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The accumulation of phosphorus (P) in plants increases their biomass and resistance/tolerance to organic pollutants. Both characteristics are mandatory for the utilization of plants in phytoremediation. Arbuscular mycorrhizal (AM) fungi improve plant P nutrition, and thus growth. However, only a few studies have focused on the dynamics of inorganic P (Pi) uptake in AM fungal-colonized plants in the presence of organic pollutants. Indeed, most of the results so far were obtained after harvesting the plants, thus by evaluating P concentration and content at a single time point. Here, we investigated the effects of the AM fungus Rhizophagus irregularis MUCL 41833 on the short-term Pi uptake dynamics of Medicago truncatula plants grown in the presence of benzo[a]pyrene (B[a]P), a polyaromatic hydrocarbon (PAH) frequently found in polluted soils. The study was conducted using a non-destructive circulatory semi-hydroponic cultivation system to investigate the short-term Pi depletion from a nutrient solution and as a corollary, the Pi uptake by the AM fungal-colonized and non-colonized plants. The growth, P concentration, and content of plants were also evaluated at harvest. The presence of B[a]P neither impacted the development of the AM fungus in the roots nor the plant growth and Pi uptake, suggesting a marked tolerance of both organisms to B[a]P pollution. A generally higher Pi uptake coupled with a higher accumulation of P in shoots and roots was noticed in AM fungal-colonized plants as compared to the non-colonized controls, irrespective of the presence or absence of B[a]P. Therefore, fungal-colonized plants showed the best growth. Furthermore, the beneficial effect provided by the presence of the AM fungus in roots was similar in presence or absence of B[a]P, thus opening the door for potential utilization in phytomanagement of PAH-polluted soils.

Insect herbivore feeding causes mechanical damage to plants, which can activate plant defense responses. Whether symbiosis with beneficial microorganisms can enhance the responses of plants to mechanical damage is of importance for plant anti-herbivore resistance. In this study, defense responses of tomato (Lycopersicon esculentum) plants to mechanical wounding was investigated after the tomato roots being infected by arbuscular mycorrhizal fungus (AMF) Funneliformis mosseae. The results showed that in response to leaf mechanical wounding, the activities of phenylalanine ammonia-lyase (PAL), superoxide dismutase (SOD), peroxidase (POD), polyphenol oxidase (PPO) and catalase (CAT) in the leaves of tomato pre-inoculated with AMF (FD), as well as transcript levels of genes encoding phenylalanine ammonia lyase (PAL) and β-1,3-glucanase (PR2) in the leaves and roots were significantly higher in relative to sole mechanical wounding (D), sole mycorrhizal inoculation (F), and control without mechanical wounding and mycorrhizal inoculation (CK). Although the activities of protective enzyme and transcript levels of the two defense-related genes were induced in the plants of sole mechanical wounding (D) and sole mycorrhizal inoculation (F), the induction was faster and stronger in the plants with leaf mechanical wounding and mycorrhizal pre-inoculation (FD). Our findings indicated that arbuscular mycorrhizal colonization could prime quicker and stronger defense responses of tomato plants to mechanical damage.

The application of 33 P or 32 P isotopes to directly trace phosphorus (P) uptake during arbuscular mycorrhizal (AM) symbiosis is limited by the radioactivity of the two P isotopes, especially under field conditions. A potential alternative method for tracing P uptake in plant-soil systems relies on the analysis of the stable oxygen (O) isotopes of ortho-phosphate (Pi); however, little is known about the fate of the P-O bond during Pi uptake in AM symbioses. This study investigated whether the abundance of 18 O in Pi extracted from the shoots of maize increased after 18 O-labeled Pi added to soil was taken up by either roots of maize or AM extraradical hyphae. A two-compartment culture system, consisting of a root and AM hyphal compartment (RHC, including both roots and AM hyphae) and an AM hyphal compartment (HC, including only hyphae) was designed, and the AM fungus Funneliformis mosseae was used to inoculate the roots of maize. Our results indicated that the abundance of 18 O in Pi extracted from the maize shoots increased significantly 3 months after the addition of 18 O-labeled Pi to the soil in the pots which only contained roots. The abundance of 18 O was much lower than expected, however, which suggests a great majority of 18 O in labeled Pi was lost in the soil or during Pi metabolism in the shoots of maize. The abundance of 18 O in Pi extracted from the maize shoots did not increase 3 months after 18 O-labeled Pi was added to the HC, and therefore, loss of 18 O in labeled Pi may also occur during Pi metabolism in AM hyphae. Use of 18 O-labeled Pi as a qualitative tracer of P uptake during AM symbiosis appears unfeasible for such a long-term (3 months) experiment, although it should be investigated in a short-term labeling experiment.

Arbuscular mycorrhizas are widespread in land plants including liverworts, some of the closest living relatives of the first plants to colonize land 500 million years ago (MYA). Previous investigations reported near-exclusive colonization of liverworts by the most recently evolved arbuscular mycorrhizal fungi, the Glomeraceae, indicating a recent acquisition from flowering plants at odds with the widely held notion that arbuscular mycorrhizal-like associations in liverworts represent the ancestral symbiotic condition in land plants. We performed an analysis of symbiotic fungi in 674 globally collected liverworts using molecular phylogenetics and electron microscopy. Here, we show every order of arbuscular mycorrhizal fungi colonizes early-diverging liverworts, with non-Glomeraceae being at least 10 times more common than in flowering plants. Arbuscular mycorrhizal fungi in liverworts and other ancient plant lineages (hornworts, lycopods, and ferns) were delimited into 58 taxa and 36 singletons, of which at least 43 are novel and specific to liverworts. The discovery that early plant lineages are colonized by early-diverging fungi supports the hypothesis that arbuscular mycorrhizas are an ancestral symbiosis for all land plants., (© 2018 The Authors.)

In a pot experiment, Medicago sativa inoculated with/without arbuscular mycorrhizal (AM) fungus Rhizophagus irregularis were grown in four levels (0, 10, 25, and 75 mg/kg) of arsenic (As)-polluted soil to investigate the influences of AM symbiosis on plant As tolerance. The results showed that mycorrhizal inoculation significantly increased plant biomass, while As addition decreased mycorrhizal colonization and hyphal length density. Mycorrhizal inoculation dramatically improved plant phosphorus (P) nutrition, restricted As uptake and retained more As in roots by upregulating the expression of the AM-induced P transporter gene MsPT4 and the metallothionein gene MsMT2. High soil As content downregulated MsPT4 expression. Dimethylarsenic acid (DMA) was detected only in the shoots of mycorrhizal plants, indicating that AM fungi likely play an essential role in As detoxification by biological methylation. The present investigation allowed deeper insights into the As detoxification mechanisms of AM associations and demonstrated the important role of AM fungi in plant resistance under As-contaminated conditions., (Copyright © 2018 Elsevier Inc. All rights reserved.)

In this study, we investigated the effects of the arbuscular mycorrhizal fungi (AMF) Funneliformis mosseae and Diversispora spurcum on the growth, antioxidant physiology, and uptake of phosphorus (P), sulfur (S), lead (Pb), zinc (Zn), cadmium (Cd), and arsenic (As) by maize (Zea mays L.) grown in heavy metal-polluted soils though a potted plant experiment. F. mosseae significantly increased the plant chlorophyll a content, height, and biomass; decreased the H 2 O 2 and malondialdehyde (MDA) contents; and enhanced the superoxide dismutase (SOD) and catalase (CAT) activities and the total antioxidant capacity (T-AOC) in maize leaves; this effect was not observed with D. spurcum. Both F. mosseae and D. spurcum promoted the retention of heavy metals in roots and increased the uptake of Pb, Zn, Cd, and As, and both fungi restricted heavy metal transfer, resulting in decreased Pb, Zn, and Cd contents in shoots. Therefore, the fungi reduced the translocation factors for heavy metal content (TF) and uptake (TF') in maize. Additionally, F. mosseae promoted P and S uptake by shoots, and D. spurcum increased P and S uptake by roots. Moreover, highly significant negative correlations were found between antioxidant capacity and the H 2 O 2 , MDA, and heavy metal contents, and there was a positive correlation with the biomass of maize leaves. These results suggested that AMF alleviated plant toxicity and that this effect was closely related to antioxidant activation in the maize leaves and increased retention of heavy metals in the roots.

