant diversity and ecosystem productivity based on the ability of more diverse plant communities to use limiting resources more fully. This is supported by recent empirical evidence. Additionally, in natural ecosystems, plant productivity is often a function of the presence and composition of mycorrhizal associations. Yet, the effect of mycorrhizal fungi on the relationship between plant diversity and productivity has not been investigated. We predict that in the presence of AMF, productivity will saturate at lower levels of species richness because AMF increase the ability of plant species to utilize nutrient resources. In this study we manipulated old-field plant species richness in the presence and absence of two species of AMF. We found that in the absence of AMF, the relationship between plant species richness and productivity is positive and linear. However, in the presence of AMF, the relationship is positive but asymptotic, even though the maximum plant biomass was significantly different between the two AMF treatments. This is consistent with the hypothesis that AMF increase the redundancy of plant species in the productivity of plant communities, and indicates that these symbionts must be considered in future investigations of plant biodiversity and ecosystem function.     

The unseen majority: soil microbes as drivers of plant diversity and productivity in terrestrial ecosystems

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 for c. 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 that c. 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.     

Microbial biofilms in nature: unlocking their potential for agricultural applications

Soil environments are dynamic and the plant rhizosphere harbours a phenomenal diversity of micro-organisms which exchange signals and beneficial nutrients. Bipartite beneficial or symbiotic interactions with host roots, such as mycorrhizae and various bacteria, are relatively well characterized. In addition, a tripartite interaction also exists between plant roots, arbuscular mycorrhizal fungi (AMF) and associated bacteria. Bacterial biofilms exist as a sheet of bacterial cells in association with AMF structures, embedded within a self-produced exopolysaccharide matrix. Such biofilms may play important functional roles within these tripartite interactions. However, the details about such interactions in the rhizosphere and their relevant functional relationships have not been elucidated. This review explores the current understanding of naturally occurring microbial biofilms, and their interaction with biotic surfaces, especially AMF. The possible roles played by bacterial biofilms and the potential for their application for a more productive and sustainable agriculture is discussed in this review.     

Disturbance is more important than seeding or grazing in determining soil microbial communities in a semiarid grassland

A primary goal of ecological restoration is often to return processes and functions to degraded ecosystems. Soil, while often ignored in restoration, supports diverse communities of organisms and is a fundamental actor in providing ecosystem processes and services. We investigated the impact of seeding and livestock grazing on plant communities, soil microorganisms, and soil fertility 3 years after the restoration of a disturbed pipeline corridor in southeastern Arizona. The initial soil disturbance and topsoil treatment, regardless of seeding or grazing, was the most influential factor in determining differences in both plant and microbial communities. Compared with the control, the disturbed and restored sites had greater plant species richness, greater total herbaceous plant cover, greater soil organic matter, higher pH, and differed in soil nutrients. Bacteria and fungi appeared to generally correlate with micro‐environment and soil physiochemical properties rather than specific plant species. The undisturbed control had a smaller proportion of bacterial functional groups associated with the breakdown of plant biomass (polysaccharide decomposition) and a smaller proportion of arbuscular mycorrhizal fungi (AMF) compared with disturbed and restored sites. The ability of the unseeded disturbed site to recover robust vegetation may be due in part to the high presence of AMF. These differences show selection for soil microorganisms that thrive in disturbed and restored sites and may contribute to increased plant productivity. Restoration of specific plant species or ecological processes and services would both benefit from better understanding of the impacts of disturbance on soil microorganisms and soil fertility.     

