Non-mycorrhizal Research Articles

Carbon allocation and partitioning in Populus tremuloides are modulated by ectomycorrhizal fungi under phosphorus limitation.

The fate of carbon (C) captured by forest trees during photosynthesis is influenced by the supply of other resources. Fixed C may be partitioned among biomolecules within the leaf and/or allocated throughout the tree to growth, storage and maintenance activities. Phosphorus (P) availability often limits tree productivity due to its high biological demand and strong interactions with soil minerals. As ectomycorrhizal (ECM) fungi play critical roles in enhancing phosphate (Pi) acquisition by their hosts, these symbioses will influence the fate of C within trees and forested ecosystems. Using Populus tremuloides Michx. (trembling aspen) in symbiosis with Laccaria bicolor (Marie) P.D. Orton or Paxillus involutus (Batsch) Fr., we assessed C acquisition, allocation and partitioning under Pi limitation, specifically focusing on primary and secondary C compounds. Both ECM fungi moderated the effects of low P on photosynthesis and C partitioning among carbohydrates and secondary metabolites by sustaining Pi uptake and translocation in P. tremuloides under Pi limitation. As leaf P declined, reductions in photosynthesis were accompanied by significant shifts in C partitioning from nonstructural carbohydrates (NSCs) to phenolic glycosides and tannins. Carbon partitioning in roots exhibited more complex patterns, with distinct increases in NSCs in nonmycorrhizal (NM) plants under Pi limitation that were not evident in plants colonized by either ECM symbiont. In general, aspen colonized by L. bicolor exhibited C partitioning patterns intermediate between those of NM and P. involutus aspen. The C cost of symbiosis was pronounced for plants supporting P. involutus, where ECM plants exhibited maintenance of photosynthesis yet reduced biomass in comparison with NM and L. bicolor aspen under Pi replete conditions. Our results indicate that the ECM symbiosis affects the disposition of C in forest trees in part by altering the acquisition of other limiting resources from soils, but also through ECM species-specific influences on host physiology. This modulation of C partitioning will have broad implications for forest ecosystem C capture, storage and cycling where nutrient resources may be limited.

Influence of arbuscular mycorrhizal fungi on the functional mechanisms associated with drought tolerance in carob (Ceratonia siliqua L.)

Arbuscular mycorrhizal symbiosis strengthened the mechanisms developed by carob to withstand drought stress, including improved water relations, increased cell wall rigidity and osmolytes accumulation, and enhanced oxidative stress alleviation. The present investigation was carried out to provide more insight into the influence of arbuscular mycorrhizal fungi (Funneliformis mosseae, Rhizophagus fasciculatus, and Rhizophagus intraradices) on drought tolerance of carob. Non-mycorrhizal (NM) and arbuscular mycorrhizal (AM) carob plants were subjected to two watering regimes, 75% of field capacity (well water) or 25% of field capacity (water stress). Obtained results showed that stressed AM plants exhibited increased performance in terms of growth and biomass production, water and nutrient acquisition, and oxidative stress alleviation compared to NM plants. In fact, under limited water regime, AM plants maintained high stomatal conductance and high relative water content (over 94%) due to their high water and nutrient uptake efficiency. Moreover, AM plants especially those associated with F. mosseae maintained high membrane integrity (over 80%), high cell wall rigidity, and high leaf water potential and osmotic potential at full turgor and at turgor loss, while these parameters steeply decreased in NM plants. Furthermore, drought-stressed AM plants showed decreased hydrogen peroxide and malondialdehyde contents associated with increased activities of superoxide dismutase, ascorbate peroxidase, guaiacol peroxidase, and catalase compared to their relative NM plants. Thus, AMF strengthened the mechanisms involved in drought tolerance of carob by improving water relations, increasing cell wall rigidity, and enhancing oxidative stress alleviation. Funneliformis mosseae was the most effective in improving carob tolerance to drought stress.