Arbuscular mycorrhiza (AM) is a widespread symbiosis between most land plants and fungi of the Glomeromycotina, which has existed for more than 400million years. AM fungi (AMF) improve plant nutrition with mineral nutrients and conversely, their growth and development is fueled by organic carbon supplied from their host. Recent studies demonstrated independently and with different experimental approaches that lipids are transferred from plants to fungi in addition to sugars, and that AMF are dependent on this lipid supply because they lack genes encoding fatty acid synthase I subunits. Dependence on host lipids or lipid parasitism occur in a range of interorganismic associations with participants from almost all kingdoms. Thus, these phenomena seem rather common in mutualistic and parasitic interactions., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

ZnO nanoparticles (NPs) are applied in a wide variety of applications and frequently accumulate in the environment, thus posing risks to the environment and human health. Arbuscular mycorrhizal (AM) fungi (AMF) associate symbiotically with roots of most higher plants, helping their host plants acquire phosphorus (P). AMF can reduce the toxicity of ZnO NPs, but the benefits of AMF to host plants highly vary with soil available P. We hypothesize that organic P may help AMF to alleviate ZnO NP phytotoxicity. Here, we investigated the effects of inoculation with Funneliformis mosseae on plant growth and Zn accumulation, using maize grown in soil-sand mix substrates spiked with ZnO NPs (0 or 500 mg kg -1 ) under different organic P supply levels (0, 20, or 50 mg kg -1 ). The results showed addition of ZnO NPs inhibited root colonization rate, increased the shoot/root P concentration ratio, and led to significant Zn accumulation in soil and plants. As predicted, AM effects on maize plants all varied with P supply levels, both with or without ZnO NP additions. Organic P interacted synergistically with AMF to promote plant growth and acquisition of P, N, K, Fe, and Cu. AM inoculation reduced the bioavailable Zn released from ZnO NPs and decreased the concentrations and translocation of Zn to maize shoots. In conclusion, ZnO NPs caused excess Zn in soil and plants, posing potential environmental risks. However, our present results first demonstrate that organic P exhibited similar positive effects to AMF and interacted synergistically with AMF to improve plant growth and nutrition, and to decrease Zn accumulation and partitioning in plants, and thus helped diminish the adverse effects induced by ZnO NPs.

Plant-soil feedback is recognized as the mutual interaction between plants and soil microorganisms, but its role on the biological invasion of the Brazilian tropical seasonal dry forest by invasive plants still remains unclear. Here, we analyzed and compared the arbuscular mycorrhizal fungi (AMF) communities and soil characteristics from the root zone of invasive and native plants, and tested how these AMF communities affect the development of four invasive plant species (Cryptostegia madagascariensis, Parkinsonia aculeata, Prosopis juliflora, and Sesbania virgata). Our field sampling revealed that AMF diversity and frequency of the Order Diversisporales were positively correlated with the root zone of the native plants, whereas AMF dominance and frequency of the Order Glomerales were positively correlated with the root zone of invasive plants. We grew the invasive plants in soil inoculated with AMF species from the root zone of invasive (I changed ) and native (I unaltered ) plant species. We also performed a third treatment with sterilized soil inoculum (control). We examined the effects of these three AMF inoculums on plant dry biomass, root colonization, plant phosphorous concentration, and plant responsiveness to mycorrhizas. We found that I unaltered and I changed promoted the growth of all invasive plants and led to a higher plant dry biomass, mycorrhizal colonization, and P uptake than control, but I changed showed better results on these variables than I unaltered . For plant responsiveness to mycorrhizas and fungal inoculum effect on plant P concentration, we found positive feedback between changed-AMF community (I changed ) and three of the studied invasive plants: C. madagascariensis, P. aculeata, and S. virgata.

The protective role of arbuscular mycorrhizal fungi (AMF) against the phytopathogen Clavibacter michiganensis subsp. michiganensis (Cmm) was examined in tomato plants. Seven different AMF isolates were used to determine which ones were able to induce effectively resistance against Cmm. Stems of seven-week tomato plants were infected with Cmm, then a disease severity index (DSI) was determined during the next three weeks. In addition to different responses to mycorrhizal inoculation, three levels of responses to the bacterial disease were recognized in treatments. Plants inoculated with Rhizophagus irregularis (Ri) showed both the highest colonization and the highest induced resistance to Cmm while the effect of Funneliformis mosseae, Gigaspora margarita and Claroideoglomus claroideum on mycorrhizal colonization and on the induced resistance were intermediate and high, respectively. Subsequently, Ri was chosen to inoculate ethylene-insensitive tomato mutant line Never ripe (Nr) and its background (Pearson) to investigate the possible role of ethylene (ET) in the mycorrhiza-induced resistance (MIR). The results showed that Ri could induce systemic resistance against Cmm in the Pearson background, whereas ET-insensitivity in Nr plants impaired MIR. These results suggest that ET is required for Ri-induced resistance against Cmm. To our knowledge, this is the first study to examine the effect of different AMF isolates on the response of tomato plants to Cmm and involvement of ET in MIR against Cmm.

It was documented that arbuscular mycorrhiza fungi (AMF) play an important role in protecting host plants against arsenic (As) contamination. However, most terrestrial ecosystems contain a considerable number of nonmycorrhizal plants. So far little information is available for the interaction of such non-host plants with AMF under As contaminations. By using a dual compartment cultivation system with a plastic board or a nylon mesh separating roots of non-host pepperweed from roots of the AM-host alfafa plants, avoiding direct root competition, the two plant species were grown separately or partially separated (with rhizosphere effects) in the presence or absence of the AMF Rhizophagus irregularis in As-contaminated soil. The results indicated that mycorrhiza caused phosphorus (P) concentration decrease in the non-host pepperweed, but promoted the P concentration of the AM host alfafa. Mycorrhiza is potentially helpful for non-host pepperweed to adapt to As contamination by decreasing root As concentration and showing no suppressing effect on biomass production. The study provides further evidence for the protective effects of AMF on non-host plants against As contamination, and improved our understanding of the potential role of AMF for non-host plant adaptation to As contaminated soils., (Copyright © 2017. Published by Elsevier B.V.)

Liquorice (Glycyrrhiza uralensis) is an important medicinal plant for which there is a huge market demand. It has been reported that arbuscular mycorrhizal (AM) symbiosis and drought stress can stimulate the accumulation of the active ingredients, glycyrrhizin and liquiritin, in liquorice plants, but the potential interactions of AM symbiosis and drought stress remain largely unknown. In the present work, we investigated mycorrhizal effects on plant growth and accumulation of glycyrrhizin and liquiritin in liquorice plants under different water regimes. The results indicated that AM plants generally exhibited better growth and physiological status including stomatal conductance, photosynthesis rate, and water use efficiency compared with non-AM plants. AM inoculation up-regulated the expression of an aquaporin gene PIP and decreased root abscisic acid (ABA) concentrations under drought stress. In general, AM plants displayed lower root carbon (C) and nitrogen (N) concentrations, higher phosphorus (P) concentrations, and therefore, lower C:P and N:P ratios but higher C:N ratio than non-AM plants. On the other hand, AM inoculation increased root glycyrrhizin and liquiritin concentrations, and the mycorrhizal effects were more pronounced under moderate drought stress than under well-watered condition or severe drought stress for glycyrrhizin accumulation. The accumulation of glycyrrhizin and liquiritin in AM plants was consistent with the C:N ratio changes in support of the carbon-nutrient balance hypothesis. Moreover, the glycyrrhizin accumulation was positively correlated with the expression of glycyrrhizin biosynthesis genes SQS1, β-AS, CYP88D6, and CYP72A154. By contrast, no significant interaction of AM inoculation with water treatment was observed for liquiritin accumulation, while we similarly observed a positive correlation between liquiritin accumulation and the expression of a liquiritin biosynthesis gene CHS. These results suggested that AM inoculation in combination with proper water management potentially could improve glycyrrhizin and liquiritin accumulation in liquorice roots and may be practiced to promote liquorice cultivation.

