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A major barrier to sustainably improving food security for a growing global population is the availability of suitable space for growing crops. Urban areas offer a potential solution to increase availability of land, however, horticultural soils often accumulate zinc. These increased levels may affect the interactions between crops and soil microbes with potential implications for crop health and nutrition. Using radio-isotope tracing, we investigated the effect of urban environmentally relevant concentrations of zinc in soils on the nutrient exchange between arbuscular mycorrhizal fungi and pea plants. At higher concentrations of zinc, transfer of phosphorus from fungi to plants and the movement of carbon from plants to fungi was dramatically decreased. Our results suggest that while urban horticulture holds promise for sustainably enhancing local food production and addressing global food security, the unchecked presence of contaminants in these soils may pose a critical hurdle to realizing the potential of urban soils.

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The minute 'dust seeds' of some terrestrial orchids preferentially germinate and develop as mycoheterotrophic protocorms near conspecific adult plants. Here we test the hypothesis that mycorrhizal mycelial connections provide a direct pathway for transfer of recent photosynthate from conspecific green orchids to achlorophyllous protocorms. Mycelial networks of Ceratobasidium cornigerum connecting green Dactylorhiza fuchsii plants with developing achlorophyllous protocorms of the same species were established on oatmeal or water agar before the shoots of green plants were exposed to 14CO2. After incubation for 48 h, the pattern of distribution of fixed carbon was visualised in intact entire autotrophic/protocorm systems using digital autoradiography and quantified in protocorms by liquid scintillation counting. Both methods of analysis revealed accumulation of 14C above background levels in protocorms, confirming that autotrophic plants supply carbon to juveniles via common mycorrhizal networks. Despite some accumulation of plant-fixed carbon in the fungal mycelium grown on oatmeal agar, a greater amount of carbon was transferred to protocorms growing on water agar, indicating that the polarity of transfer may be influenced by sink strength. We suggest this transfer pathway may contribute significantly to the pattern and processes determining localised orchid establishment in nature, and that 'parental nurture' via common mycelial networks may be involved in these processes.

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Long computation times in vegetation and climate models hamper our ability to evaluate the potentially powerful role of plants on weathering and carbon sequestration over the Phanerozoic Eon. Simulated vegetation over deep time is often homogenous, and disregards the spatial distribution of plants and the impact of local climatic variables on plant function. Here we couple a fast vegetation model (FLORA) to a spatially-resolved long-term climate-biogeochemical model (SCION), to assess links between plant geographical range, the long-term carbon cycle and climate. Model results show lower rates of carbon fixation and up to double the previously predicted atmospheric CO2 concentration due to a limited plant geographical range over the arid Pangea supercontinent. The Mesozoic dispersion of the continents increases modelled plant geographical range from 65% to > 90%, amplifying global CO2 removal, consistent with geological data. We demonstrate that plant geographical range likely exerted a major, under-explored control on long-term climate change.

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The most studied plant-fungal symbioses to date are the interactions between plants and arbuscular mycorrhizal (AM) fungi of the Glomeromycotina clade. Advancements in phylogenetics and microbial community profiling have distinguished a group of symbiosis-forming fungi that resemble AM fungi as belonging instead to the Mucoromycotina. These enigmatic fungi are now known as Mucoromycotina 'fine root endophytes' and could provide a means to understand the origins of plant-fungal symbioses. Most of our knowledge of the mechanisms of fungal symbiosis comes from investigations using AM fungi. Here, we argue that inclusion of Mucoromycotina fine root endophytes in future studies will expand our understanding of the mechanisms, evolution, and ecology of plant-fungal symbioses.

