Increased Carbon Partitioning to Secondary Metabolites Under Phosphorus Deficiency in Glycyrrhiza uralensis Fisch. Is Modulated by Plant Growth Stage and Arbuscular Mycorrhizal Symbiosis.

Phosphorus (P) is one of the macronutrients limiting plant growth. Plants regulate carbon (C) allocation and partitioning to cope with P deficiency, while such strategy could potentially be influenced by plant growth stage and arbuscular mycorrhizal (AM) symbiosis. In a greenhouse pot experiment using licorice (Glycyrrhiza uralensis) as the host plant, we investigated C allocation belowground and partitioning in roots of P-limited plants in comparison with P-sufficient plants under different mycorrhization status in two plant growth stages. The experimental results indicated that increased C allocation belowground by P limitation was observed only in non-AM plants in the early growth stage. Although root C partitioning to secondary metabolites (SMs) in the non-AM plants was increased by P limitation as expected, trade-off patterns were different between the two growth stages, with C partitioning to SMs at the expense of non-structural carbohydrates (NSCs) in the early growth stage but at the expense of root growth in the late growth stage. These changes, however, largely disappeared because of AM symbiosis, where more root C was partitioned to root growth and AM fungus without any changes in C allocation belowground and partitioning to SMs under P limitations. The results highlighted that besides assisting with plant P acquisition, AM symbiosis may alter plant C allocation and partitioning to improve plant tolerance to P deficiency.

Contrasting Nitrogen Fertilisation Rates Alter Mycorrhizal Contribution to Barley Nutrition in a Field Trial.

Controlled environment studies show that arbuscular mycorrhizal fungi (AMF) may contribute to plant nitrogen (N) uptake, but the role of these near-ubiquitous symbionts in crop plant N nutrition under natural field conditions remains largely unknown. In a field trial, we tested the effects of N fertilisation and barley (Hordeum vulgare L.) cultivar identity on the contribution of AMF to barley N uptake using 15N tracers added to rhizosphere soil compartments. AMF were shown capable of significantly increasing plant 15N acquisition from root exclusion zones, and this was influenced by nitrogen addition type, N fertiliser application rate and barley cultivar identity. Our data demonstrate a previously overlooked potential route of crop plant N uptake which may be influenced substantially and rapidly in response to shifting agricultural management practices.

Aphids Influence Soil Fungal Communities in Conventional Agricultural Systems.

Arbuscular mycorrhizal fungi (AMF) form symbioses with the roots of most plant species, including cereals. AMF can increase the uptake of nutrients including nitrogen (N) and phosphorus (P), and of silicon (Si) as well as increase host resistance to various stresses. Plants can simultaneously interact with above-ground insect herbivores such as aphids, which can alter the proportion of plant roots colonized by AMF. However, it is unknown if aphids impact the structure of AMF communities colonizing plants or the extent of the extraradical mycelium produced in the soil, both of which can influence the defensive and nutritional benefit a plant derives from the symbiosis. This study investigated the effect of aphids on the plant-AMF interaction in a conventionally managed agricultural system. As plants also interact with other soil fungi, the non-AMF fungal community was also investigated. We hypothesized that aphids would depress plant growth, and reduce intraradical AMF colonization, soil fungal hyphal density and the diversity of AM and non-AM fungal communities. To test the effects of aphids, field plots of barley enclosed with insect proof cages were inoculated with Sitobion avenae or remained uninoculated. AMF specific and total fungal amplicon sequencing assessed root fungal communities 46 days after aphid addition. Aphids did not impact above-ground plant biomass, but did increase the grain N:P ratio. Whilst aphid presence had no impact on AMF intraradical colonization, soil fungal hyphal length density, or AMF community characteristics, there was a trend for the aphid treatment to increase vesicle numbers and the relative abundance of the AMF family Gigasporaceae. Contrary to expectations, the aphid treatment also increased the evenness of the total fungal community. This suggests that aphids can influence soil communities in conventional arable systems, a result that could have implications for multitrophic feedback loops between crop pests and soil organisms across the above-below-ground interface.

Arbuscular Mycorrhizal Fungi and Plant Chemical Defence: Effects of Colonisation on Aboveground and Belowground Metabolomes.