The presence of dark septate endophytes (DSEs) or arbuscular mycorrhizal fungi (AMF) in plant roots and their effects on plant fitness have been extensively described. However, little is known about their interactions when they are simultaneously colonizing a plant root, especially in trace element (TE)-polluted soils. We therefore investigated the effects of Cadophora sp. and Funneliformis mosseae on ryegrass (Lolium perenne) growth and element uptake in a Cd/Zn/Pb-polluted soil. The experiment included four treatments, i.e., inoculation with Cadophora sp., inoculation with F. mosseae, co-inoculation with Cadophora sp. and F. mosseae, and no inoculation. Ryegrass biomass and shoot Na, P, K, and Mg concentrations significantly increased following AMF inoculation as compared to non-inoculated controls. Similarly, DSE inoculation increased shoot Na concentration, whereas dual inoculation significantly decreased shoot Cd concentration. Moreover, oxidative stress determined by ryegrass leaf malondialdehyde concentration was alleviated both in the AMF and dual inoculation treatments. We used quantitative PCR and microscope observations to quantify colonization rates. They demonstrated that DSEs had no effect on AMF colonization, while AMF colonization slightly decreased DSE frequency. We also monitored fluorescein diacetate (FDA) hydrolysis and alkaline phosphatase (AP) activity in the rhizosphere soils. FDA hydrolysis remained unchanged in the three inoculated treatments, but AMF colonization increased AP activity and P mobility in the soil whereas DSE colonization did not alter AP activity. In this experiment, we unveiled the interactions between two ecologically important fungal groups likely to occur in roots which involved a decrease of oxidative stress and Cd accumulation in shoots. These results open promising perspectives on the fungal-based phytomanagement of TE-contaminated sites by the production of uncontaminated and marketable plant biomass.

Phytoremediation of polluted sites can be improved by co-inoculation with mycorrhizal and endophytic fungi. In this study, the effects of single- and co-inoculation of Lactuca serriola with an arbuscular mycorrhizal (AM) fungus, Rhizoglomus intraradices, and endophytic fungi, Mucor sp. or Trichoderma asperellum, on plant growth, vitality, toxic metal accumulation, sesquiterpene lactone production and flavonoid concentration in the presence of toxic metals were evaluated. Inoculation with the AM fungus increased biomass yield of the plants grown on non-polluted and polluted substrate. Co-inoculation with the AM fungus and Mucor sp. resulted in increased biomass yield of plants cultivated on the polluted substrate, whereas co-inoculation with T. asperellum and the AM fungus increased plant biomass on the non-polluted substrate. In the presence of Mucor sp., mycorrhizal colonization and arbuscule richness were increased in the non-polluted substrate. Co-inoculation with the AM fungus and Mucor sp. increased Zn concentration in leaves and roots. The concentration of sesquiterpene lactones in plant leaves was decreased by AM fungus inoculation in both substrates. Despite enhanced host plant costs caused by maintaining symbiosis with numerous microorganisms, interaction of wild lettuce with both mycorrhizal and endophytic fungi was more beneficial than that with a single fungus. The study shows the potential of double inoculation in unfavourable environments, including agricultural areas and toxic metal-polluted areas.

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.

We investigated the effects of the arbuscular mycorrhizal fungus (AMF) Funneliformis mosseae on the growth and root architecture of plantlets of the grape rootstock 41B MGt under hydroponic conditions, and analyzed the concomitant expression of putative mycorrhizal-specific phosphate transporter 1 (PHT1) genes. In vitro propagated plantlets were acclimatized to ex vitro culture before AMF inoculation and grown under low phosphate (Pi) nutrition conditions during 6 weeks. Grape roots could be efficiently colonized by F. mosseae in this culture system, as shown by high mycorrhization frequency and intensity. The presence of many arbuscules in the cortex was coupled with high-level expression of two PHT1 genes in grape roots. These two very similar genes, named VvPht1-1 and VvPht1-2, present P1BS and MYCS regulatory motifs in their promoter, consistent with a specific role in the mycorrhizal pathway of Pi uptake. Although AMF inoculation significantly increased shoot growth, no effect on root biomass was observed. However, inoculated grapes exhibited an enhanced branched root system compared with non-inoculated controls, with a twofold higher number of tips and a higher proportion of fine roots usually involved in nutrient uptake from the soil. Taken together, these results suggest that root colonization by F. mosseae improved grape growth by favoring the uptake of Pi from the substrate via VvPht1-1 and VvPht1-2 high-level expression.

Root-hair growth and development regulated by soil microbes is associated with auxin. In this background, we hypothesized that mycorrhizal fungal inoculation induces greater root-hair growth through stimulated auxin synthesis and transport under water stress conditions. Trifoliate orange (Poncirus trifoliata) was inoculated with an arbuscular mycorrhizal (AM) fungus (Funneliformis mosseae) under well-watered (WW) and drought stress (DS) for 9 weeks. Compared with non-AM seedlings, AM seedlings displayed significantly higher density, length, and diameter of root hairs and root indoleacetic acid (IAA) level, whereas lower total root IAA efflux, regardless of soil moisture status. Root PtYUC3 and PtYUC8 involved in IAA biosynthesis were up-regulated by mycorrhization under WW and DS, whereas AM-modulated expression in PtTAA1, PtTAR2, PtYUC4, and PtYUC6 depended on status of soil moisture. Mycorrhizal inoculation down-regulated the transcript level of root auxin efflux carriers like PtPIN1 and PtPIN3, whereas significantly up-regulated the expression of root auxin-species influx carriers like PtABCB19 and PtLAX2 under DS. These results indicated that AMF-stimulated greater root-hair growth of trifoliate orange under DS that is independent on AMF species is related with mycorrhiza-modulated auxin synthesis and transport, which benefits the host plant to enhance drought tolerance.

Arbuscular mycorrhizal fungi (AMF; phylum Gomeromycota) associate with plants forming one of the most successful microbe-plant associations. The fungi promote plant diversity and have a potentially important role in global agriculture. Plant growth depends on both inter- and intra-specific variation in AMF. It was recently reported that an unusually large number of AMF taxa have an intercontinental distribution, suggesting long-distance gene flow for many AMF species, facilitated by either long-distance natural dispersal mechanisms or human-assisted dispersal. However, the intercontinental distribution of AMF species has been questioned because the use of very low-resolution markers may be unsuitable to detect genetic differences among geographically separated AMF, as seen with some other fungi. This has been untestable because of the lack of population genomic data, with high resolution, for any AMF taxa. Here we use phylogenetics and population genomics to test for intra-specific variation in Rhizophagus irregularis, an AMF species for which genome sequence information already exists. We used ddRAD sequencing to obtain thousands of markers distributed across the genomes of 81 R. irregularis isolates and related species. Based on 6 888 variable positions, we observed significant genetic divergence into four main genetic groups within R. irregularis, highlighting that previous studies have not captured underlying genetic variation. Despite considerable genetic divergence, surprisingly, the variation could not be explained by geographical origin, thus also supporting the hypothesis for at least one AMF species of widely dispersed AMF genotypes at an intercontinental scale. Such information is crucial for understanding AMF ecology, and how these fungi can be used in an environmentally safe way in distant locations.

Background: Lignin and cellulose, organic constituents of the plant or plant-based material not commonly used for feeding purpose are referred as Biomass. Patents suggest that this can be used as the best resource of renewable energy. Vesicular Arbuscular Mycorrhizae (VAM) fungi can play an effective role in biomass manufacturing through activated metabolism of the plant under dual symbiosis. During C acclimatization, mycorrhizal inoculated plants existent greater number of leaves with a height of plants as compared to non-mycorrhizal plants. The current article discloses the search of the natural resources for C assimilation into biomass using mycorrhizal symbiosis., Methods: The pot experiment was conducted in the natural environment for extraction of more bioenergy through biomass of Conocarpus erectus L under VAM (Glomus fasciculatum) inoculation in various environmental conditions with replicates., Results: It observed that these fungal engineered plants showed distinctive prospective to offer, enhanced biomass to energy couple with a strong network for sinking CO2 from the atmosphere via strong roots and large surface area of leaves. There was an increase in biomass (9-17% respectively) of the plant under drought-VAM, VAM inoculation and VAM- enriched CO2 conditions in same period in comparison to control plants through lignin, cellulose and carbohydrate contents. It was followed by enhanced enzyme activities and nutrient ions in dual symbiosis., Conclusion: Coupling biomass-originated energy may recover environmental conditions and commercial value for sustainable growth in energy consumption sector. The green energy from fungal engineered plants may replace high demand of fossil fuel as a young biofuel and make the cities more productive in the fabrication of bioenergy too in the form of biomass or biofuel with C impartial atmosphere., (Copyright© Bentham Science Publishers; For any queries, please email at epub@benthamscience.org.)