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Arbuscular mycorrhizal fungi colonize the roots of most plants, forming a near-ubiquitous symbiosis1 that is typically characterized by the bi-directional exchange of fungal-acquired nutrients for plant-fixed carbon.2 Mycorrhizal fungi can form below-ground networks3,4,5,6 with potential to facilitate the movement of carbon, nutrients, and defense signals across plant communities.7,8,9 The importance of neighbors in mediating carbon-for-nutrient exchange between mycorrhizal fungi and their plant hosts remains equivocal, particularly when other competing pressures for plant resources are present. We manipulated carbon source and sink strengths of neighboring pairs of host plants through exposure to aphids and tracked the movement of carbon and nutrients through mycorrhizal fungal networks with isotope tracers. When carbon sink strengths of both neighboring plants were increased by aphid herbivory, plant carbon supply to extraradical mycorrhizal fungal hyphae was reduced, but mycorrhizal phosphorus supply to both plants was maintained, albeit variably, across treatments. However, when the sink strength of only one plant in a pair was increased, carbon supply to mycorrhizal fungi was restored. Our results show that loss of carbon inputs into mycorrhizal fungal hyphae from one plant may be ameliorated through inputs of a neighbor, demonstrating the responsiveness and resilience of mycorrhizal plant communities to biological stressors. Furthermore, our results indicate that mycorrhizal nutrient exchange dynamics are better understood as community-wide interactions between multiple players rather than as strict exchanges between individual plants and their symbionts, suggesting that mycorrhizal C-for-nutrient exchange is likely based more on unequal terms of trade than the "fair trade" model for symbiosis.

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For more than 400 million years, mycorrhizal fungi and plants have formed partnerships that are crucial to the emergence and functioning of global ecosystems. The importance of these symbiotic fungi for plant nutrition is well established. However, the role of mycorrhizal fungi in transporting carbon into soil systems on a global scale remains under-explored. This is surprising given that ∼75% of terrestrial carbon is stored belowground and mycorrhizal fungi are stationed at a key entry point of carbon into soil food webs. Here, we analyze nearly 200 datasets to provide the first global quantitative estimates of carbon allocation from plants to the mycelium of mycorrhizal fungi. We estimate that global plant communities allocate 3.93 Gt CO2e per year to arbuscular mycorrhizal fungi, 9.07 Gt CO2e per year to ectomycorrhizal fungi, and 0.12 Gt CO2e per year to ericoid mycorrhizal fungi. Based on this estimate, 13.12 Gt of CO2e fixed by terrestrial plants is, at least temporarily, allocated to the underground mycelium of mycorrhizal fungi per year, equating to ∼36% of current annual CO2 emissions from fossil fuels. We explore the mechanisms by which mycorrhizal fungi affect soil carbon pools and identify approaches to increase our understanding of global carbon fluxes via plant-fungal pathways. Our estimates, although based on the best available evidence, are imperfect and should be interpreted with caution. Nonetheless, our estimations are conservative, and we argue that this work confirms the significant contribution made by mycorrhizal associations to global carbon dynamics. Our findings should motivate their inclusion both within global climate and carbon cycling models, and within conservation policy and practice.

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Most plants form mycorrhizal associations with mutualistic soil fungi. Through these partnerships, resources are exchanged including photosynthetically fixed carbon for fungal-acquired nutrients. Recently, it was shown that the diversity of associated fungi is greater than previously assumed, extending to Mucoromycotina fungi. These Mucoromycotina 'fine root endophytes' (MFRE) are widespread and generally co-colonise plant roots together with Glomeromycotina 'coarse' arbuscular mycorrhizal fungi (AMF). Until now, this co-occurrence has hindered the determination of the direct function of MFRE symbiosis. To overcome this major barrier, we developed new techniques for fungal isolation and culture and established the first monoxenic in vitro cultures of MFRE colonising a flowering plant, clover. Using radio- and stable-isotope tracers in these in vitro systems, we measured the transfer of 33 P, 15 N and 14 C between MFRE hyphae and the host plant. Our results provide the first unequivocal evidence that MFRE fungi are nutritional mutualists with a flowering plant by showing that clover gained both 15 N and 33 P tracers directly from fungus in exchange for plant-fixed C in the absence of other micro-organisms. Our findings and methods pave the way for a new era in mycorrhizal research, firmly establishing MFRE as both mycorrhizal and functionally important in terrestrial ecosystems.

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Earth's long-term climate may have profoundly influenced plant evolution. Local climatic factors, including water availability, light, and temperature, play a key role in plant physiology and growth, and have fluctuated substantially over geological time. However, the impact of these key climate variables on global plant biomass across the Phanerozoic has not yet been established. Linking climate and dynamic vegetation modelling, we identify two key 'windows of opportunity' during the Ordovician and Jurassic-Paleogene capable of supporting dramatic expansions of potential plant biomass. These conditions are driven by continental dispersion, paleolatitude of continental area and a lack of glaciation, allowing for an intense hydrological cycle and greater water availability. These windows coincide with the initial expansion of land plants and the later angiosperm radiation. Our findings suggest that the timing and expansion of habitable space for plants played an important role in plant evolution and diversification.