Ectomycorrhizal fungi reduce the light compensation point and promote carbon fixation of Pinus thunbergii seedlings to adapt to shade environments

We examined the effects of three ectomycorrhizal (ECM) symbionts on the growth and photosynthesis capacity of Japanese black pine (Pinus thunbergii) seedlings and estimated physiological and photosynthetic parameters such as the light compensation point (LCP), biomass, and phosphorus (Pi) concentration of P. thunbergii seedlings. Through this investigation, we documented a new role of ectomycorrhizal (ECM) fungi: enhancement of the survival and competitiveness of P. thunbergii seedlings under low-light condition by reducing the LCP of seedlings. At a CO2 concentration of 400 ppm, the LCP of seedlings with ECM inoculations was 40–70 μmol photons m−2 s−1, significantly lower than that of non-mycorrhizal (NM) seedlings (200 μmol photons m−2 s−1). In addition, photosynthetic carbon fixation (Pn) increased with light intensity and CO2 level, and the Pn of ECM seedlings was significantly higher than that of NM seedlings; Pisolithus sp. (Pt)- and Laccaria amethystea (La)-mycorrhizal seedlings had significantly lower Pn than Cenococcum geophilum (Cg)-mycorrhizal seedlings. However, La-mycorrhizal seedlings exhibited the highest fresh weight, relative water content (RWC), and the lowest LCP in the mycorrhizal group. Concomitantly, ECM seedlings showed significantly increased chlorophyll content of needles and higher Pi concentrations compared to NM seedlings. Overall, ECM symbionts promoted growth and photosynthesis while reducing the LCP of P. thunbergii seedlings. These findings indicate that ECM fungi can enhance the survival and competitiveness of host seedlings under low light.

Arbuscular Mycorrhiza Augments Arsenic Tolerance in Wheat (Triticum aestivum L.) by Strengthening Antioxidant Defense System and Thiol Metabolism.

Arbuscular mycorrhiza (AM) can help plants to tolerate arsenic (As) toxicity. However, plant responses are found to vary with the host plant and the AM fungal species. The present study compares the efficacy of two AM fungi Rhizoglomus intraradices (M1) and Glomus etunicatum (M2) in amelioration of As stress in wheat (Triticum aestivum L. var. HD-2967). Mycorrhizal (M) and non-mycorrhizal (NM) wheat plants were subjected to four levels of As (0, 25, 50, and 100 mg As kg-1 soil). Although As additions had variable effects on the percentage of root colonized by the two fungal inoculants, each mycobiont conferred benefits to the host plant. Mycorrhizal plants continued to display better growth than NM plants. Formation of AM helped the host plant to overcome As-induced P deficiency and maintained favorable P:As ratio. Inoculation of AMF had variable effects on the distribution of As in plant tissues. While As translocation factor decreased in low As (25 mg kg-1 soil), it increased under high As (50 and 100 mg As kg-1 soil). Further As translocation to grain was reduced (As grain:shoot ratio) in M plants compared with NM plants. Arsenic-induced oxidative stress (generation of H2O2 and lipid peroxidation) in plants reduced significantly by AMF inoculation. The alleviation potential of AM was more evident with increase in severity of As stress. Colonization of AMF resulted in higher activities of the antioxidant enzymes (superoxide dismutase, catalase, and guaiacol peroxidase). It increased the concentrations of the antioxidant molecules (carotenoids, proline, and α-tocopherol) than their NM counterparts at high As addition level. Comparatively higher activities of enzymes of glutathione-ascorbate cycle in M plants led to higher ascorbate:dehydroascorbate (AsA:DHA) and glutathione:glutathione disulphide (GSH:GSSG) ratios. Inoculation by AMF also augmented the glyoxalase system by increasing the activities of both glyoxalase I and glyoxalase II enzymes. Mycorrhizal colonization increased concentrations of cysteine, glutathione, non-protein thiols, and activity of glutathione-S-transferase that facilitated sequestration of As into non-toxic complexes. The study reveals multifarious role of AMF in alleviation of As toxicity.

Plant mycorrhizal status, but not type, shifts with latitude and elevation in Europe