Changes in the diversity of soil arbuscular mycorrhizal fungi after cultivation for biofuel production in a Guantanamo (Cuba) tropical system

The arbuscular mycorrhizal fungi (AMF) are a key, integral component of the stability, sustainability and functioning of ecosystems. In this study, we characterised the AMF biodiversity in a native vegetation soil and in a soil cultivated with Jatropha curcas or Ricinus communis, in a tropical system in Guantanamo (Cuba), in order to verify if a change of land use to biofuel plant production had any effect on the AMF communities. We also asses whether some soil properties related with the soil fertility (total N, Organic C, microbial biomass C, aggregate stability percentage, pH and electrical conductivity) were changed with the cultivation of both crop species. The AM fungal small sub-unit (SSU) rRNA genes were subjected to PCR, cloning, sequencing and phylogenetic analyses. Twenty AM fungal sequence types were identified: 19 belong to the Glomeraceae and one to the Paraglomeraceae. Two AMF sequence types related to cultured AMF species (Glo G3 for Glomus sinuosum and Glo G6 for Glomus intraradices-G. fasciculatum-G. irregulare) did not occur in the soil cultivated with J. curcas and R. communis. The soil properties (total N, Organic C and microbial biomass C) were higher in the soil cultivated with the two plant species. The diversity of the AMF community decreased in the soil of both crops, with respect to the native vegetation soil, and varied significantly depending on the crop species planted. Thus, R. communis soil showed higher AMF diversity than J. curcas soil. In conclusion, R. communis could be more suitable for the long-term conservation and sustainable management of these tropical ecosytems.     

SeSaMe:Metagenome Sequence Classification of Arbuscular Mycorrhizal Fungi-associated Microorganisms

Arbuscular mycorrhizal fungi (AMF) are plant root symbionts that play key roles in plant growth and soil fertility. They are obligate biotrophic fungi that form coenocytic multinucleated hyphae and spores. Numerous studies have shown that diverse microorganisms live on the surface of and inside their mycelia, resulting in a metagenome when whole-genome sequencing (WGS) data are obtained from sequencing AMF cultivated in vivo. The metagenome contains not only the AMF sequences, but also those from associated microorganisms. In this study, we introduce a novel bioinformatics program, Spore-associated Symbiotic Microbes (SeSaMe), designed for taxonomic classification of short sequences obtained by next-generation DNA sequencing. A genus-specific usage bias database was created based on amino acid usage and codon usage of a three consecutive codon DNA 9-mer encoding an amino acid trimer in a protein secondary structure. The program distinguishes between coding sequence (CDS) and non-CDS, and classifies a query sequence into a genus group out of 54 genera used as reference. The mean percentages of correct predictions of the CDS and the non-CDS test sets at the genus level were 71% and 50% for bacteria, 68% and 73% for fungi (excluding AMF), and 49% and 72% for AMF (Rhizophagus irregularis), respectively. SeSaMe provides not only a means for estimating taxonomic diversity and abundance but also the gene reservoir of the reference taxonomic groups associated with AMF. Therefore, it enables users to study the symbiotic roles of associated microorganisms. It can also be applicable to other microorganisms as well as soil metagenomes. SeSaMe is freely available at www.fungalsesame.org.     

丛枝菌根真菌参与下植物-土壤系统的养分交流及调控

近几年随着有机农业的发展,丛枝菌根的作用受到特别关注。丛枝菌根是由植物根系与丛枝菌根真菌(AMF)形成的一种共生体。在植物-AMF-土壤系统中,AMF为植物提供N、P等营养的同时从根系得到所需的C。概述了植物-AMF-土壤系统中C、N、P等营养物质的交流以及AMF与土壤微生物的互作关系。丛枝菌根的形成可显著提高植物对P的吸收,且在高P条件下多余的P可储存于AMF中。AMF对土壤N循环的影响相当复杂,可能参与调控N循环的多个过程,如硝化作用、反硝化作用和氨氧化作用等。在有机质丰富的土壤中AMF菌丝可快速扩增并吸收其中的N,主要供菌丝自身所需,只有一小部分传递给植物。AMF对土壤C库的影响尚存争议,可能存在时间尺度的差异。短期内可活化土壤C,而在长期尺度上可能有利于土壤C的储存。AMF能够通过改变土壤微生物群落结构而影响植物-土壤体系的物质交流。AMF与解磷菌、根瘤菌和放线菌的协同增效作用可促进土壤有机质的降解或增强其固氮能力;AMF对氨氧化菌的抑制作用可降低氨的氧化减少N2O的释放。AMF与外生共生真菌EMF共存时,表现出协同增效作用,但EMF的优先定殖会限制AMF的侵染。AMF不同类群之间则主要表现为竞争和拮抗关系。AMF与土壤微生物之间的互作关系受土壤无机环境的影响,在养分亏缺条件下微生物之间往往表现为竞争关系。因植物、AMF与土壤微生物之间存在复杂的互作关系,为此AMF并不总是表现出其对植物营养的促进作用。目前关于AMF的作用机理仍以假说为主,需要进一步的实验验证。在植物-AMF-土壤系统中N与C的交流和P与C的交流并未表现出一致性,对N、P循环相互关系的进一步探讨有助于深入理解植物-土壤体系中的养分循环。植物、AMF和土壤微生物的养分来源及其对养分的相对需求强度和吸收效率尚未可知,因此无法深入理解AMF在植物-土壤体系中养分交流和转化的作用。在方法上,传统的土壤学方法在养分动态研究中存在局限性,现代分子生物学手段和化学计量学的结合值得尝试。     

