Towards growth of arbuscular mycorrhizal fungi independent of a plant host

When surface-sterilized spores of the arbuscular mycorrhizal fungus (AMF) Glomus intraradices Sy167 were germinated on agar plates in the slightly modified minimum mineral medium described by G. Bécard and J. A. Fortin (New Phytol. 108:211-218, 1988), slime-forming bacteria, identified as Paenibacillus validus, frequently grew up. These bacteria were able to support growth of the fungus on the agar plates. In the presence of P. validus, hyphae branched profusely and formed coiled structures. These were much more densely packed than the so-called arbuscule-like structures which are formed by AMF grown in coculture with carrot roots transformed with T-DNA from Agrobacterium rhizogenes. The presence of P. validus alone also enabled G. intraradices to form new spores, mainly at the densely packed hyphal coils. The new spores were not as abundant as and were smaller than those formed by AMF in the monoxenic culture with carrot root tissues, but they also contained lipid droplets and a large number of nuclei. In these experiments P. validus could not be replaced by bacteria such as Escherichia coli K-12 or Azospirillum brasilense Sp7. Although no conditions under which the daughter spores regerminate and colonize plants have been found yet, and no factor(s) from P. validus which stimulates fungal growth has been identified, the present findings might be a significant step forward toward growth of AMF independent of any plant host.     

Toxicity of diesel fuel to germination,growth and colonization of <Emphasis Type="Italic">Glomus intraradices</Emphasis> in soil and <Emphasis Type="Italic">in vitro</Emphasis> transformed carrot root cultures

Phytoremediation is the use of selected plants to decontaminate polluted environments. Arbuscular mycorrhizal fungi (AMF) may potentially be useful for phytoremediation, but it is not known how petroleum hydrocarbons influence AMF spore germination and hyphal growth. To address this question, germination of spores and germ tube growth of Glomus intraradices Schenck and Smith and Glomus aggregatum Schenck and Smith were assessed in soil contaminated with up to 3% (w/v) of F2 diesel oil or HAGO reference oil. Hyphal growth, colonization and progeny spore production were assessed in vitro using transformed root cultures of Daucus carota and G. intraradices spores in a F2 diesel contaminated medium. In addition, extraradical hyphal growth of G. intraradices colonizing Daucus carota in the presence of F2 diesel was studied. Neither F2 diesel nor HAGO reference oil affected spore germination or germ tube growth in soil. However, in the presence of plant roots, germ tube growth of G. intraradices was reduced and delayed in the presence of F2 diesel and root colonization was not detected. Hyphal growth of pre-colonized carrot roots by G. intraradices was reduced and delayed in F2 contaminated medium compared to controls. F2 diesel did not inhibit spore germination of these AMF species but did reduce colonization, germ tube and hyphal growth. These results suggest that AMF inoculum can be established in petroleum-contaminated sites. However, it may prove beneficial to plant pre-colonized plants to increase the probability of sufficient AMF colonization and growth. The likely mechanism(s) of petroleum toxicity in this plant-microbe system was discussed.     

Differential host plant performance as a function of soil arbuscular mycorrhizal fungal communities: experimentally manipulating co-occurring Glomus species

In order to evaluate host plant performance relative to different soil arbuscular mycorrhizal fungal (AMF) communities, Andropogon gerardii seedlings were grown with nine different AMF communities. The communities consisted of 0, 10, or 20 spores of Glomus etunicatum and 0, 10, or 20 spores of Glomus intraradices in all possible combinations. Spores were produced by fungal cultures originating on A. gerardii in a serpentine plant community; seeds of A. gerardii were collected at the same site. The experiment was performed in the greenhouse using a mixture of sterilized serpentine soil and sand to which naturally occurring non-mycorrhizal microbes were added. There was no difference in root AMF colonization rates between single species communities of either G. etunicatum or G. intraradices, but G. intraradices enhanced plant growth and G. etunicatum did not. However, plants grew larger with some combinations of G.␣intraradices plus G. etunicatum than with the same quantity of G. intraradices alone. These results suggest the potential for niche complementarity in the mycorrhizal fungi. That G. etunicatum only increased plant growth in the presence of G. intraradices could be illustrative of why AMF that appear to be parasitic or benign when examined in isolation are maintained within multi-species mycorrhizal communities in nature.     