Arbuscular mycorrhizal fungal (AMF) colonisation of plant roots is one of the most ancient and widespread interactions in ecology, yet the systemic consequences for plant secondary chemistry remain unclear. We performed the first metabolomic investigation into the impact of AMF colonisation by Rhizophagus irregularis on the chemical defences, spanning above- and below-ground tissues, in its host-plant ragwort (Senecio jacobaea). We used a non-targeted metabolomics approach to profile, and where possible identify, compounds induced by AMF colonisation in both roots and shoots. Metabolomics analyses revealed that 33 compounds were significantly increased in the root tissue of AMF colonised plants, including seven blumenols, plant-derived compounds known to be associated with AMF colonisation. One of these was a novel structure conjugated with a malonyl-sugar and uronic acid moiety, hitherto an unreported combination. Such structural modifications of blumenols could be significant for their previously reported functional roles associated with the establishment and maintenance of AM colonisation. Pyrrolizidine alkaloids (PAs), key anti-herbivore defence compounds in ragwort, dominated the metabolomic profiles of root and shoot extracts. Analyses of the metabolomic profiles revealed an increase in four PAs in roots (but not shoots) of AMF colonised plants, with the potential to protect colonised plants from below-ground organisms.

Arbuscular mycorrhizal fungi reduce nitrous oxide emissions from N2 O hotspots.

Nitrous oxide (N2 O) is a potent, globally important, greenhouse gas, predominantly released from agricultural soils during nitrogen (N) cycling. Arbuscular mycorrhizal fungi (AMF) form a mutualistic symbiosis with two-thirds of land plants, providing phosphorus and/or N in exchange for carbon. As AMF acquire N, it was hypothesized that AMF hyphae may reduce N2 O production. AMF hyphae were either allowed (AMF) or prevented (nonAMF) access to a compartment containing an organic matter and soil patch in two independent microcosm experiments. Compartment and patch N2 O production was measured both before and after addition of ammonium and nitrate. In both experiments, N2 O production decreased when AMF hyphae were present before inorganic N addition. In the presence of AMF hyphae, N2 O production remained low following ammonium application, but increased in the nonAMF controls. By contrast, negligible N2 O was produced following nitrate application to either AMF treatment. Thus, the main N2 O source in this system appeared to be via nitrification, and the production of N2 O was reduced in the presence of AMF hyphae. It is hypothesized that AMF hyphae may be outcompeting slow-growing nitrifiers for ammonium. This has significant global implications for our understanding of soil N cycling pathways and N2 O production.

Using the Evidence-Based Practice Service Nursing Bundle to Increase Patient Satisfaction.

ABSTRACTINTRODUCTION: Patient satisfaction and patient experience goals are often linked to financial consequences. Although the link does exist, the bottom line is not only about money; it's about providing a quality experience for ED patients and creating an environment that engages staff. Evidence-based practice (EBP) strategies that have positive impact on patient perceptions of their ED care and increased satisfaction ratings include AIDET, Hourly Rounding, and Bedside Shift Report, which incorporate updates of test results and explanations of events occurring during a patient's visit. In addition to these outcomes, Hourly Rounding and Bedside Shift Report have been linked to patient safety improvements. Combining these strategies, our team created and implemented the service nursing bundle as a quality improvement (QI) initiative, with the goal of having a positive effect on patient experiences in emergency departments, represented by at least a 5% increase in overall quality-of-care ranking and ratings. METHODS: This QI project involved comparing professional research consultant (PRC) patient- satisfaction phone survey ratings from patients before and after ED staff members completed a 1-hour service nursing bundle class. In addition to the patient-satisfaction ratings, 1,104 audits evaluating staff use of the service bundle implementation were collected over an 8-week period.reading document: RESULTS: The random observational audits showed the adoption of the service nursing bundle as staff compliance started at 65% in week 1 compared with 100% by week 8. Before intervention (July 2015): 50% of patients rated their overall quality of care as excellent, yielding a benchmark ranking of 42.5 percentile. Postservice bundle education implementation (September 2015): 60% of patients rated their overall quality of care as excellent, increasing our ranking to the 85.5 percentile. The postservice bundle group was 1.5 times more likely to respond "excellent" to all 5 survey questions, which was statistically significant (z =2.82, P = 0.004). The patients' perceptions of total time spent in the emergency department and ratings of "excellent" revealed a significant statistical difference (before: 35.0%, after: 49.5%, X2 (1) = 4.24, P < 0.05). DISCUSSION: With the implementation of the bundle, our emergency department experienced an 11.8% increase in the number of patients rating their overall quality of care as excellent. This upsurge resulted in a 40% increase in overall quality of care, propelling our emergency department's ranking to the 85th percentile.

Carbon and phosphorus exchange may enable cooperation between an arbuscular mycorrhizal fungus and a phosphate-solubilizing bacterium.