AbstractAimIdentifying the factors that drive large‐scale patterns of biotic interaction is fundamental for understanding how communities respond to changing environmental conditions. Mycorrhizal symbiosis is a key interaction between fungi and most vascular plants. Whether plants are obligately (OM) or facultatively (FM) mycorrhizal, and which mycorrhizal type they form – arbuscular mycorrhizal (AM), ectomycorrhizal (ECM), ericoid mycorrhizal (ERM) or non‐mycorrhizal (NM) – can have strong implications for plant species distribution at the continental scale and on the responses of plants to environmental gradients.LocationEurope, north of 43° latitude and excluding Russia, Belarus and Moldova.Time periodUndefined.Major taxa studiedVascular plants.MethodsUsing published sources, we compiled the most complete dataset yet of plant mycorrhizal and geographical information for Europe, comprising 1442 plant species. We mapped the European distributions of plant mycorrhizal status (OM and FM) and type (AM, ECM, ERM and NM) and analysed their relationships with climatic, edaphic and plant productivity drivers on a 50 km × 50 km equal‐area grid.ResultsThe distribution of mycorrhizal types in Europe was driven by mean temperature, soil pH and productivity. AM plant species predominated throughout the region, but at higher latitudes the share of NM and, to a lesser extent, ECM and ERM species increased. FM species predominated over OM species, and this increased with latitude and was dependent on temperature drivers. The high share of OM species in the central European mountains indicates a possible influence of historical glacial refugia.Main conclusionsOur results challenge the prevailing view of parallel trends in the latitudinal and elevational distribution of mycorrhizal types and demonstrate distinctive responses of plants with different mycorrhizal status to climatic, edaphic and biogeographical drivers at the European scale.

Comparative transcriptome analysis between Solanum lycopersicum L. and Lotus japonicus L. during arbuscular mycorrhizal development

ABSTRACTArbuscular mycorrhizal (AM) fungi form symbiotic associations with diverse plant species. The AM fungi enhance mineral uptake from the soil, which benefits the growth of the host plants. Previous microarray and RNA-seq analyses have identified a large number of AM-responsive plant genes. However, little is known whether the gene expression profile of mycorrhiza is different among genetically distant plant species. The aim of this study was to assess the conservation and divergence of AM-responsive genes between two different hosts, Solanum lycopersicum L. and Lotus japonicus L., during AM development using RNA-seq data. In each host plant, gene expression was compared between AM roots and non-mycorrhizal (NM) roots. Potential orthologs of AM-responsible genes between S. lycopersicum and L. japonicus were identified with reciprocal BLAST searches. Only one quarter to one third of the AM-inducible genes in each plant species were co-upregulated in both species. The co-upregulated genes included those known to be essential for AM development and function. The co-upregulated genes exhibited a wide range of fold changes in the AM symbiosis, and the fold change value for individual co-upregulated genes was positively correlated between the two hosts. Most of the species-dependent upregulated genes exhibited low levels of induction. We also analyzed gene expression in AM fungi colonizing roots of S. lycopersicum and L. japonicus. Overall, the gene expression profiles of Rhizophagus irregularis were similar among the roots of the two different hosts, although hundreds of fungal genes were differentially expressed between the two hosts. In particular, genes related to the mitochondrial electron transport chain were highly expressed in AM fungi colonizing L. japonicus roots, indicating that adenosine triphosphate (ATP) production was enhanced in the L. japonicus–R. irregularis symbiosis. Overall, these results show that a certain proportion of AM-responsive genes is conserved across plant species. The species-dependent AM-responsive genes may be related to the physiological differences between AM and NM roots in each plant species.

Arbuscular mycorrhizal fungus colonization in Nicotiana tabacum decreases the rate of both carboxylate exudation and root respiration and increases plant growth under phosphorus limitation

Under phosphorus (P) limitation, plants tend to maximize their efficiency of P acquisition by increasing the exudation of root carboxylates, such as citrate, whose synthesis is mediated by respiration via the alternative oxidase (AOX) in cluster roots. However, high respiratory costs related to nutrient uptake are associated with slower plant growth, whereas arbuscular mycorrhizal (AM) fungus colonization increases plant growth and decreases the exudation of citrate and malate. Thus, the present research is based on the hypothesis that AM fungus colonization will decrease root respiration via the alternative pathway and the amount of carboxylates in the rhizosphere, and increase plant growth. We used the oxygen-isotope-fractionation technique to study the in vivo respiratory activities of the cytochrome and the alternative oxidase pathways (COP and AOP) in AM and nonmycorrhizal (NM) tobacco plants grown under P-sufficient (0.25 mM) and P-limiting (0.025 mM) conditions in sand. The amount of root exudates in the rhizosphere, total biomass and root P content were determined. Under sufficient P, the amount of citrate and malate was higher in NM plants, while no differences were found in respiration between NM and AM plants. On the other hand, low P increases the exudation of citrate and respiration via AOP in NM plants, while it does not affect the amount of carboxylates, nor the respiratory rate in AM plants. Biomass production was reduced only in NM plants under low P. Our results highlight that AM fungus colonization decreases the rate of root respiration and the exudation of citrate and malate, whilst increasing plant growth. Conversely, respiration, via AOX, is associated with the exudation of citrate and less biomass production in NM roots.