Perception of arbuscular mycorrhizal fungi (AMF) triggers distinct plant signalling responses for parallel establishment of symbiosis and induction of lateral root formation. Rice receptor kinase CHITIN ELICITOR RECEPTOR KINASE 1 (CERK1) and α/β-fold hydrolase DWARF14-LIKE (D14L) are involved in pre-symbiotic fungal perception. After 6 wk post-inoculation with Rhizophagus irregularis, root developmental responses, fungal colonization and transcriptional responses were monitored in two independent cerk1 null mutants; a deletion mutant lacking D14L, and with D14L complemented as well as their respective wild-type cultivars (cv Nipponbare and Nihonmasari). Here we show that although essential for symbiosis, D14L is dispensable for AMF-induced root architectural modulation, which conversely relies on CERK1. Our results demonstrate uncoupling of symbiosis and the symbiotic root developmental signalling during pre-symbiosis with CERK1 required for AMF-induced root architectural changes., (© 2017 The Authors. New Phytologist © 2017 New Phytologist Trust.)

In arbuscular mycorrhizal (AM) roots, the plasma membrane (PM) of the host plant is involved in all developmental stages of the symbiotic interaction, from initial recognition to intracellular accommodation of intra-radical hyphae and arbuscules. Although the role of the PM as the agent for cellular morphogenesis and nutrient exchange is especially accentuated in endosymbiosis, very little is known regarding the PM protein composition of mycorrhizal roots. To obtain a global overview at the proteome level of the host PM proteins as modified by symbiosis, we performed a comparative protein profiling of PM fractions from Medicago truncatula roots either inoculated or not with the AM fungus Rhizophagus irregularis. PM proteins were isolated from root microsomes using an optimized discontinuous sucrose gradient; their subsequent analysis by liquid chromatography followed by mass spectrometry (MS) identified 674 proteins. Cross-species sequence homology searches combined with MS-based quantification clearly confirmed enrichment in PM-associated proteins and depletion of major microsomal contaminants. Changes in protein amounts between the PM proteomes of mycorrhizal and non-mycorrhizal roots were monitored further by spectral counting. This workflow identified a set of 82 mycorrhiza-responsive proteins that provided insights into the plant PM response to mycorrhizal symbiosis. Among them, the association of one third of the mycorrhiza-responsive proteins with detergent-resistant membranes pointed at partitioning to PM microdomains. The PM-associated proteins responsive to mycorrhization also supported host plant control of sugar uptake to limit fungal colonization, and lipid turnover events in the PM fraction of symbiotic roots. Because of the depletion upon symbiosis of proteins mediating the replacement of phospholipids by phosphorus-free lipids in the plasmalemma, we propose a role of phosphate nutrition in the PM composition of mycorrhizal roots.

Arbuscular mycorrhizal fungi (AMF) colonize up to 90% of all land plants and facilitate the acquisition of mineral nutrients by their hosts. Inorganic orthophosphate (P i ) and nitrogen (N) are the major nutrients transferred from the fungi to plants. While plant P i transporters involved in nutrient transfer at the plant-fungal interface have been well studied, the plant N transporters participating in this process are largely unknown except for some ammonium transporters (AMT) specifically assigned to arbuscule-colonized cortical cells. In plants, many nitrate transporter 1/peptide transporter family (NPF) members are involved in the translocation of nitrogenous compounds including nitrate, amino acids, peptides and plant hormones. Whether NPF members respond to AMF colonization, however, is not yet known. Here, we investigated the transcriptional regulation of 82 rice (Oryza sativa) NPF genes in response to colonization by the AMF Rhizophagus irregularis in roots of plants grown under five different nutrition regimes. Expression of the four OsNPF genes NPF2.2/PTR2, NPF1.3, NPF6.4 and NPF4.12 was strongly induced in mycorrhizal roots and depended on the composition of the fertilizer solution, nominating them as interesting candidates for nutrient signaling and exchange processes at the plant-fungal interface.

Aluminum (Al) toxicity can induce oxidative and nitrosative stress, which limits growth and yield of crop plants. Nevertheless, plant tolerance to stress may be improved by symbiotic associations including arbuscular mycorrhiza (AM). Nitric oxide (NO) is a signaling molecule involved in physiological processes and plant responses to abiotic and biotic stresses. However, almost no information about the NO metabolism has been gathered about AM. In the present work, Medicago truncatula seedlings were inoculated with Rhizophagus irregularis, and 7-week-old plants were treated with 50μM AlCl 3 for 3h. Cytochemical and molecular techniques were used to measure the components of the NO metabolism, including NO content and localization, expression of genes encoding NO-synthesis (MtNR1, MtNR2 and MtNIR1) and NO-scavenging (MtGSNOR1, MtGSNOR2, MtHB1 and MtHB2) enzymes and the profile of protein tyrosine nitration (NO 2 -Tyr) in Medicago roots. For the first time, NO and NO 2 -Tyr accumulation was connected with fungal structures (arbuscules, vesicles and intercellular hyphae). Expression analysis of genes encoding NO-synthesis enzymes indicated that AM symbiosis results in lower production of NO in Al-treated roots in comparison to non-mycorrhizal roots. Elevated levels of transcription of genes encoding NO-scavenging enzymes indicated more active NO scavenging in AMF-inoculated Al-treated roots compared to non-inoculated roots. These results were confirmed by less NO accumulation and lower protein nitration in Al-stressed mycorrhizal roots in comparison to non-mycorrhizal roots. This study provides a new insight in NO metabolism in response to arbuscular mycorrhiza under normal and metal stress conditions. Our results suggest that mycorrhizal fungi decrease NO and tyrosine nitrated proteins content in Al-treated Medicago roots, probably via active NO scavenging system., (Copyright © 2017 Elsevier GmbH. All rights reserved.)

Hexavalent chromium is a potent carcinogen, while phosphorus is an essential nutrient. The role of arbuscular mycorrhizal fungi (AMF) in the uptake of P is well known and was also reported, at low levels, for Cr. However, it is unclear whether the uptake of Cr can impact the short-term uptake dynamics of P since both elements have a similar chemical structure and may thus potentially compete with each other during the uptake process. This study investigated the impact of Cr(VI) on short-term P uptake by the AMF Rhizophagus irregularis MUCL 41833 in Medicago truncatula. Bi-compartmented Petri plates were used to spatially separate a root compartment (RC) from a hyphal compartment (HC) using a whole plant in vitro culture system. The HC was supplemented with Cr(VI). Chromium(VI) as well as total Cr and P were monitored during 16 h within the HC and their concentrations determined by the end of the experiment within roots and shoots. Our results indicated that the uptake and translocation of Cr from hyphae to roots was a fast process: roots in which the extraradical mycelium (ERM) was exposed to Cr(VI) accumulated more Cr than roots of which the ERM was not exposed to Cr(VI) or was dead. Our results further confirmed that dead ERM immobilized more Cr than alive ERM. Finally our results demonstrated that the short exposure to Cr(VI) was sufficient to stimulate P uptake by the ERM and that the stimulation process began within the first 4 h of exposure., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Growth and maintenance partly depend on both respiration and ATP production during oxidative phosphorylation in leaves. Under stress, ATP is needed to maintain the accumulated biomass. ATP production mostly proceeds from the cytochrome oxidase pathway (COP), while respiration via the alternative oxidase pathway (AOP) may decrease the production of ATP per oxygen consumed, especially under phosphorus (P) limitation and salinity conditions. Symbiosis with arbuscular mycorrhizal (AM) fungi is reputed by their positive effect on plant growth under stress at mature stages of colonization; however, fungal colonization may decrease plant growth at early stages. Thus, the present research is based on the hypothesis that AM fungus colonization will increase both foliar respiration and ATP production at mature stages of plant growth while decreasing them both at early stages. We used the oxygen-isotope-fractionation technique to study the in vivo respiratory activities and ATP production of the COP and AOP in AM and non-AM (NM) tobacco plants grown under P-limiting and saline conditions in sand at different growth stages (14, 28 and 49days). Our results suggest that AM symbiosis represents an ATP cost detrimental for shoot growth at early stages, whilst it represents a benefit on ATP allowing for faster rates of growth at mature stages, even under salinity conditions., (Copyright © 2017 Elsevier GmbH. All rights reserved.)