Arbuscular mycorrhizal fungi (AMF) transfer plant photosynthate underground which can stimulate soil microbial growth. In this study, we examined whether there was a potential link between carbon (C) release from an AMF and phosphorus (P) availability via a phosphate-solubilizing bacterium (PSB). We investigated the outcome of the interaction between the AMF and the PSB by conducting a microcosm and two Petri plate experiments. An in vitro culture experiment was also conducted to determine the direct impact of AMF hyphal exudates on growth of the PSB. The AMF released substantial C to the environment, triggering PSB growth and activity. In return, the PSB enhanced mineralization of organic P, increasing P availability for the AMF. When soil available P was low, the PSB competed with the AMF for P, and its activity was not stimulated by the fungus. When additional P was added to increase soil available P, the PSB enhanced AMF hyphal growth, and PSB activity was also stimulated by the fungus. Our results suggest that an AMF and a free-living PSB interacted to the benefit of each other by providing the C or P that the other microorganism required, but these interactions depended upon background P availability.

Resolving the 'nitrogen paradox' of arbuscular mycorrhizas: fertilization with organic matter brings considerable benefits for plant nutrition and growth.

Arbuscular mycorrhizal fungi (AMF) can transfer nitrogen (N) to host plants, but the ecological relevance is debated, as total plant N and biomass do not generally increase. The extent to which the symbiosis is mutually beneficial is thought to rely on the stoichiometry of N, phosphorus (P) and carbon (C) availability. While inorganic N fertilization has been shown to elicit strong mutualism, characterized by improved plant and fungal growth and mineral nutrition, similar responses following organic N addition are lacking. Using a compartmented microcosm experiment, we determined the significance to a mycorrhizal plant of placing a (15) N-labelled, nitrogen-rich patch of organic matter in a compartment to which only AMF hyphae had access. Control microcosms denied AMF hyphal access to the patch compartment. When permitted access to the patch compartment, the fungus proliferated extensively in the patch and transferred substantial quantities of N to the plant. Moreover, our data demonstrate that allowing hyphal access to an organic matter patch enhanced total plant N and P contents, with a simultaneous and substantial increase in plant biomass. Furthermore, we demonstrate that organic matter fertilization of arbuscular mycorrhizal plants can foster a mutually beneficial symbiosis based on nitrogen transfer, a phenomenon previously thought irrelevant.

Fishing for nutrients in heterogeneous landscapes: modelling plant growth trade-offs in monocultures and mixed communities.

The problem of how best to find and exploit essential resources, the quality and locations of which are unknown, is common throughout biology. For plants, the need to grow an efficient root system so as to acquire patchily distributed soil nutrients is typically complicated by competition between plants, and by the costs of maintaining the root system. Simple mechanistic models for root growth can help elucidate these complications, and here we argue that these models can be usefully informed by models initially developed for foraging fish larvae. Both plant and fish need to efficiently search a spatio-temporally variable environment using simple algorithms involving only local information, and both must perform this task against a backdrop of intra- and inter-specific competition and background mortality. Here we develop these parallels by using simple stochastic models describing the growth and efficiency of four contrasting idealized root growth strategies. We show that plants which grow identically in isolation in homogeneous substrates will typically perform very differently when grown in monocultures, in heterogeneous nutrient landscapes and in mixed-species competition. In particular, our simulations show a consistent result that plants which trade-off rapid growth in favour of a more efficient and durable root system perform better, both on average and in terms of the best performing individuals, than more rapidly growing ephemeral root systems. Moreover, when such slower growing but more efficient plants are grown in competition, the overall community productivity can exceed that of the constituent monocultures. These findings help to disentangle many of the context-dependent behaviours seen in the experimental literature, and may form a basis for future studies at the level of complex population dynamics and life history evolution.

Phosphate concentration and arbuscular mycorrhizal colonisation influence the growth, yield and expression of twelve PHT1 family phosphate transporters in foxtail millet (Setaria italica).

Phosphorus (P) is an essential element which plays several key roles in all living organisms. Setaria italica (foxtail millet) is a model species for panacoid grasses including several millet species widely grown in arid regions of Asia and Africa, and for the bioenergy crop switchgrass. The growth responses of S. italica to different levels of inorganic phosphate (Pi) and to colonisation with the arbuscular mycorrhizal fungus Funneliformis mosseae (syn. Glomus mosseae) were studied. Phosphate is taken up from the environment by the PHT1 family of plant phosphate transporters, which have been well characterized in several plant species. Bioinformatic analysis identified 12 members of the PHT1 gene family (SiPHT1;1-1;12) in S. italica, and RT and qPCR analysis showed that most of these transporters displayed specific expression patterns with respect to tissue, phosphate status and arbuscular mycorrhizal colonisation. SiPHT1;2 was found to be expressed in all tissues and in all growth conditions tested. In contrast, expression of SiPHT1;4 was induced in roots after 15 days growth in hydroponic medium of low Pi concentration. Expression of SiPHT1;8 and SiPHT1;9 in roots was selectively induced by colonisation with F. mosseae. SiPHT1;3 and SiPHT1;4 were found to be predominantly expressed in leaf and root tissues respectively. Several other transporters were expressed in shoots and leaves during growth in low Pi concentrations. This study will form the basis for the further characterization of these transporters, with the long term goal of improving the phosphate use efficiency of foxtail millet.