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All cereal crops engage in arbuscular mycorrhizal symbioses which can have profound, but sometimes deleterious, effects on plant nutrient acquisition and growth. The mechanisms underlying variable mycorrhizal responsiveness in cereals are not well characterised or understood. Adapting crops to realise mycorrhizal benefits could reduce fertiliser requirements and improve crop nutrition where fertiliser is unavailable. We conducted a phenotype screen in wheat (Triticum aestivum L.), using 99 lines of an Avalon × Cadenza doubled-haploid mapping population. Plants were grown with or without a mixed inoculum containing 5 species of arbuscular mycorrhizal fungi. Plant growth, nutrition and mycorrhizal colonisation were quantified. Plant growth response to inoculation was remarkably varied among lines, ranging from more than 30% decrease to 80% increase in shoot biomass. Mycorrhizal plants did not suffer decreasing shoot phosphorus concentration with increasing biomass as observed in their non-mycorrhizal counterparts. The extent to which mycorrhizal inoculation was beneficial for individual lines was negatively correlated with shoot biomass in the non-mycorrhizal state but was not correlated with the extent of mycorrhizal colonisation of roots. Highly variable mycorrhizal responsiveness among closely related wheat lines and the identification of several QTL for these traits suggests the potential to breed for improved crop-mycorrhizal symbiosis.

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Root exudates and rhizosheaths of attached soil are important features of growing roots. To elucidate factors involved in rhizosheath formation, wild-type (WT) barley (Hordeum vulgare L. cv. Pallas) and a root hairless mutant, bald root barley (brb), were investigated with a combination of physiological, biochemical, and immunochemical assays. When grown in soil, WT barley roots bound ∼5-fold more soil than brb per unit root length. High molecular weight (HMW) polysaccharide exudates of brb roots had less soil-binding capacity than those of WT root exudates. Carbohydrate and glycan monoclonal antibody analyses of HMW polysaccharide exudates indicated differing glycan profiles. Relative to WT plants, root exudates of brb had reduced signals for arabinogalactan-protein (AGP), extensin, and heteroxylan epitopes. In contrast, the root exudate of 2-week-old brb plants contained ∼25-fold more detectable xyloglucan epitope relative to WT. Root system immunoprints confirmed the higher levels of release of the xyloglucan epitope from brb root apices and root axes relative to WT. Epitope detection with anion-exchange chromatography indicated that the increased detection of xyloglucan in brb exudates was due to enhanced abundance of a neutral polymer. Conversely, brb root exudates contained decreased amounts of an acidic polymer, with soil-binding properties, containing the xyloglucan epitope and glycoprotein and heteroxylan epitopes relative to WT. We, therefore, propose that, in addition to physically structuring soil particles, root hairs facilitate rhizosheath formation by releasing a soil-binding polysaccharide complex.

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Mycorrhizal associations between fungi and plant roots have globally significant impacts on nutrient cycling. Mucoromycotina 'fine root endophytes' (MFRE) are a distinct and recently characterised group of mycorrhiza-forming fungi that associate with the roots of a range of host plant species. Given their previous misidentification and assignment as arbuscular mycorrhizal fungi (AMF) of the Glomeromycotina, it is now important to untangle the specific form and function of MFRE symbioses. In particular, relatively little is known about the nature of MFRE colonisation and its role in N uptake and transfer to host plants. Even less is known about the mechanisms by which MFRE access and assimilate N, and how this N is processed and subsequently exchanged with host plants for photosynthates. Here, we summarise and contrast the structures formed by MFRE and arbuscular mycorrhizal fungi in host plants as well as compare the N source preference of each mycorrhizal fungal group with what is currently known for MFRE N uptake. We compare the mechanisms of N assimilation and transfer to host plants utilised by the main groups of mycorrhizal fungi and hypothesise potential mechanisms for MFRE N assimilation and transfer, outlining directions for future research.