Mycorrhizal fungi influence on silver uptake and membrane protein gene expression following silver nanoparticle exposure

The rapid growth of nanotechnology and the high demand for nanomaterial use have greatly increased the risk of particle release into the environment. Understanding nanomaterial interactions with crop species and their associated microorganisms is critical to food safety and security. In the current study, tomato was inoculated with mycorrhizal fungi and subsequently exposed to 12, 24, or 36 mg/kg of 2- or 15-nm silver nanoparticles (Ag-NPs). Mycorrhizal (M) and non-mycorrhizal (NM) tomatoes exposed to 36 mg/kg of 2-nm Ag-NPs accumulated 1300 and 1600 μg/g silver in their tissues, respectively. Mycorrhizal plants accumulated 14% less silver compared to non-mycorrhizal plants. To begin to understand the mechanisms by which plants accumulate NPs, the expression of two aquaporin channel genes, the plasma membrane intrinsic protein (PIP) and the tonoplast membrane intrinsic protein (TIP), and one potassium channel (KC) gene were studied. In non-mycorrhizal plants, the expression of KC, PIP, and TIP was eight, five, and nine times higher than the control, respectively. These expressions for mycorrhizal plants were 5.8, 3.5, and 2 times higher than controls, respectively. The expression of KC and PIP, which are located on the plasma membrane, was 3.5 and 2.5, respectively, times higher than TIP, which is located on the tonoplast. PIP expression was significantly higher in NM tomatoes exposed to 12 mg/kg of 2-nm Ag-NPs compared to M plants. These results show that mycorrhizal colonization decreases Ag accumulation in NP-exposed plants and also moderates changes in expression level of membrane transport proteins.

Paxillus involutus-Facilitated Cd2+ Influx through Plasma Membrane Ca2+-Permeable Channels Is Stimulated by H2O2 and H+-ATPase in Ectomycorrhizal Populus × canescens under Cadmium Stress.

Using a Non-invasive Micro-test Technique, flux profiles of Cd2+, Ca2+, and H+ were investigated in axenically grown cultures of two strains of Paxillus involutus (MAJ and NAU), ectomycorrhizae formed by these fungi with the woody Cd2+-hyperaccumulator, Populus × canescens, and non-mycorrhizal (NM) roots. The influx of Cd2+ increased in fungal mycelia, NM and ectomycorrhizal (EM) roots upon a 40-min shock, after short-term (ST, 24 h), or long-term (LT, 7 days) exposure to a hydroponic environment of 50 μM CdCl2. Cd2+ treatments (shock, ST, and LT) decreased Ca2+ influx in NM and EM roots but led to an enhanced influx of Ca2+ in axenically grown EM cultures of the two P. involutus isolates. The susceptibility of Cd2+ flux to typical Ca2+ channel blockers (LaCl3, GdCl3, verapamil, and TEA) in fungal mycelia and poplar roots indicated that the Cd2+ entry occurred mainly through Ca2+-permeable channels in the plasma membrane (PM). Cd2+ treatment resulted in H2O2 production. H2O2 exposure accelerated the entry of Cd2+ and Ca2+ in NM and EM roots. Cd2+ further stimulated H+ pumping activity benefiting NM and EM roots to maintain an acidic environment, which favored the entry of Cd2+ across the PM. A scavenger of reactive oxygen species, DMTU, and an inhibitor of PM H+-ATPase, orthovanadate, decreased Ca2+ and Cd2+ influx in NM and EM roots, suggesting that the entry of Cd2+ through Ca2+-permeable channels is stimulated by H2O2 and H+ pumps. Compared to NM roots, EM roots exhibited higher Cd2+-fluxes under shock, ST, and LT Cd2+ treatments. We conclude that ectomycorrhizal P. × canescens roots retained a pronounced H2O2 production and a high H+-pumping activity, which activated PM Ca2+ channels and thus facilitated a high influx of Cd2+ under Cd2+ stress.