Nitrogen cycling in agroecosystems is heavily dependent upon arbuscular mycorrhizal fungi (AMF) present in the soil microbiome. These fungi develop obligate symbioses with various host plant species, thus increasing their ability to acquire nutrients. However, AMF are particularly sensitive to physical, chemical and biological disturbances caused by human actions that limit their establishment. For a more sustainable agriculture, it will be necessary to further investigate which agricultural practices could be favorable to maximize the benefits of AMF to improve crop nitrogen use efficiency (NUE), thus reducing nitrogen (N) fertilizer usage. Direct seeding, mulch-based cropping systems prevent soil mycelium disruption and increase AMF propagule abundance. Such cropping systems lead to more efficient root colonization by AMF and thus a better establishment of the plant/fungal symbiosis. In addition, the use of continuous cover cropping systems can also enhance the formation of more efficient interconnected hyphal networks between mycorrhizae colonized plants. Taking into account both fundamental and agronomic aspects of mineral nutrition by plant/AMF symbioses, we have critically described, how improving fungal colonization through the reduction of soil perturbation and maintenance of an ecological balance could be helpful for increasing crop NUE., (Copyright © 2017 Elsevier B.V. All rights reserved.)

The effects of arbuscular mycorrhizal (AM) fungi on plant-associated microbes are poorly known. We tested the hypothesis that colonization by an AM fungus affects microbial species richness and microbial community composition of host plant tissues. We grew the grass, Deschampsia flexuosa in a greenhouse with or without the native AM fungus, Claroideoglomus etunicatum. We divided clonally produced tillers into two parts: one inoculated with AM fungus spores and one without AM fungus inoculation (non-mycorrhizal, NM). We characterized bacterial (16S rRNA gene) and fungal communities (internal transcribed spacer region) in surface-sterilized leaf and root plant compartments. AM fungus inoculation did not affect microbial species richness or diversity indices in leaves or roots, but the AM fungus inoculation significantly affected bacterial community composition in leaves. A total of three OTUs in leaves belonging to the phylum Firmicutes positively responded to the presence of the AM fungus in roots. Another six OTUs belonging to the Proteobacteria (Alpha, Beta, and Gamma) and Bacteroidetes were significantly more abundant in NM plants when compared to AM fungus-inoculated plants. Further, there was a significant correlation between plant dry weight and leaf microbial community compositional shift. Also, there was a significant correlation between leaf bacterial community compositional shift and foliar nitrogen content changes due to AM fungus inoculation. The results suggest that AM fungus colonization in roots has a profound effect on plant physiology that is reflected in leaf bacterial community composition.

Arbuscular mycorrhizal (AM) fungi experience oxidative stress during the plant-fungal interaction, due to endogenous reactive oxygen species (ROS) produced by fungal metabolism and exogenous ROS produced by plant cells. Here, we examine the responses to H 2 O 2 in Gigaspora margarita, an AM fungus containing the endobacterial symbiont Candidatus Glomeribacter gigasporarum (CaGg). Previous studies revealed that G. margarita with its endobacterium produces more ATP and has higher respiratory activity than a cured line that lacks the endobacterium. This higher bioenergetic potential leads to higher production of ROS and to a higher ROS-detoxifying capacity, suggesting a direct or indirect role of the endobacterium in modulating fungal antioxidant responses. To test the hypothesis that the fungal-endobacterial symbiosis may enhance the fitness of the AM fungus in the presence of oxidative stress, we treated the fungus with a sublethal concentration of H 2 O 2 and performed RNA-seq analysis. Our results demonstrate that (i) irrespective of the endobacterium presence, G. margarita faces oxidative stress by activating multiple metabolic processes (methionine oxidation, sulfur uptake, the pentose phosphate pathway, activation of ROS-scavenger genes); (ii) in the presence of its endobacterium, G. margarita upregulates some metabolic pathways, like chromatin status modifications and iron metabolism; and (iii) contrary to our hypothesis, the cured line responds to H 2 O 2 by activating the transcription of specific ROS scavengers. We confirmed the RNA-seq findings by measuring the glutathione and ascorbate concentration, which was the same in both lines after H 2 O 2 treatment. We conclude that both fungal lines may face oxidative stress, but they activate alternative strategies.

Bacterial wilt caused by Ralstonia solanacearum is one of the world's most important soil-borne plant diseases. In Martinique, French West Indies, a highly virulent new pathogenic variant of this bacterium (phylotype IIB/4NPB) severely impacts tomato production. Here we report on the effect of R. solanacearum CFBP 6783, classified in phytotype IIB/4NPB, on tomato plantlets grown under strict in vitro culture conditions in the presence or absence of the arbuscular mycorrhizal fungus Rhizophagus irregularis MUCL 41833. A mycelium donor plant (i.e. Crotalaria spectabilis) was used for rapid, uniform mycorrhization of tomato plantlets that were subsequently infected by the bacterium. Bacterial wilt was significantly delayed and the incidence of the disease consequently reduced in the mycorrhizal tomato plantlets. Conversely, R. solanacearum did not affect root colonization by the AMF within the 16 days of the experiment. These results suggested that the mycorrhizal fungus was able to reduce bacterial wilt symptoms, probably by eliciting defence mechanisms in the plant.

Although sandy coastal plains are important buffer zones to protect the coast line and maintain biological diversity and ecosystem services, these ecosystems have been endangered by anthropogenic activities. Thus, information on coastal biodiversity and forces shaping coastal biological diversity are extremely important for effective conservation strategies. In this study, we aimed to compare arbuscular mycorrhizal (AM) fungal communities from soil samples collected on the mainland and nearby islands located in Brazilian sandy coastal plain ecosystems (Restingas) to get information about AM fungal biogeography and identify factors shaping these communities. Soil samples were collected in 2013 and 2014 on the beachfront of the tropical sandy coastal plain at six sites (three island and three mainland locations) across the northeast, southeast, and south regions of Brazil. Overall, we recorded 53 AM fungal species from field and trap culture samples. The richness and diversity of AM fungal species did not differ between mainland and island locations, but AM fungal community assemblages were different between mainland and island environments and among most sites sampled. Glomeromycota communities registered from island samples showed higher heterogeneity than communities from mainland samples. Sandy coastal plains harbor diverse AM fungal communities structured by climatic, edaphic, and spatial factors, while the distance from the colonizing source (mainland environments) does not strongly affect the AM fungal communities in Brazilian coastal environments.