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Plants simultaneously interact with a range of biotrophic symbionts, ranging from mutualists such as arbuscular mycorrhizal fungi (AMF), to parasites such as the potato cyst nematode (PCN). The exchange of mycorrhizal-acquired nutrients for plant-fixed carbon (C) is well studied; however, the impact of competing symbionts remains underexplored. In this study, we examined mycorrhizal nutrient and host resource allocation in potato with and without AMF and PCN using radioisotope tracing, whilst determining the consequences of such allocation. The presence of PCN disrupted C for nutrient exchange between plants and AMF, with plant C overwhelmingly obtained by the nematodes. Despite this, AMF maintained transfer of nutrients on PCN-infected potato, ultimately losing out in their C for nutrient exchange with the host. Whilst PCN exploited the greater nutrient reserves to drive population growth on AMF-potato, the fungus imparted tolerance to allow the host to bear the parasitic burden. Our findings provide important insights into the belowground dynamics of plant-AMF symbioses, where simultaneous nutritional and nonnutritional benefits conferred by AMF to hosts and their parasites are seldom considered in plant community dynamics. Our findings suggest this may be a critical oversight, particularly in the consideration of C and nutrient flows in plant and soil communities.

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Production and heavy application of chemical-based fertilizers to maintain crop yields is unsustainable due to pollution from run-off, high CO2 emissions, and diminishing yield returns. Access to fertilizers will be limited in the future due to rising energy costs and dwindling rock phosphate resources. A growing number of companies produce and sell arbuscular mycorrhizal fungal (AMF) inoculants, intended to help reduce fertilizer usage by facilitating crop nutrient uptake through arbuscular mycorrhizas. However, their success has been variable. Here, we present information about the efficacy of a commercially available AMF inoculant in increasing AMF root colonization and fungal contribution to plant nutrient uptake, which are critical considerations within the growing AMF inoculant industry. Summary Arable agriculture needs sustainable solutions to reduce reliance on large inputs of nutrient fertilizers while continuing to improve crop yields. By harnessing arbuscular mycorrhizal symbiosis, there is potential to improve crop nutrient assimilation and growth without additional inputs, although the efficacy of commercially available mycorrhizal inocula in agricultural systems remains controversial.Using stable and radioisotope tracing, carbon-for-nutrient exchange between arbuscular mycorrhizal fungi and three modern cultivars of wheat was quantified in a non-sterile, agricultural soil, with or without the addition of a commercial mycorrhizal inoculant.While there was no effect of inoculum addition on above-ground plant biomass, there was increased root colonization by arbuscular mycorrhizal fungi and changes in community structure. Inoculation increased phosphorus uptake across all wheat cultivars by up to 30%, although this increase was not directly attributable to mycorrhizal fungi. Carbon-for-nutrient exchange between symbionts varied substantially between the wheat cultivars.Plant tissue phosphorus increased in inoculated plants potentially because of changes induced by inoculation in microbial community composition and/or nutrient cycling within the rhizosphere. Our data contribute to the growing consensus that mycorrhizal inoculants could play a role in sustainable food production systems of the future.

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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).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.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.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.

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Non-vascular plants associating with arbuscular mycorrhizal (AMF) and Mucoromycotina 'fine root endophyte' (MFRE) fungi derive greater benefits from their fungal associates under higher atmospheric [CO2] (a[CO2]) than ambient; however, nothing is known about how changes in a[CO2] affect MFRE function in vascular plants. We measured movement of phosphorus (P), nitrogen (N) and carbon (C) between the lycophyte Lycopodiella inundata and Mucoromycotina fine root endophyte fungi using 33P-orthophosphate, 15 N-ammonium chloride and 14CO2 isotope tracers under ambient and elevated a[CO2] concentrations of 440 and 800 ppm, respectively. Transfers of 33P and 15 N from MFRE to plants were unaffected by changes in a[CO2]. There was a slight increase in C transfer from plants to MFRE under elevated a[CO2]. Our results demonstrate that the exchange of C-for-nutrients between a vascular plant and Mucoromycotina FRE is largely unaffected by changes in a[CO2]. Unravelling the role of MFRE in host plant nutrition and potential C-for-N trade changes between symbionts under different abiotic conditions is imperative to further our understanding of the past, present and future roles of plant-fungal symbioses in ecosystems.