Mutualism Persistence and Abandonment during the Evolution of the Mycorrhizal Symbiosis.

Mutualistic symbioses with mycorrhizal fungi are widespread in plants. The majority of plant species associate with arbuscular mycorrhizal (AM) fungi. By contrast, the minority associate with ectomycorrhizal (EM) fungi, have abandoned the symbiosis and are nonmycorrhizal (NM), or engage in an intermediate, weakly AM symbiosis (AMNM). To understand the processes that maintain the mycorrhizal symbiosis or cause its loss, we reconstructed its evolution using a ∼3,000-species seed plant phylogeny integrated with mycorrhizal state information. Reconstruction indicated that the common ancestor of seed plants most likely associated with AM fungi and that the EM, NM, and AMNM states descended from the AM state. Direct transitions from the AM state to the EM and NM states were infrequent and generally irreversible, implying that natural selection or genetic constraint could promote stasis once a particular state evolved. However, the evolution of the NM state was more frequent via an indirect pathway through the AMNM state, suggesting that weakening of the AM symbiosis is a necessary precursor to mutualism abandonment. Nevertheless, reversions from the AMNM state back to the AM state were an order of magnitude more likely than transitions to the NM state, suggesting that natural selection favors the AM symbiosis over mutualism abandonment.

Sensitivity of jarrah (Eucalyptus marginata) to phosphate, phosphite, and arsenate pulses as influenced by fungal symbiotic associations.

Many plant species adapted to P-impoverished soils, including jarrah (Eucalyptus marginata), develop toxicity symptoms when exposed to high doses of phosphate (Pi) and its analogs such as phosphite (Phi) and arsenate (AsV). The present study was undertaken to investigate the effects of fungal symbionts Scutellospora calospora, Scleroderma sp., and Austroboletus occidentalis on the response of jarrah to highly toxic pulses (1.5 mmol kg−1 soil) of Pi, Phi, and AsV. S. calospora formed an arbuscular mycorrhizal (AM) symbiosis while both Scleroderma sp. and A. occidentalis established a non-colonizing symbiosis with jarrah plants. All these interactions significantly improved jarrah growth and Pi uptake under P-limiting conditions. The AM fungal colonization naturally declines in AM-eucalypt symbioses after 2–3 months; however, in the present study, the high Pi pulse inhibited the decline of AM fungal colonization in jarrah. Four weeks after exposure to the Pi pulse, plants inoculated with S. calospora had significantly lower toxicity symptoms compared to non-mycorrhizal (NM) plants, and all fungal treatments induced tolerance against Phi toxicity in jarrah. However, no tolerance was observed for AsV-treated plants even though all inoculated plants had significantly lower shoot As concentrations than the NM plants. The transcript profile of five jarrah high-affinity phosphate transporter (PHT1 family) genes in roots was not altered in response to any of the fungal species tested. Interestingly, plants exposed to high Pi supplies for 1 day did not have reduced transcript levels for any of the five PHT1 genes in roots, and transcript abundance of four PHT1 genes actually increased. It is therefore suggested that jarrah, and perhaps other P-sensitive perennial species, respond positively to Pi available in the soil solution through increasing rather than decreasing the expression of selected PHT1 genes. Furthermore, Scleroderma sp. can be considered as a fungus with dual functional capacity capable of forming both ectomycorrhizal and non-colonizing associations, where both pathways are always accompanied by evident growth and nutritional benefits.

Effectiveness of arbuscular mycorrhizal fungi in phytoremediation of lead- contaminated soil by vetiver grass

ABSTRACTA greenhouse experiment was conducted to evaluate the effectiveness of arbuscular mycorrhizal (AM) fungi in phytoremediation of lead (Pb)-contaminated soil by vetiver grass. Experiment was a factorial arranged in a completely randomized design. Factors included four Pb levels (50, 200, 400, and 800 mg kg−1) as Pb (NO3)2, AM fungi at three levels (non mycorrhizal (NM) control, Rhizophagus intraradices, Glomus versiforme). Shoot and root dry weights (SDW and RDW) decreased as Pb levels increased. Mycorrhizal inoculation increased SDW and RDW compared to NM control. With mycorrhizal inoculation and increasing Pb levels, Pb uptake of shoot and root increased compared to those of NM control. Root colonization increased with mycorrhizal inoculation but decreased as Pb levels increased. Phosphorus concentration and uptake in shoot of plants inoculated with AM fungi was significantly higher than NM control at 200 and 800 mg Pb kg−1. The Fe concentration, Fe and Mn uptake of shoot in plants inoculated with Rhizophagus intraradices in all levels of Pb were significantly higher than NM control. Mycorrhizal inoculation increased Pb extraction, uptake and translocation efficiencies. Lead translocation factor decreased as Pb levels increased; however inoculation with AM fungi increased Pb translocation.