Arbuscular mycorrhizal symbiosis is an ancient interaction between plants and fungi of the phylum Glomeromycota. In exchange for photosynthetically fixed carbon, the fungus provides the plant host with greater access to soil nutrients via an extensive network of root-external hyphae. Here, to determine the impact of the symbiosis on the host ionome, the concentration of 19 elements was determined in the roots and leaves of a panel of 30 maize varieties, grown under phosphorus-limiting conditions, with or without inoculation with the fungus Funneliformis mosseae. Although the most recognized benefit of the symbiosis to the host plant is greater access to soil phosphorus, the concentration of a number of other elements responded significantly to inoculation across the panel as a whole. In addition, variety-specific effects indicated the importance of plant genotype to the response. Clusters of elements were identified that varied in a co-ordinated manner across genotypes, and that were maintained between non-inoculated and inoculated plants., (© The Author 2017. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Salinity stress leads to the production of reactive oxygen species (ROS) which can cause oxidative damage in plants. A correlation between antioxidant capacity and salt tolerance has been demonstrated in several plant species, which may be enhanced by inoculation with arbuscular mycorrhizal fungi (AMF). However, plant responses to mycorrhization may differ depending on the host plant as well as AMF isolate. It has been proposed that AMF sourced from stressed environments may be better suited as stress ameliorators than non-native/exotic ones. The present study compared the effectiveness of a native inoculum from saline soil and two exotic single isolates, Funneliformis mossseae and Rhizophagus irregularis (single or dual mix), and associated their effectiveness with modulation of antioxidant defence, in two Cajanus cajan (pigeonpea) genotypes (salt sensitive-Paras, salt tolerant-Pusa 2002) under NaCl stress. Plants subjected to NaCl (0-100 mM) recorded a substantial build-up of ROS, more in Paras than Pusa 2002. Although mycorrhization with all AMF improved plant biomass and reduced oxidative burst by strengthening antioxidant enzymatic activities, inoculation with R. irregularis (alone or in combination with F. mosseae) resulted in higher biomass accumulation which correlated with its higher root colonization and improved redox stability through rapid recycling of reduced ascorbate and glutathione. The study thus suggested that mitigation of salt-induced oxidative burden by increased activation of scavenging antioxidants is an important mechanism that determined the higher effectiveness of R. irregularis over the native saline mix in pigeonpea plants.

The growth of plant roots and arbuscular mycorrhizal fungi (AMF) can be inhibited by low pH; however, it is largely unknown which is more sensitive to low pH. This study aimed to compare the physiological and molecular responses of external hyphae (EH) and roots to low pH in terms of growth, development and functioning. We established AM symbiosis in a two-compartmented system (root compartment, RC; hyphal compartment, HC) using AMF and transformed hairy roots and exposed them to pH 6.5 and/or pH 4.5. The results showed that pH 4.5 significantly decreased root cell viability, while EH at pH 6.5 attenuated the effect. In either RC or HC, pH 4.5 reduced biomass, P content, colonization, ALP activity in roots, and ALP activity and polyphosphate accumulation in EH. GintPT expression in EH was inhibited by pH 4.5 in HC but not in RC. The expression of mycorrhiza-responsive LePTs was significantly reduced by the lower colonization due to decreased pH in either RC or HC, while the expression of non-mycorrhiza-responsive LePTs was not affected. Variation partitioning analysis indicated that EH was less sensitive to low pH than roots. The interactions between roots and EH under low pH stress merit further investigation., (© 2017 Society for Applied Microbiology and John Wiley & Sons Ltd.)

A greenhouse pot experiment was conducted to investigate the effects of arbuscular mycorrhizal (AM) fungi Claroideoglomus etunicatum (CE) and Rhizophagus intraradices (RI) on AM colonization rate, biomass, nutrient uptake, C:N:P stoichiometry, and the uptake and transport of lanthanum (La) and lead (Pb) by maize ( Zea mays L.) grown in La-and Pb-contaminated soils (combined La-Pb concentrations of 50, 200, and 800 mg·kg -1 ). The aim was to provide a scientific basis for the remediation of soils contaminated by rare earth elements and heavy metals. The results indicated that symbiotic associations were successfully established between the two isolates and maize, and the average AM colonization rate ranged from 26.7% to 95.8%. The increasing concentrations of La and Pb in soils significantly decreased the mycorrhizal colonization rate, biomass, and mineral nutrition concentrations of the maize, and significantly increased C:P and N:P ratios and the concentrations of La and Pb in shoots and roots of maize. The shoot and root dry weights of maize were significantly increased by 17.8%-158.9% with two AM fungi inoculations, while the P concentration of shoots and roots of the maize were significantly increased by 24.5%-153.8%. Inoculation with two AM fungi decreased the C:P and N:P ratios, consistent with the growth rate hypothesis. With AM fungi inoculation in three types of La-Pb co-contaminated soils, root Pb concentrations of the maize significantly increased by 51.3%-67.7%; shoot Pb concentrations of the maize significantly decreased by 16.0%-67.7%; and the transport rate of Pb from root to shoot of the maize decreased by 31.5%-54.7%. Meanwhile, inoculation with AM fungi significantly increased the shoot La concentrations in the maize grown in soils mildly contaminated with La-Pb, while it significantly decreased shoot La concentrations, increased root La concentrations of maize, and inhibited the transport of La from root to shoot of the maize grown in soils moderately contaminated with La-Pb, but had no significant effect in severely contaminated soils. The results showed that AM fungi had the potential to promote phytoremediation of soils contaminated with rare earth elements and heavy metals, with potential applications to revegetate such contaminated soil ecosystems.

Arbuscular mycorrhizas (AM) are the most common symbiotic associations between a plant's root compartment and fungi. They provide nutritional benefit (mostly inorganic phosphate [P i ]), leading to improved growth, and nonnutritional benefits, including defense responses to environmental cues throughout the host plant, which, in return, delivers carbohydrates to the symbiont. However, how transcriptional and metabolic changes occurring in leaves of AM plants differ from those induced by P i fertilization is poorly understood. We investigated systemic changes in the leaves of mycorrhized Medicago truncatula in conditions with no improved P i status and compared them with those induced by high-P i treatment in nonmycorrhized plants. Microarray-based genome-wide profiling indicated up-regulation by mycorrhization of genes involved in flavonoid, terpenoid, jasmonic acid (JA), and abscisic acid (ABA) biosynthesis as well as enhanced expression of MYC2 , the master regulator of JA-dependent responses. Accordingly, total anthocyanins and flavonoids increased, and most flavonoid species were enriched in AM leaves. Both the AM and P i treatments corepressed iron homeostasis genes, resulting in lower levels of available iron in leaves. In addition, higher levels of cytokinins were found in leaves of AM- and P i -treated plants, whereas the level of ABA was increased specifically in AM leaves. Foliar treatment of nonmycorrhized plants with either ABA or JA induced the up-regulation of MYC2 , but only JA also induced the up-regulation of flavonoid and terpenoid biosynthetic genes. Based on these results, we propose that mycorrhization and P i fertilization share cytokinin-mediated improved shoot growth, whereas enhanced ABA biosynthesis and JA-regulated flavonoid and terpenoid biosynthesis in leaves are specific to mycorrhization., (© 2017 American Society of Plant Biologists. All Rights Reserved.)

Little is known about the adaptive capacity of arbuscular mycorrhizal (AM) fungi to novel hosts. Here we assessed the possibility of two heterospecific AM fungal isolates to adaptively change, in terms of host biomass response, as a function of host plant identity, over the course of a growing season. First, we produced pure inocula of Rhizophagus clarus and Rhizophagus intraradices, each starting from a single spore. Second, we "trained" each isolate individually in a community with two plants, sudangrass (Sorgum bicolour subsp. drummondii) and leek (Aliium ampeloprasum var. porrum), using a dual-compartment system to allow the establishment of a common mycorrhizal network between the two hosts. Third, we conducted a greenhouse experiment to reciprocally test each "trained" clone, obtained from each compartment, either with the same (home), or the other host (away) under two contrasting phosphorus levels. Overall, results did not support adaptive responses of the AM fungi to their hosts (i.e., greater host biomass under "home" relative to "away" conditions), but the opposite (i.e., greater host biomass under "away" relative to "home" conditions) was more frequently observed. These changes in AM fungal symbiotic functioning open the possibility for relatively rapid genetic change of arbuscular mycorrhizal fungi in response to new hosts, which represents one step forward from in vitro experiments.