Production and specificity of polyclonal antibodies against soluble proteins from the arbuscular mycorrhizal fungus Glomus intraradices

Rabbit polyclonal antibodies were produced against a soluble protein fraction from a vesicle and spore mixture of the arbuscular mycorrhizal fungus (AMF) Glomus intraradices. The protocol for isolation of vesicles and spores from plant roots was optimized to minimize debris contamination. Protein extract purification and preparation for immunization was adapted to increase protein content and immunogenicity. Active antisera were produced starting from the second boost immunization. Antibodies obtained were specific for surface antigens of AMF and revealed different patterns of soluble protein antigens in G. intraradices, G. constrictum and an unidentified Glomus species. Accepted: 6 December 2000     

Biological Control of Fusarium Wilt in Tomato Caused by Fusarium oxysporum f. sp. lycopersici by AMF Glomus intraradices and some Rhizobacteria

In the present study, the effects of the arbuscular mycorrhizal fungus (AMF) Glomus intraradices Schenck & Smith and four rhizobacteria (RB; 58/1 and D/2: Pseudomonas fluorescens biovar II; 17: P. putida; 21: Enterobacter cloacae), which are the important members of the rhizosphere microflora and biological control agents against plant diseases, were examined in the pathosystem of Fusarium oxysporum f. sp. lycopersici [(Sacc) Syd. et Hans] (FOL) and tomato with respect to morphological parameters (fresh and dry root weight) and phosphorous (P) concentration in the roots. Treatments with single and dual inoculation with G. intraradices and RB strains reduced disease severity by 8.6–58.6%. Individual bacteria inoculations were more effective than both the single AMF and dual (G. intraradices + RB) inoculations. In addition, the RB and G. intraradices enhanced dry root weight effectively. Significant increases in root weights were recorded particularly in the triple inoculations compared with single or dual inoculations. Compared with the non‐treated controls all biological control agents increased P‐content of treated roots of plants. Colonization with RB increased especially in triple (FOL + G. intraradices + RB) inoculations whereas colonization of G. intraradices was significantly decreased in treatment of FOL + G. intraradices compared with triple inoculations. The results suggest that suitable combinations of these biocontrol agents may ameliorate plant growth and health.     

Dependence of Arbuscular-Mycorrhizal Fungi on Their Plant Host for Palmitic Acid Synthesis

Lipids are the major form of carbon storage in arbuscular-mycorrhizal fungi. We studied fatty acid synthesis by Glomus intraradices and Gigaspora rosea. [¹⁴C]Acetate and [¹⁴C]sucrose were incorporated into a synthetic culture medium to test fatty acid synthetic ability in germinating spores (G. intraradices and G. rosea), mycorrhized carrot roots, and extraradical fungal mycelium (G. intraradices). Germinating spores and extraradical hyphae could not synthesize 16-carbon fatty acids but could elongate and desaturate fatty acids already present. The growth stimulation of germinating spores by root exudates did not stimulate fatty acid synthesis. 16-Carbon fatty acids (16:0 and 16:1) were synthesized only by the fungi in the mycorrhized roots. Our data strongly suggest that the fatty acid synthase activity of arbuscular-mycorrhizal fungi is expressed exclusively in the intraradical mycelium and indicate that fatty acid metabolism may play a major role in the obligate biotrophism of arbuscular-mycorrhizal fungi.     

Elemental composition of arbuscular mycorrhizal fungi at high salinity

We investigated the elemental composition of spores and hyphae of arbuscular mycorrhizal fungi (AMF) collected from two saline sites at the desert border in Tunisia, and of Glomus intraradices grown in vitro with or without addition of NaCl to the medium, by proton-induced X-ray emission. We compared the elemental composition of the field AMF to those of the soil and the associated plants. The spores and hyphae from the saline soils showed strongly elevated levels of Ca, Cl, Mg, Fe, Si, and K compared to their growth environment. In contrast, the spores of both the field-derived AMF and the in vitro grown G. intraradices contained lower or not elevated Na levels compared to their growth environment. This resulted in higher K:Na and Ca:Na ratios in spores than in soil, but lower than in the associated plants for the field AMF. The K:Na and Ca:Na ratios of G. intraradices grown in monoxenic cultures were also in the same range as those of the field AMF and did not change even when those ratios in the growth medium were lowered several orders of magnitude by adding NaCl. These results indicate that AMF can selectively take up elements such as K and Ca, which act as osmotic equivalents while they avoid uptake of toxic Na. This could make them important in the alleviation of salinity stress in their plant hosts.     