Contributions of an arbuscular mycorrhizal fungus to growth and physiology of loquat (Eriobotrya japonica) plants subjected to drought stress

Loquat (Eriobotrya japonica) is an evergreen tree with a shallow root system subjected to drought stress. We have found that AM fungi can alleviate drought stress by improving loquat nutrient uptake. However, the physiological mechanisms of improving drought tolerance have not been described so far in loquat mycorrhiza symbiosis. Funneliformis mosseae was used as arbuscular mycorrhizal fungus and loquat was selected as a model for an evergreen, woody plant. Thus, a pot experiment with four treatments was conducted. Growth, leaf water status, solute accumulation, oxidative damage to lipids, antioxidant activities, and phytohormones were evaluated by non-mycorrhizal (NM) and arbuscular mycorrhizal (AM) loquat plants growing under well-watered or drought-stressed conditions. Results showed that AM plants had higher dry-biomass production and leaf water potential than NM plants under drought-stressed conditions. The drought-stressed AM roots accumulated more proline than in NM roots, while not in leaves. Lipid peroxides of leaves and roots in drought-stressed AM plants were 26 and 61 % lower than in NM plants. The AM symbiosis may enhance osmotic adjustment in roots, contributing to maintaining a water potential gradient and water absorption from soil into the roots. The cumulative effects increased the AM plant tolerance to drought stress. The results are compared to findings reported hitherto mainly from short-lived, herbaceous AM plants in the literature.

Uptake Kinetics of Arsenic in Upland Rice Cultivar Zhonghan 221 Inoculated with Arbuscular Mycorrhizal Fungi

Arbuscular mycorrhizal fungi (AMF) appear to be highly associated with arsenic (As) uptake in host plants because arsenate (As(V)) and phosphorus (P) share the same transporter, whereby AMF can enhance P uptake. A short-term experiment was conducted for low- (0 to 0.05 mM As) and high-affinity (0 to 2.5 mM As) uptake systems, to investigate the AMF role on As uptake mechanism in plants, which may explain As uptake kinetics in upland rice cultivar: Zhonghan 221. When concentration of As ranged from 0 to 0.05 mM, Funneliformis geosporum (Fg) significantly decreased arsenite (As(III)) and monomethylarsonicacid (MMA) uptake when (p < 0.05) compared to non-mycorrhizal (NM) treatment, since the major route for (As(III)) in rice roots—rice silicon transporter Lsi1 would be influenced by Fg inoculation at high As concentrations. Fg can also reduce As(V) uptake significantly (p < 0.05) under both uptake systems relative to NM treatment, whereas, Funneliformis mosseae (Fm) increased As(V) and MMA uptake in rice roots, with MMA uptake rate generally lower than As(III) and As(V). Using suitable AMF species inoculation with rice, As uptake and accumulation in rice grains can be reduced and the risk to human health, once consumed, can be minimized.

Growth and photosynthetic responses of ectomycorrhizal pine seedlings exposed to elevated Cu in soils.

It is still controversial whether ectomycorrhizal (ECM) mycelia filter out toxic metals in nutrient absorption of host trees. In this study, pine (Pinus densiflora) seedlings colonized by Cu-sensitive and Cu-tolerant ECM species were exposed to a wide spectrum of soil Cu concentrations to investigate functions of ECM fungi under Cu stress. The photosynthetic rates of intact needles were monitored in situ periodically. The biomass and elements of plants were also measured after harvest. The ameliorating effect of ECM infection on host plants exposed to toxic stress was metal concentration specific. Under lower-level Cu stress, ECM fungi increased seedling performance, while ECM seedlings accumulated more Cu than nonmycorrhizal (NM) seedlings. Under higher-level Cu stress, photosynthesis decreased well before visible symptoms of Cu toxicity appeared. The reduced photosynthesis and biomass in ECM seedlings compared to NM seedlings under higher Cu conditions were also accompanied by lower phosphorus in needles. There was no marked difference between the two fungal species. Our results indicate that the two ECM fungi studied in our system may not have an ability to selectively eliminate Cu in nutrient absorption and may not act as effective barriers that decrease toxic metal uptake into host plants.