Interactions between arbuscular mycorrhizal fungi and organic material substrates.

Arbuscular mycorrhizal (AM) associations are widespread and form between ca. two-thirds of all land plants and fungi in the phylum Glomeromycota. The association is a mutualistic symbiosis with the fungi enhancing nutrient capture for the plant while obtaining carbon in return. Although arbuscular mycorrhizal fungi (AMF) lack any substantial saprophytic capability they do preferentially associate with various organic substrates and respond by hyphal proliferation, indicating the fungus derives a benefit from the organic substrate. AMF may also enhance decomposition of the organic material. The benefit to the host plant of this hyphal proliferation is not always apparent, particularly regarding nitrogen (N) transfer, and there may be circumstances under which both symbionts compete for the N released given both have a large demand for N. The results of various studies examining AMF responses to organic substrates and the interactions with other members of the soil community will be discussed.

An arbuscular mycorrhizal fungus significantly modifies the soil bacterial community and nitrogen cycling during litter decomposition.

Arbuscular mycorrhizal fungi (AMF) perform an important ecosystem service by improving plant nutrient capture from soil, yet little is known about how AMF influence soil microbial communities during nutrient uptake. We tested whether an AMF modifies the soil microbial community and nitrogen cycling during litter decomposition. A two-chamber microcosm system was employed to create a root-free soil environment to control AMF access to (13) C- and (15) N-labelled root litter. Using a 16S rRNA gene microarray, we documented that approximately 10% of the bacterial community responded to the AMF, Glomus hoi. Taxa from the Firmicutes responded positively to AMF, while taxa from the Actinobacteria and Comamonadaceae responded negatively to AMF. Phylogenetic analyses indicate that AMF may influence bacterial community assembly processes. Using nanometre-scale secondary ion mass spectrometry (NanoSIMS) we visualized the location of AMF-transported (13) C and (15) N in plant roots. Bulk isotope ratio mass spectrometry revealed that the AMF exported 4.9% of the litter (15) N to the host plant (Plantago lanceolata L.), and litter-derived (15) N was preferentially exported relative to litter-derived (13) C. Our results suggest that the AMF primarily took up N in the inorganic form, and N export is one mechanism by which AMF could modify the soil microbial community and decomposition processes.

Interactions between an arbuscular mycorrhizal fungus and a soil microbial community mediating litter decomposition.

We investigated arbuscular mycorrhizal fungi (AMF) alteration of microbial mediation of litter decomposition. AMF (Glomus hoi) were either allowed access to or excluded from Plantago lanceolata L. root litter embedded in soil; litter was labeled with either (13) C only or (13) C and (15) N. Plant N uptake was significantly increased if AMF accessed the litter, and (15) N analysis of the plant material indicated that 2-3% of plant N originated from litter. Succession of the soil community mediating decomposition was assessed by phospholipid fatty acids (PLFA) combined with (13) C-PLFA. During the first 21 days of decomposition, saprotrophic fungi and Gram-negative bacteria were the dominant consumers of litter C. As decomposition progressed however, (13) C content of the fungal biomarkers declined substantially, and Gram-negative and Gram-positive bacteria became the primary reservoirs of labeled litter C. The putative PLFA marker for AMF (16:1ω5c) originated primarily from non-AMF sources. In AMF-invaded root litter, Gram-negative, Gram-positive, and 16:1ω5c markers became less (13) C-enriched relative to their counterparts in non-AMF-invaded microcosms during active decomposition. These patterns of (13) C: (12) C enrichment may result from AMF supply of (12) C from the plant to the decomposing soil microbial community; such C inputs could alter the microbial mediation of litter decomposition.

Growth and symbiotic effectiveness of an arbuscular mycorrhizal fungus in organic matter in competition with soil bacteria.