Influence of arbuscular mycorrhizal fungi on microbes and enzymes of soils from different cultivated densities of red clover

Effects of arbuscular mycorrhizal fungus (AMF)Glomus mosseae on plant growth, soil microbial populations and enzymes activities of soils were studied in red clover (Trifolium pratense L.) grown in pots at different cultivated densities. Seeds of red clover were sown with 50 g inoculums ofG. mosseae per pot. After a week, the plants were thinned to 20, 30, 40, 50 and 60 seedlings per pot. Three months after treatment, AMF inoculation significantly stimulated plant growth. Quantities of vesicles and spores, arbuscules and hyphae were the highest when 30 and 50 seedlings were grown per pot, respectively. However, no root was infected in control plants. In all the soil sites, the numbers of fungi and bacteria were followed in the order: root > root surface > rhizospheric. It was indicated that arbuscular mycorrhizal fungus decreased the numbers of fungi and bacteria but improved growth of actinomycetes. Compared to control plants, AMF stimulated activities of phosphatase and urease but decreased invertase.     

Response of mycorrhizal grapevine to Armillaria mellea inoculation: disease development and polyamines

A study was conducted with the vine rootstock Richter 110 (Vitis berlandieri Planch. x Vitis rupestris L.) in order to assess whether the colonisation by the arbuscular mycorrhizal fungus (AMF) Glomus intraradices (BEG 72) can delay the disease development in plants inoculated with the root-rot fungus Armillaria mellea (Vahl:Fr) Kummer, and to elucidate if the levels of polyamines (PAs) are modified in response to G. intraradices, A. mellea or by the dual infection. Four treatments were considered: control and G. intraradices-inoculated plants infected or not with A. mellea. Plant growth, mycorrhizal colonisation and disease development were monitored throughout the experiment. High performance liquid chromatography (HPLC) in combination with fluorescence spectrophotometry was used to separate and quantify free root and leaf polyamines. The slower development of pathogenic symptoms and the higher plant biomass of mycorrhizal plants inoculated with A. mellea indicate an increase of tolerance due to the AMF inoculation. The variations in free PA levels detected at the beginning of the pathogenic infection suggest that PAs may have a potential role in the signalling mechanisms of the tolerance of mycorrhizal plants against A. mellea.     

Aquaporin genes GintAQPF1 and GintAQPF2 from Glomus intraradices contribute to plant drought tolerance

Arbuscular mycorrhizal (AM) symbiosis, established between AM fungi (AMF) and roots of higher plants, occurs in most terrestrial ecosystems. It has been well demonstrated that AM symbiosis can improve plant performance under various environmental stresses, including drought stress. However, the molecular basis for the direct involvement of AMF in plant drought tolerance has not yet been established. Most recently, we cloned two functional aquaporin genes, GintAQPF1 and GintAQPF2, from AM fungus Glomus intraradices. By heterologous gene expression in yeast, aquaporin localization, activities and water permeability were examined. Gene expressions during symbiosis in expose to drought stress were also analyzed. Our data strongly supported potential water transport via AMF to host plants. As a complement, here we adopted the monoxenic culture system for AMF, in which carrot roots transformed by Ri-T DNA were cultured with Glomus intraradices in two-compartment Petri dishes, to verify the aquaporin gene functions in assisting AMF survival under polyethylene glycol (PEG) treatment. Our results showed that 25% PEG significantly upregulated the expression of two aquaporin genes, which was in line with the gene functions examined in yeast. We therefore concluded that the aquaporins function similarly in AMF as in yeast subjected to osmotic stress. The study provided further evidence to the direct involvement of AMF in improving plant water relations under drought stresses.     

Interaction between root growth allocation and mycorrhizal fungi in soil with patchy P distribution

Aims and Background

Many plants preferentially grow roots into P-enriched soil patches, but little is known about how the presence of arbuscular mycorrhizal fungi (AMF) affects this response.