Effects of Arbuscular Mycorrhizal Fungi on Metals Uptake, Physiological and Biochemical Response of &amp;lt;i&amp;gt;Medicago Sativa&amp;lt;/i&amp;gt; L. with Increasing Zn and Cd Concentrations in Soil

The effect of mycorrhizal symbiosis on metal accumulation and plant tolerance are not commonly studied in medicinal plants under metal stress. The objective of this study was to assess the impact of mycorrhiza on alfalfa plants with the increase of Zn and Cd toxicity. The experiment was conducted under controlled laboratory conditions. Zinc (Zn) and cadmium (Cd) uptake, some biochemical and physiological parameters were studied in eight-week-old alfalfa plants in response to inoculation or not with arbuscular mycorrhizal fungi (AMF) and with the increase of Zn (0, 100, 300, 900 mg·kg-1) and Cd concentrations (0, 100, 300, 600 mg·kg-1) in soil. The results showed that mycorrhizal (M) plants exhibited tolerance to Zn and Cd up to 300 mg·kg-1 in comparison to non-mycorrhizal (NM) plants which exhibited a significant growth reduction at the same soil Zn and Cd level. M inoculation reduced the Zn and Cd accumulation in shoot and showed higher Zn and Cd contents in roots which showed a different Zn and Cd distribution in AMF associated or non-associated plants. Mycorrhizal plants increased phosphorus (P) contents at all Zn and Cd concentrations except the highest (600 and 900 mg·kg-1) leading significant alterations in biochemical contents such as proline, antioxidant enzymes in leaves and also in nutrients (N, P, K, Cu, Ni, Fe, Mn). Zn and cadmium toxicity cause to increase the proline content in shoot of NM plants, however, proline contents are lower in M plants. Results confirmed that AMF protected alfalfa plants against Zn and Cd toxicity. Mycorrhizal colonization was able to form an efficient symbiosis with alfalfa plants in moderately contaminated Zn and Cd soils (300 mg·kg-1) and play an important role in food quality and safety.

Mycorrhizas effects on nutrient interception in two riparian grass species

Effects of arbuscular mycorrhizal (AM) fungi on plant growth and soil nutrient depletion are well known, but their roles as nutrient interceptor in riparian areas are less clear. The effects of AM fungi on growth, soil nutrient depletion and nutrient leaching were investigated in columns with two riparian grass species. Mycorrhizal and non mycorrhizal (NM) plants were grown in a mixture of riparian soil and sand (60% and 40%, w/w respectively) for 8 weeks under glasshouse conditions. Mycorrhizal colonization, AM external hyphae development, plant growth, nutrient uptake and NO 3 , NH 4 and available P in soil and leachate were measured. Mycorrhizal fungi highly colonized roots of exotic grass Phalaris aquatica and significantly increased plant growth and nutrient uptake. Columns containing of AM Phalaris aquatica had higher levels of AM external hyphae, lower levels of NO 3 , NH 4 and available P in soil and leachate than NM columns. Although roots of native grass Austrodanthonia caespitosa had moderately high levels of AM colonization and AM external hyphae in soil, AM inoculation had no significant effects on plant growth, soil and leachate concentration of NO 3 and NH 4 . But AM inoculation decreased available soil P concentration in deeper soil layer and had no effects on dissolved P in leachate. Although both grass species had nearly the same biomass, results showed that leachate collected from Austrodanthonia caespitosa columns significantly had lower levels of NO 3 , NH 4 and dissolve P than leachate from exotic Phalaris aquatica columns. Taken together, these data shows that native plant species intercept higher nutrient than exotic plant species and had no responsiveness to AM fungi related to nutrient leaching, but AM fungi play an important role in interception of nutrient in exotic plant species.

Mineral nutrition of campos rupestres plant species on contrasting nutrient-impoverished soil types.