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Assuntos

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Associations formed between plants and arbuscular mycorrhizal (AM) fungi are characterized by the bi-directional exchange of fungal-acquired soil nutrients for plant-fixed organic carbon compounds. Mycorrhizal-acquired nutrient assimilation by plants may be symmetrically linked to carbon (C) transfer from plant to fungus or governed by sink-source dynamics. Abiotic factors, including atmospheric CO2 concentration ([CO2]), can affect the relative cost of resources traded between mutualists, thereby influencing symbiotic function. Whether biotic factors, such as insect herbivores that represent external sinks for plant C, impact mycorrhizal function remains unstudied. By supplying 33P to an AM fungus (Rhizophagus irregularis) and 14CO2 to wheat, we tested the impact of increasing C sink strength (i.e., aphid herbivory) and increasing C source strength (i.e., elevated [CO2]) on resource exchange between mycorrhizal symbionts. Allocation of plant C to the AM fungus decreased dramatically following exposure to the bird cherry-oat aphid (Rhopalosiphum padi), with high [CO2] failing to alleviate the aphid-induced decline in plant C allocated to the AM fungus. Mycorrhizal-mediated uptake of 33P by plants was maintained regardless of aphid presence or elevated [CO2], meaning insect herbivory drove asymmetry in carbon for nutrient exchange between symbionts. Here, we provide direct evidence that external biotic C sinks can limit plant C allocation to an AM fungus without hindering mycorrhizal-acquired nutrient uptake. Our findings highlight the context dependency of resource exchange between plants and AM fungi and suggest biotic factors-individually and in combination with abiotic factors-should be considered as powerful regulators of symbiotic function.

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An accurate understanding of the diversity and distribution of fungal symbioses in land plants is essential for mycorrhizal research. Here we update the seminal work of Wang and Qiu (Mycorrhiza 16:299-363, 2006) with a long-overdue focus on early-diverging land plant lineages, which were considerably under-represented in their survey, by examining the published literature to compile data on the status of fungal symbioses in liverworts, hornworts and lycophytes. Our survey combines data from 84 publications, including recent, post-2006, reports of Mucoromycotina associations in these lineages, to produce a list of at least 591 species with known fungal symbiosis status, 180 of which were included in Wang and Qiu (Mycorrhiza 16:299-363, 2006). Using this up-to-date compilation, we estimate that fewer than 30% of liverwort species engage in symbiosis with fungi belonging to all three mycorrhizal phyla, Mucoromycota, Basidiomycota and Ascomycota, with the last being the most widespread (17%). Fungal symbioses in hornworts (78%) and lycophytes (up to 100%) appear to be more common but involve only members of the two Mucoromycota subphyla Mucoromycotina and Glomeromycotina, with Glomeromycotina prevailing in both plant groups. Our fungal symbiosis occurrence estimates are considerably more conservative than those published previously, but they too may represent overestimates due to currently unavoidable assumptions.

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Arbuscular mycorrhizal fungi (AMF) form symbioses with most crops, potentially improving their nutrient assimilation and growth. The effects of cultivar and atmospheric CO2 concentration ([CO2 ]) on wheat-AMF carbon-for-nutrient exchange remain critical knowledge gaps in the exploitation of AMF for future sustainable agricultural practices within the context of global climate change. We used stable and radioisotope tracers (15 N, 33 P, 14 C) to quantify AMF-mediated nutrient uptake and fungal acquisition of plant carbon in three wheat (Triticum aestivum L.) cultivars. We grew plants under current ambient (440 ppm) and projected future atmospheric CO2 concentrations (800 ppm). We found significant 15 N transfer from fungus to plant in all cultivars, and cultivar-specific differences in total N content. There was a trend for reduced N uptake under elevated atmospheric [CO2 ]. Similarly, 33 P uptake via AMF was affected by cultivar and atmospheric [CO2 ]. Total P uptake varied significantly among wheat cultivars and was greater at the future than current atmospheric [CO2 ]. We found limited evidence of cultivar or atmospheric [CO2 ] effects on plant-fixed carbon transfer to the mycorrhizal fungi. Our results suggest that AMF will continue to provide a route for nutrient uptake by wheat in the future, despite predicted rises in atmospheric [CO2 ]. Consideration should therefore be paid to cultivar-specific AMF receptivity and function in the development of climate smart germplasm for the future.

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