Methods

Lotus japonicus (L.) was grown in a low-P soil with (a) no additional P, (b) homogeneous P (28 mg pot^?1), (c) low heterogeneous P (9.3 mg pot^?1), and (d) high heterogeneous P (28 mg pot^?1). Each P treatment was combined with one of three mycorrhiza treatments: no mycorrhizae, Glomus intraradices, indigenous AMF. Real-time PCR was used to assess the abundance of G. intraradices and the indigeneous AMF G. mosseae and G. claroideum.

Results

Mycorrhization and P fertilization strongly increased plant growth. Homogeneous P supply enhanced growth in both mycorrhizal treatments, while heterogeneous P fertilization increased biomass production only in treatments with indigenous AMF inoculation. Preferential root allocation into P-enriched soil was significant only in absence of AMF. The abundance of AMF species was similar in P-enriched and unfertilized soil patches.

Conclusion

Mycorrhization may completely override preferential root growth responses of plants to P- patchiness in soil. The advantage of this effect for the plants is to give roots more freedom to forage for other resources in demand for growth and to adapt to variable soil conditions.     

The Role of Mycorrhizae Associated with Vetiver Grown in Pb-/Zn-Contaminated Soils: Greenhouse Study

The effects of mycorrhizae on growth and uptake of N, P, Zn, and Pb by plants were investigated in a greenhouse trial using vetiver grass (Vetiveria zizanioides) as host. Inoculation of the host plants with arbuscular mycorrhizal fungi (AMF), Glomus mosseae and G. intraradices spores, significantly increased the growth and P uptake. Mycorrhizal colonization increased Pb and Zn uptake by plants under low soil metal concentrations (at 0 and 10 mg/kg of Pb or Zn), whereas under higher concentrations (at 100 and 1,000 mg/kg of Pb or Zn), it decreased Pb and Zn uptake. P concentration in soil was negatively correlated with mycorrhizal colonization as well as Zn or Pb concentrations. The results showed that inoculation of the host plants with AMF protects them from the potential toxicity caused by increased uptake of Pb and Zn, but the degree of protection varied according to the fungus and host plant combination. The potential of arbuscular mycorrhizae in phytoremediation of the Zn‐ or the Pb‐contaminated soils is discussed in this article.     

Functional diversity in arbuscular mycorrhizas: exploitation of soil patches with different phosphate enrichment differs among fungal species

Most terrestrial plant species form associations with arbuscular mycorrhizal fungi (AMF) that transfer soil P to the plant via their external hyphae. The distribution of nutrients in soils is typically patchy (heterogeneous) but little is known about the ability of AMF to exploit P patches in soil. This was studied by growing symbioses of Linum usitatissimum and three AMF (Glomus intraradices, G. mosseae and Gigaspora margarita) in pots with two side-arms, which were accessible to hyphae, but not to roots. Soil in one side-arm was either unamended (P0) or enriched with P; simultaneous labelling of this soil with ³²P revealed that G. intraradices responded to P enrichment both in terms of hyphal proliferation and P uptake, whereas the other AMF did not. Labelling with ³³P of P0 soil in the other side arm revealed that the increased P uptake by G. intraradices from the P-enriched patch was paralleled by decreased P uptake by other parts of the mycelium. This is the first demonstration of variation in growth and nutrient uptake by an AMF as influenced by a localized P enrichment of the soil. The results are discussed in the context of functional diversity of AMF.     

Role of native and exotic mycorrhizal symbiosis to develop morphological,physiological and biochemical responses coping with water drought of date palm,Phoenix dactylifera

Key Message

Arbuscular mycorrhizal (AM) symbiosis can improve date palm growth and alleviate drought-related impacts than non-mycorrhizal plants due to the ability of AMF for modifying plant metabolism and physiology.