In Brazil, the campos rupestres occur over the Brazilian shield, and are characterized by acidic nutrient-impoverished soils, which are particularly low in phosphorus (P). Despite recognition of the campos rupestres as a global biodiversity hotspot, little is known about the diversity of P-acquisition strategies and other aspects of plant mineral nutrition in this region. To explore nutrient-acquisition strategies and assess aspects of plant P nutrition, we measured leaf P and nitrogen (N) concentrations, characterized root morphology and determined the percentage arbuscular mycorrhizal (AM) colonization of 50 dominant species in six communities, representing a gradient of soil P availability. Leaf manganese (Mn) concentration was measured as a proxy for carboxylate-releasing strategies. Communities on the most P-impoverished soils had the highest proportion of nonmycorrhizal (NM) species, the lowest percentage of mycorrhizal colonization, and the greatest diversity of root specializations. The large spectrum of leaf P concentration and variation in root morphologies show high functional diversity for nutritional strategies. Higher leaf Mn concentrations were observed in NM compared with AM species, indicating that carboxylate-releasing P-mobilizing strategies are likely to be present in NM species. The soils of the campos rupestres are similar to the most P-impoverished soils in the world. The prevalence of NM strategies indicates a strong global functional convergence in plant mineral nutrition strategies among severely P-impoverished ecosystems.

Arbuscular mycorrhiza increase artemisinin accumulation in Artemisia annua by higher expression of key biosynthesis genes via enhanced jasmonic acid levels.

It is becoming increasingly evident that the formation of arbuscular mycorrhiza (AM) enhances secondary metabolite production in shoots. Despite mounting evidence, relatively little is known about the underlying mechanisms. This study suggests that increase in artemisinin concentration in Artemisia annua colonized by Rhizophagus intraradices is due to altered trichome density as well as transcriptional patterns that are mediated via enhanced jasmonic acid (JA) levels. Mycorrhizal (M) plants had higher JA levels in leaf tissue that may be due to induction of an allene oxidase synthase gene (AOS), encoding one of the key enzymes for JA production. Non-mycorrhizal (NM) plants were exogenously supplied with a range of methyl jasmonic acid concentrations. When leaves of NM and M plants with similar levels of endogenous JA were compared, these matched closely in terms of shoot trichome density, artemisinin concentration, and transcript profile of artemisinin biosynthesis genes. Mycorrhization increased artemisinin levels by increasing glandular trichome density and transcriptional activation of artemisinin biosynthesis genes. Transcriptional analysis of some rate-limiting enzymes of mevalonate and methyl erythritol phosphate (MEP) pathways revealed that AM increases isoprenoids by induction of the MEP pathway. A decline in artemisinin concentration in shoots of NM and M plants treated with ibuprofen (an inhibitor of JA biosynthesis) further confirmed the implication of JA in the mechanism of artemisinin production.

Increased photosynthetic acclimation in alfalfa associated with arbuscular mycorrhizal fungi (AMF) and cultivated in greenhouse under elevated CO2

Medicago sativa L. (alfalfa) can exhibit photosynthetic down-regulation when grown in greenhouse conditions under elevated atmospheric CO2. This forage legume can establish a double symbiosis with nitrogen fixing bacteria and arbuscular mycorrhizal fungi (AMF), which may increase the carbon sink effect of roots. Our aim was to assess whether the association of alfalfa with AMF can avoid, diminish or delay the photosynthetic acclimation observed in previous studies performed with nodulated plants. The results, however, showed that mycorrhizal (M) alfalfa at the end of their vegetative period had lower carbon (C) discrimination than non-mycorrhizal (NM) controls, indicating photosynthetic acclimation under ECO2 in plants associated with AMF. Decreased C discrimination was due to the acclimation of conductance, since the amount of Rubisco and the expression of genes codifying both large and small subunits of Rubisco were similar or slightly higher in M than in NM plants. Moreover, M alfalfa accumulated a greater amount of soluble sugars in leaves than NM plants, thus favoring a down-regulation effect on photosynthetic rates. The enhanced contents of sugars in leaves coincided with a reduced percentage of arbuscules in roots, suggesting decreased sink of carbohydrates from shoots to roots in M plants. The shorter life cycle of alfalfa associated with AMF in comparison with the NM controls may also be related to the accelerated photosynthetic acclimation in M plants. Further research is needed to clarify to what extent this behavior could be extrapolated to alfalfa cultivated in the field and subjected to periodic cutting of shoots under climatic change scenarios.