Mycorrhizal benefits on native plants of the Caatinga,a Brazilian dry tropical forest

In the terrestrial ecosystems, soil is an important component, characterized by holding high diversity of microorganisms which play a key role for productivity and vegetal composition. The group of symbionts microorganisms stands out for contributing directly to the growth and plant nutrition, and among them, the arbuscular mycorrhizal fungi form one of the oldest and well established associations. In order to increase the knowledge and contribute for further research with AMF and plants of Caatinga, in this review we compile data from previous studies on the effects of symbiosis between arbuscular mycorrhizal fungi (AMF) and plants of the Caatinga, a type of dry tropical forest found in the northeast of Brazil. These studies collected data under various experimental conditions, emphasizing fungal efficiency and host responsiveness in soils with varied fertility. From our analysis we conclude that in general the symbiotic efficiency on these plants depends on many factors, such as the plant-fungi combination, fertility and soil type. Furthermore, in leguminosae the impact of a joint inoculation with nitrogen fixing bacteria must be taken into account. Claroideoglomus etunicatum was the most tested AMF species benefiting almost all plants tested. Approximately 30 plant species were studied regarding possible benefits provided by AMF and of these only Hymenea courbaril and Aspidosperma pyrifolium did not respond to mycorrhization. Higher efficiency of the mycorrhizal symbiosis can be obtained in soils with low P levels, emphasizing the essential role of these microorganisms in the growth and survival of plant species from the Caatinga biome.     

The influence of competing root symbionts on below‐ground plant resource allocation

1. Plants typically interact with multiple above‐ and below‐ground organisms simultaneously, with their symbiotic relationships spanning a continuum ranging from mutualism, such as with arbuscular mycorrhizal fungi (AMF), to parasitism, including symbioses with plant‐parasitic nematodes (PPN).
2. Although research is revealing the patterns of plant resource allocation to mutualistic AMF partners under different host and environmental constraints, the root ecosystem, with multiple competing symbionts, is often ignored. Such competition is likely to heavily influence resource allocation to symbionts.
3. Here, we outline and discuss the competition between AMF and PPN for the finite supply of host plant resources, highlighting the need for a more holistic understanding of the influence of below‐ground interactions on plant resource allocation. Based on recent developments in our understanding of other symbiotic systems such as legume–rhizobia and AMF‐aphid‐plant, we propose hypotheses for the distribution of plant resources between contrasting below‐ground symbionts and how such competition may affect the host.
4. We identify relevant knowledge gaps at the physiological and molecular scales which, if resolved, will improve our understanding of the true ecological significance and potential future exploitation of AMF‐PPN‐plant interactions in order to optimize plant growth. To resolve these outstanding knowledge gaps, we propose the application of well‐established methods in isotope tracing and nutrient budgeting to monitor the movement of nutrients between symbionts. By combining these approaches with novel time of arrival experiments and experimental systems involving multiple plant hosts interlinked by common mycelial networks, it may be possible to reveal the impact of multiple, simultaneous colonizations by competing symbionts on carbon and nutrient flows across ecologically important scales.