Most plants entertain mutualistic interactions known as arbuscular mycorrhiza (AM) with soil fungi (Glomeromycota) which provide them with mineral nutrients in exchange for reduced carbon from the plant. Mycorrhizal roots represent strong carbon sinks in which hexoses are transferred from the plant host to the fungus. However, most of the carbon in AM fungi is stored in the form of lipids. The absence of the type I fatty acid synthase (FAS-I) complex from the AM fungal model species Rhizophagus irregularis suggests that lipids may also have a role in nutrition of the fungal partner. This hypothesis is supported by the concerted induction of host genes involved in lipid metabolism. We explore the possible roles of lipids in the light of recent literature on AM symbiosis., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

Arbuscular mycorrhizal fungi (AMF) are widespread soil fungi that can form endosymbiotic structures with the root systems of most plants and can improve the tolerance of host plants to heavy metals. In the present study, we investigated the effects of AMF (Glomus coronatum) inoculation on the tolerance of Tagetes patula L. to Cu. Almost all of the non-mycorrhizal plants exposed to 100 μM Cu died after 3 d, whereas phytotoxicity was only observed in mycorrhizal plants that were exposed to Cu concentrations greater than 100 μM. Analysing the dynamic accumulation of Cu indicated that, after 7 d of Cu exposure, less Cu was absorbed or accumulated by mycorrhizal plants than by control plants, and significantly less Cu was translocated to the shoots. Meanwhile, analysing the root morphology, the integrity of the root plasma membranes, the photosynthesis rate, and the content of essential elements of plants growing in cultures with 50 μM Cu revealed that AMF inoculation markedly alleviated the toxic effects of Cu stress on root system activity, photosynthesis rate, and mineral nutrient accumulation. In addition, to understand the Cu allocation, an energy spectrum analysis of Cu content at the transverse section of root tips was conducted and subsequently provided direct evidence that intraradical hyphae at the root endodermis could selectively immobilise large amounts of Cu. Indeed, the sorption and barrier mechanisms of AMF hyphae reduce Cu toxicity in the roots of T. patula and eventually enhance the plants' Cu tolerance.

Arbuscular mycorrhizal fungi are able to improve plant establishment in polluted soils but little is known about the genes involved in the plant protection against pollutant toxicity by mycorrhization, in particular in the presence of polycyclic aromatic hydrocarbons (PAH). The present work aims at studying in both symbiotic partners, Medicago truncatula and Rhizophagus irregularis: (i) expression of genes putatively involved in PAH tolerance (MtSOD, MtPOX, MtAPX, MtGST, MtTFIIS, and MtTdp1α), (ii) activities of antioxidant (SOD, POX) and detoxification (GST) enzymes, and (iii) H 2 O 2 and the heavy PAH, benzo[a]pyrene (B[a]P) accumulation. In the presence of B[a]P, whereas induction of the enzymatic activities was detected in R. irregularis and non-mycorrhizal roots as well as upregulation of the gene expressions in the non-mycorrhizal roots, downregulation of the gene expressions and decrease of enzyme activities were observed in mycorrhizal roots. Moreover, B[a]P increased H 2 O 2 production in non-mycorrhizal roots and in R. irregularis but not in mycorrhizal roots. In addition, a lower B[a]P bioaccumulation in mycorrhizal roots was measured in comparison with non-mycorrhizal roots. Being less affected by pollutant toxicity, mycorrhizal roots did not activate any defense mechanism either at the gene expression regulation level or at the enzymatic level.

The effect of arbuscular mycorrhiza (AM) symbiosis on plant growth is associated with the balance between costs and benefits. A feedback regulation loop has been described in which the higher carbohydrate cost to plants for AM symbiosis is compensated by increases in their photosynthetic rates. Nevertheless, plant carbon balance depends both on photosynthetic carbon uptake and respiratory carbon consumption. The hypothesis behind this research was that the role of respiration in plant growth under AM symbiosis may be as important as that of photosynthesis. This hypothesis was tested in Arundo donax L. plantlets inoculated with Rhizophagus irregularis and Funneliformis mosseae. We tested the effects of AM inoculation on both photosynthetic capacity and in vivo leaf and root respiration. Additionally, analyses of the primary metabolism and ion content were performed in both leaves and roots. AM inoculation increased photosynthesis through increased CO 2 diffusion and electron transport in the chloroplast. Moreover, respiration decreased only in AM roots via the cytochrome oxidase pathway (COP) as measured by the oxygen isotope technique. This decline in the COP can be related to the reduced respiratory metabolism and substrates (sugars and tricarboxylic acid cycle intermediates) observed in roots., (© 2017 John Wiley & Sons Ltd.)

Arbuscular mycorrhizal (AM) fungi facilitate plant uptake of mineral nutrients and draw organic nutrients from the plant. Organic nutrients are thought to be supplied primarily in the form of sugars. Here we show that the AM fungus Rhizophagus irregularis is a fatty acid auxotroph and that fatty acids synthesized in the host plants are transferred to the fungus to sustain mycorrhizal colonization. The transfer is dependent on RAM2 (REQUIRED FOR ARBUSCULAR MYCORRHIZATION 2) and the ATP binding cassette transporter-mediated plant lipid export pathway. We further show that plant fatty acids can be transferred to the pathogenic fungus Golovinomyces cichoracerum and are required for colonization by pathogens. We suggest that the mutualistic mycorrhizal and pathogenic fungi similarly recruit the fatty acid biosynthesis program to facilitate host invasion., (Copyright © 2017, American Association for the Advancement of Science.)

The effects of inoculation with an arbuscular mycorrhizal (AM) fungus on Cd and Ni tolerance and uptake in Medicago sativa, an AM host, and Sesuvium portulacastrum, a non-host plant, were investigated in a greenhouse experiment. The plants were cultivated in sterilized sand in a two-compartmented system, which prevented root competition but enabled colonization of the whole substrate by AM fungal extraradical mycelium. M. sativa was either left non-inoculated or inoculated with the AM fungus Rhizophagus irregularis, and both plants were either cultivated without heavy metal (HM) addition or supplied with cadmium (Cd) or nickel (Ni), each in two doses. Additional pots with singly cultivated plants were established to control for the effect of the co-cultivation. AM significantly enhanced the growth of M. sativa and substantially increased its uptake of both HMs. The roots of S. portulacastrum became colonized by AM fungal hyphae and vesicles. The presence of the AM fungus in the cultivation system tended to increase the HM uptake of S. portulacastrum, but the effect was less consistent and pronounced than that in M. sativa. We conclude that AM fungal mycelium radiating from M. sativa did not negatively affect the growth and HM uptake of S. portulacastrum. On the contrary, we hypothesize that it stimulated the absorption and translocation of Cd and Ni in the non-host species. Thus, our results suggest that AM fungal mycelium radiating from mycorrhizal plants does not decrease the HM uptake of non-host plants, many of which are considered promising candidate plants for phytoremediation., (Copyright © 2016 Elsevier Ltd. All rights reserved.)

Plants usually interact with other plants, and the outcome of such interaction ranges from facilitation to competition depending on the identity of the plants, including their sexual expression. Arbuscular mycorrhizal (AM) fungi have been shown to modify competitive interactions in plants. However, few studies have evaluated how AM fungi influence plant intraspecific and interspecific interactions in dioecious species. The competitive abilities of female and male plants of Antennaria dioica were examined in a greenhouse experiment. Females and males were grown in the following competitive settings: (i) without competition, (ii) with intrasexual competition, (iii) with intersexual competition, and (iv) with interspecific competition by Hieracium pilosella - a plant with similar characteristics to A. dioica. Half of the pots were grown with Claroideoglomus claroideum, an AM fungus isolated from the same habitat as the plant material. We evaluated plant survival, growth, flowering phenology, and production of AM fungal structures. Plant survival was unaffected by competition or AM fungi. Competition and the presence of AM fungi reduced plant biomass. However, the sexes responded differently to the interaction between fungal and competition treatments. Both intra- and interspecific competition results were sex-specific, and in general, female performance was reduced by AM colonization. Plant competition or sex did not affect the intraradical structures, extraradical hyphae, or spore production of the AM fungus. These findings suggest that plant sexual differences affect fundamental processes such as competitive ability and symbiotic relationships with AM fungi., (© 2016 German Botanical Society and The Royal Botanical Society of the Netherlands.)