Arbuscular mycorrhizal (AM) fungi can enhance the rate of decomposition of organic material, and can acquire nitrogen (N) from organic sources, although they are not saprotrophs. These fungi may instead indirectly influence decomposition through interactions with other soil microorganisms. We investigated the impact of both AM hyphae and a bacterial filtrate on N capture by a host plant from sterilized organic material (Lolium perenne shoots dual labelled with (15) N and (13) C), using compartmented microcosms. The addition of a bacterial filtrate considerably suppressed AM hyphal growth in the patch and reduced the root phosphorus content, demonstrating that bacterial populations can reduce symbiotic effectiveness. In contrast, AM hyphae had only a limited impact on bacterial community structure. Uptake of (15) N greatly exceeded that of (13) C, demonstrating that fungi acquired N in an inorganic form. We also examined the ability of AM fungi in gnotobiotic hairy root culture to acquire N directly from organic materials of varying complexities (glutamic acid, urea, bacterial lysate and L. perenne shoots). AM colonization did not enhance root N capture from these materials, although the bacterial lysate reduced both total AM colonization and arbuscule frequency. Collectively, these data demonstrate antagonistic interactions between AM fungi and bacteria that reflect resource competition for decomposition products.

Substantial nitrogen acquisition by arbuscular mycorrhizal fungi from organic material has implications for N cycling.

Arbuscular mycorrhizal (AM) fungi are obligate biotrophs that acquire carbon (C) solely from host plants. AM fungi can proliferate hyphae in, and acquire nitrogen (N) from, organic matter. Although they can transfer some of that N to plants, we tested the hypothesis that organic matter is an important N source for the AM fungi themselves. We grew pairs of plants with and without the AM fungus Glomus hoi in microcosms that allowed only the fungus access to a 15N/13C-labeled organic patch; in some cases, one plant was shaded to reduce C supply to the fungus. The fungal hyphae proliferated vigorously in the patch, irrespective of shading, and increased plant growth and N content; approximately 3% of plant N came from the patch. The extraradical mycelium of the fungus was N-rich (3-5% N) and up to 31% of fungal N came from the patch, confirming the hypothesis. The fungus acquired N as decomposition products, because hyphae were not 13C-enriched. In a second experiment, hyphae of both G. hoi and Glomus mosseae that exploited an organic material patch were also better able to colonize a new host plant, demonstrating a fungal growth response. These findings show that AM fungi can obtain substantial amounts of N from decomposing organic materials and can enhance their fitness as a result. The large biomass and high N demand of AM fungi means that they represent a global N pool equivalent in magnitude to fine roots and play a substantial and hitherto overlooked role in the nitrogen cycle.

Mycorrhizas and biomass crops: opportunities for future sustainable development.

Central to soil health and plant productivity in natural ecosystems are in situ soil microbial communities, of which mycorrhizal fungi are an integral component, regulating nutrient transfer between plants and the surrounding soil via extensive mycelial networks. Such networks are supported by plant-derived carbon and are likely to be enhanced under coppiced biomass plantations, a forestry practice that has been highlighted recently as a viable means of providing an alternative source of energy to fossil fuels, with potentially favourable consequences for carbon mitigation. Here, we explore ways in which biomass forestry, in conjunction with mycorrhizal fungi, can offer a more holistic approach to addressing several topical environmental issues, including 'carbon-neutral' energy, ecologically sustainable land management and CO(2) sequestration.

Temperature dependence of respiration in roots colonized by arbuscular mycorrhizal fungi.

* The arbuscular mycorrhizal (AM) symbiosis is ubiquitous, and the fungus represents a major pathway for carbon movement in the soil-plant system. Here, we investigated the impacts of AM colonization of Plantago lanceolata and temperature on the regulation of root respiration (R). * Warm-grown AM plants exhibited higher rates of R than did nonAM plants, irrespective of root mass. AM plants exhibited higher maximal rates of R (R(max)-R measured in the presence of an uncoupler and exogenous substrate) and greater proportional use of R(max) as a result of increased energy demand and/or substrate supply. The higher R values exhibited by AM plants were not associated with higher maximal rates of cytochrome c oxidase (COX) or protein abundance of either the COX or the alternative oxidase. * Arbuscular mycorrhizal colonization had no effect on the short-term temperature dependence (Q(10)) of R. Cold-acclimated nonAM plants exhibited higher rates of R than their warm-grown nonAM counterparts. By contrast, chilling had a negligible effect on R of AM-plants. Thus, AM plants exhibited less cold acclimation than their nonAM counterparts. * Overall, these results highlight the way in which AM colonization alters the underlying components of respiratory metabolism and the response of root R to sustained changes in growth temperature.