Aphid Herbivory Drives Asymmetry in Carbon for Nutrient Exchange between Plants and an Arbuscular Mycorrhizal Fungus

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

A commercial arbuscular mycorrhizal inoculum increases root colonization across wheat cultivars but does not increase assimilation of mycorrhiza‐acquired nutrients

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

The distribution and evolution of fungal symbioses in ancient lineages of land plants

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

Nutrient acquisition by symbiotic fungi governs Palaeozoic climate transition

Fossil evidence from the Rhynie chert indicates that early land plants, which evolved in a high-CO2 atmosphere during the Palaeozoic Era, hosted diverse fungal symbionts. It is hypothesized that the rise of early non-vascular land plants, and the later evolution of roots and vasculature, drove the long-term shift towards a high-oxygen, low CO2 climate that eventually permitted the evolution of mammals and, ultimately, humans. However, very little is known about the productivity of the early terrestrial biosphere, which depended on the acquisition of the limiting nutrient phosphorus via fungal symbiosis. Recent laboratory experiments have shown that plant–fungal symbiotic function is specific to fungal identity, with carbon-for-phosphorus exchange being either enhanced or suppressed under superambient CO2. By incorporating these experimental findings into a biogeochemical model, we show that the differences in these symbiotic nutrient acquisition strategies could greatly alter the plant-driven changes to climate, allowing drawdown of CO2 to glacial levels, and altering the nature of the rise of oxygen. We conclude that an accurate depiction of plant–fungal symbiotic systems, informed by high-CO2 experiments, is key to resolving the question of how the first terrestrial ecosystems altered our planet

REI:An integrated measure for software reusability

To capitalize upon the benefits of software reuse, an efficient selection among candidate reusable assets should be performed in terms of functional fitness and adaptability. The reusability of assets is usually measured through reusability indices. However, these do not capture all facets of reusability, such as structural characteristics, external quality attributes, and documentation. In this paper, we propose a reusability index (REI) as a synthesis of various software metrics and evaluate its ability to quantify reuse, based on IEEE Standard on Software Metrics Validity. The proposed index is compared with existing ones through a case study on 80 reusable open-source assets. To illustrate the applicability of the proposed index, we performed a pilot study, where real-world reuse decisions have been compared with decisions imposed by the use of metrics (including REI). The results of the study suggest that the proposed index presents the highest predictive and discriminative power; it is the most consistent in ranking reusable assets and the most strongly correlated to their levels of reuse. The findings of the paper are discussed to understand the most important aspects in reusability assessment (interpretation of results), and interesting implications for research and practice are provided

Xyloglucan is released by plants and promotes soil particle aggregation.

Soil is a crucial component of the biosphere and is a major sink for organic carbon. Plant roots are known to release a wide range of carbon-based compounds into soils, including polysaccharides, but the functions of these are not known in detail. Using a monoclonal antibody to plant cell wall xyloglucan, we show that this polysaccharide is secreted by a wide range of angiosperm roots, and relatively abundantly by grasses. It is also released from the rhizoids of liverworts, the earliest diverging lineage of land plants. Using analysis of water-stable aggregate size, dry dispersion particle analysis and scanning electron microscopy, we show that xyloglucan is effective in increasing soil particle aggregation, a key factor in the formation and function of healthy soils. To study the possible roles of xyloglucan in the formation of soils, we analysed the xyloglucan contents of mineral soils of known age exposed upon the retreat of glaciers. These glacial forefield soils had significantly higher xyloglucan contents than detected in a UK grassland soil. We propose that xyloglucan released from plant rhizoids/roots is an effective soil particle aggregator and may, in this role, have been important in the initial colonization of land

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

Functional complementarity of ancient plant–fungal mutualisms: contrasting nitrogen, phosphorus and carbon exchanges between Mucoromycotina and Glomeromycotina fungal symbionts of liverworts

Liverworts, which are amongst the earliest divergent plant lineages and important ecosystem pioneers, often form nutritional mutualisms with arbuscular mycorrhiza‐forming Glomeromycotina and fine‐root endophytic Mucoromycotina fungi, both of which coevolved with early land plants. Some liverworts, in common with many later divergent plants, harbour both fungal groups, suggesting these fungi may complementarily improve plant access to different soil nutrients. We tested this hypothesis by growing liverworts in single and dual fungal partnerships under a modern atmosphere and under 1500 ppm [CO2], as experienced by early land plants. Access to soil nutrients via fungal partners was investigated with 15N‐labelled algal necromass and 33P orthophosphate. Photosynthate allocation to fungi was traced using 14CO2. Only Mucoromycotina fungal partners provided liverworts with substantial access to algal 15N, irrespective of atmospheric CO2 concentration. Both symbionts increased 33P uptake, but Glomeromycotina were often more effective. Dual partnerships showed complementarity of nutrient pool use and greatest photosynthate allocation to symbiotic fungi. We show there are important functional differences between the plant–fungal symbioses tested, providing new insights into the functional biology of Glomeromycotina and Mucoromycotina fungal groups that form symbioses with plants. This may explain the persistence of the two fungal lineages in symbioses across the evolution of land plants

Field's Logic of Truth

Saving Truth from Paradox is a re-exciting development. The 70s and 80s were a time of excitement among people working on the semantic paradoxes. There were continual formal developments, with the constant hope that these results would yield deep insights. The enthusiasm wore off, however, as people became more cognizant of the disparity between what they had accomplished, impressive as it was, and what they had hoped to accomplish. They moved onto other problems that they hoped would prove more yielding. That, at least, was how it seemed to me, so I was delighted to see a dramatically new formal development that is likely to rekindle our enthusiasm

A mycorrhizal revolution.

It has long been postulated that symbiotic fungi facilitated plant migrations onto land through enhancing the scavenging of mineral nutrients and exchanging these for photosynthetically fixed organic carbon. Today, land plant-fungal symbioses are both widespread and diverse. Recent discoveries show that a variety of potential fungal associates were likely available to the earliest land plants, and that these early partnerships were probably affected by changing atmospheric CO2 concentrations. Here, we evaluate current hypotheses and knowledge gaps regarding early plant-fungal partnerships in the context of newly discovered fungal mutualists of early and more recently evolved land plants and the rapidly changing views on the roles of plant-fungal symbioses in the evolution and ecology of the terrestrial biosphere