Arbuscular mycorrhizal fungi (AMF) can enhance drought tolerance in plants, whereas little is known regarding AMF contribution to sucrose and proline metabolisms under drought stress (DS). In this study, Funneliformis mosseae and Paraglomus occultum were inoculated into trifoliate orange (Poncirus trifoliata) under well watered and DS. Although the 71-days DS notably (P < 0.05) inhibited mycorrhizal colonization, AMF seedlings showed significantly (P < 0.05) higher plant growth performance and leaf relative water content, regardless of soil water status. AMF inoculation significantly (P < 0.05) increased leaf sucrose, glucose and fructose concentration under DS, accompanied with a significant increase of leaf sucrose phosphate synthase, neutral invertase, and net activity of sucrose-metabolized enzymes and a decrease in leaf acid invertase and sucrose synthase activity. AMF inoculation produced no change in leaf ornithine-δ-aminotransferase activity, but significantly (P < 0.05) increased leaf proline dehydrogenase activity and significantly (P < 0.05) decreased leaf both Δ 1 -pyrroline-5-carboxylate reductase and Δ 1 -pyrroline-5-carboxylate synthetase activity, resulting in lower proline accumulation in AMF plants under DS. Our results therefore suggest that AMF strongly altered leaf sucrose and proline metabolism through regulating sucrose- and proline-metabolized enzyme activities, which is important for osmotic adjustment of the host plant., Competing Interests: The authors declare no competing financial interests.

While a number of recent studies have revealed that arbuscular mycorrhizal fungi (AMF) can mediate invasive plant success, the influence of these symbionts on the most successful and high-impact invaders is largely unexplored. Two perennial herbs of this category of invasive plants, Rudbeckia laciniata and Solidago gigantea (Asteraceae), were thus tested in a pot experiment to determine whether AMF influence their growth, the concentration of phosphorus in biomass, and photosynthesis. The following treatments, including three common AMF species, were prepared on soils representative of two habitats that are frequently invaded by both plants, namely fallow and river valley: (1) control-soil without AMF, (2) Rhizophagus irregularis, (3) Funneliformis mosseae, and (4) Claroideoglomus claroideum. The invaders were strongly dependent on AMF for their growth. The mycorrhizal dependency of R. laciniata was 88 and 63 % and of S. gigantea 90 and 82 % for valley and fallow soils, respectively. The fungi also increased P concentration in their biomass. However, we found different effects of the fungal species in the stimulation of plant growth and P acquisition, with R. irregularis and C. claroideum being the most and least effective symbionts, respectively. None of AMF species had an impact on the photosynthetic performance indexes of both plants. Our findings indicate that AMF have a direct effect on the early stages of R. laciniata and S. gigantea growth. The magnitude of the response of both plant species to AMF was dependent on the fungal and soil identities. Therefore, the presence of particular AMF species in a site may determine the success of their invasion., Competing Interests: The authors declare that they have no conflict of interest.

Soil pollution is an important ecological problem worldwide. Phytoremediation is an environmental-friendly option for reducing metal pollution. A greenhouse experiment was conducted to determine the growth and physiological response, metal uptake, and the phytostabilization potential of a nontoxic Jatropha curcas L. genotype when grown in multimetal-polluted conditions. Plants were established on a mine residue (MR) amended or not amended with corn biochar (B) and inoculated or not inoculated with the mycorrhizal fungus Acaulospora sp. (arbuscular mycorrhizal fungus, AMF). J. curcas was highly capable of growing in an MR and showed no phytotoxic symptoms. After J. curcas growth (105 days), B produced high desorption of Cd and Pb from the MR; however, no increases in metal shoot concentrations were observed. Therefore, Jatropha may be useful for phytostabilization of metals in mine tailings. The use of B is recommended because improved MR chemical properties conduced to plant growth (cation-exchange capacity, organic matter content, essential nutrients, electrical conductivity, water-holding capacity) and plant growth development (higher biomass, nutritional and physiological performance). Inoculation with an AMF did not improve any plant growth or physiological plant characteristic. Only higher Zn shoot concentration was observed, but it was not phytotoxic. Future studies of B use and its long-term effect on MR remediation should be conducted under field conditions.

Root colonization by arbuscular mycorrhizal (AM) fungi is a complex and finely tuned process. Previous studies have shown that, among other plant hormones, auxin plays a role in this process but the specific involvement of Aux/IAAs, the key regulators of auxin responses, is still unknown. In this study, we addressed the role of the tomato Sl-IAA27 during AM symbiosis by using Sl-IAA27-RNAi and pSL-IAA27::GUS stable tomato lines. The data show that Sl-IAA27 expression is up-regulated by the AM fungus and that silencing of Sl-IAA27 has a negative impact on AM colonization. Sl-IAA27-silencing resulted in down-regulation of three genes involved in strigolactone synthesis, NSP1, D27 and MAX1, and treatment of Sl-IAA27-silenced plants with the strigolactone analog GR24 complemented their mycorrhizal defect phenotype. Overall, the study identified an Aux/IAA gene as a new component of the signaling pathway controlling AM fungal colonization in tomato. This gene is proposed to control strigolactone biosynthesis via the regulation of NSP1., (No claim to original US government works New Phytologist © 2016 New Phytologist Trust.)

Arbuscular mycorrhizal (AM) fungi have been used to alleviate heavy metal stress on plant growth and uptake of micro- and macroelements. A greenhouse pot experiment was conducted to verify the effects of AM fungus Rhizophagus irregularis on the growth, physiological characteristics, total Cd, and element uptake of Phragmites australis under different Cd stress (in the range of 0-20 mg L -1 ). The results showed that the symbiosis could effectively alleviate Cd toxicity with greater root biomass, higher photosynthesis rate, and lower levels of malonaldehyde (MDA) and proline than non-mycorrhizal plants could. However, reduced transpiration rate (Tr) and stomatal conductance (g s ) indicated R. irregularis protected host plants from Cd stress (≥5 mg L -1 ) via the stomatal closure. Although micro- and macroelements displayed differently in the presence of Cd, higher concentrations were still detected in mycorrhizal plants in contrast to non-mycorrhizal plants. Moreover, step multiple regression significantly demonstrated Pn max , stem diameter (Sd), and g s were the important factors with regard to total Cd uptake in the symbiosis, but Mn affected to non-mycorrhizal plants. These results suggested R. irregularis could alleviate the competition between Mn and Cd by altering plant physiology. This work clearly demonstrated that R. irregularis can be able to support P. australis growth better even though under high Cd stress (>1 mg L -1 ), suggesting its good potential for practical use in high Cd-contaminated areas.

Plant roots are the first parts of plants to face drought stress (DS), and thus root modification is important for plants to adapt to drought. We hypothesized that the roots of arbuscular mycorrhizal (AM) plants exhibit better adaptation in terms of morphology and phytohormones under DS. Trifoliate orange seedlings inoculated with Diversispora versiformis were subjected to well-watered (WW) and DS conditions for 6 weeks. AM seedlings exhibited better growth performance and significantly greater number of 1 st , 2 nd , and 3 rd order lateral roots, root length, area, average diameter, volume, tips, forks, and crossings than non-AM seedlings under both WW and DS conditions. AM fungal inoculation considerably increased root hair density under both WW and DS and root hair length under DS, while dramatically decreased root hair length under WW but there was no change in root hair diameter. AM plants had greater concentrations of indole-3-acetic acid, methyl jasmonate, nitric oxide, and calmodulin in roots, which were significantly correlated with changes in root morphology. These results support the hypothesis that AM plants show superior adaptation in root morphology under DS that is potentially associated with indole-3-acetic acid, methyl jasmonate, nitric oxide, and calmodulin levels.

Atrazine residue in the environment continually damages plants and therefore requires immediate attention and effective development of methods for its decontamination. The effects of Funnelliformis mosseae inoculation on growth and physiology in atrazine-treated Canna indica L. var. flava Roxb. were investigated. At atrazine concentrations up to 15 mg L -1 , the growth of C. indica plants were negatively affected. Inoculation with F. mosseae alleviated the atrazine inhibition of plant growth and biomass. Furthermore, the chlorophyll content and root function increased under F. mosseae inoculation, and the oxidative stress of malondialdehyde, peroxidase, and superoxide dismutase activities induced by atrazine were also alleviated by F. mosseae inoculation. The removal rate of atrazine by untreated C. indica was significant, with removal rates of 20.5-55.3% by the end of a 14-day experiment; however, F. mosseae inoculation increased the removal rate to 35.6-75.1%. In conclusion, F. mosseae inoculation can alleviate the damage induced by atrazine in C. indica. Accordingly, C. indica inoculated with F. mosseae has excellent potential to be used in phytoremediation in habitats polluted by high atrazine concentrations.