Abstract

Date palm (Phoenix dactylifera L.) is an important agricultural and commercial crop in the North of Africa and Middle Eastern countries. During the last decade, date palm plantations were subjected to degradation due to an extensive exploitation and to drastic environmental conditions such as drought. Currently, there is a growing interest in the valorization of water due to environmental problems and economic aspects. The use of arbuscular mycorrhizal fungi (AMF) can offer a possibility to overcome these problems. The objective of this study was to study the influence of different Glomus species—Glomus intraradices, G. mosseae and Complex Aoufous (native AMF)—on the development of date palm grown under two water regimes (optimal irrigation, 75 % of field capacity or water deficit, 25 % of field capacity). Our results revealed that the beneficial effect of mycorrhizal symbiosis on plant growth depended on the fungal species and the water regime applied to the palm date seedling. While the native Complex Aoufous was the most effective in increasing the shoot height and biomass under well-watered conditions, G. intraradices was the most beneficial fungus for improving growth of plants that undergo restricted water supply. This positive effect of G. intraradices under drought conditions was not related to an enhancement of the antioxidant enzymatic activities in leaves; the association of palm date with G. intradices caused an increase in the elasticity of cell walls in leaves and allowed maintaining high water content in leaves without lowering leaf water potential under stressful conditions. The adequate selection of the AMF species is crucial for improving growth of palm date seedlings, and it must be in accordance with the water regime that will be applied to plants.     

Localization and speciation of arsenic in Glomus intraradices by synchrotron radiation spectroscopic analysis

The protective mechanisms employed by arbuscular mycorrhizal fungi (AMF) to reduce the toxic effects of arsenic on host plants remain partially unknown. The goal of this research was identifying the in situ localization and speciation of arsenic (As) in the AM fungus Rhizophagus intraradices [formerly named Glomus intraradices] exposed to arsenate [As(V)]. By using a two-compartment in vitro fungal cultures of R. intraradices-transformed carrot roots, microspectroscopic X-ray fluorescence (μ-XRF), and microspectroscopic X-ray absorption near edge structure (μ-XANES), we observed that As(V) is absorbed after 1 h in the hyphae of AMF. Three hours after exposure a decrease in the concentration of As was noticed and after 24 and 72 h no detectable As concentrations were perceived suggesting that As taken up was pumped out from the hyphae. No As was detected within the roots or hyphae in the root compartment zone three or 45 h after exposure. This suggests a dual protective mechanism to the plant by rapidly excluding As from the fungus and preventing As translocation to the plant root. μ-XANES data showed that gradual As(V) reduction occurred in the AM hyphae between 1 and 3 h after arsenic exposure and was completed after 6 h. Principal component analysis (PCA) and linear combination fitting (LCF) of μ-XANES data showed that the dominant species after reduction of As(V) by R. intraradices extra-radical hyphal was As(III) complexed with a reduced iron(II) carbonate compound. The second most abundant As species present was As(V)–iron hydroxides. The remaining As(III) compounds identified by the LCF analyses suggested these molecules were made of reduced As and S. These results increase our knowledge on the mechanism of As transport in AMF and validate our hypotheses that R. intraradices directly participates in arsenic detoxification. These fungal mechanisms may help AMF colonized plants to increase their tolerance to As at contaminated sites.     

Chemical alteration of the rhizosphere of the mycorrhizal-colonized wheat root

Plexiglass pot growth chamber experiments were conducted to evaluate the chemical alterations in the rhizosphere of mycorrhizal wheat roots after inoculation with Glomus intraradices [arbuscular mycorrhizal fungus (AMF)]. Exchange resins were used as sinks for nutrients to determine whether the inoculated plant can increase the solubility and the uptake of P and micronutrients. Treatments included: (1) soil (bulk soil); (2) AMF inoculation no P addition (I–P); (3) no inoculation with no P addition (NI–P); (4) AMF inoculation with addition of 50 mg P (kg soil)^–1 (I+P), and (5) no inoculation with addition of 50 mg P (kg soil)^–1 (NI+P). The AMF inoculum was added at a rate of four spores of G. intraradices (g soil)^–1. The exchange resin membranes were inserted vertically 5 cm apart in the middle of Plexiglass pots. Spring wheat (Triticum aestivum cv. Len) was planted in each Plexiglass pot and grown for 2 weeks in a growth chamber where water was maintained at field capacity. Rhizosphere pH and redox potential (Eh), nutrient bioavailability indices and mycorrhizal colonization were determined. Mycorrhizal inoculation increased the colonization more when P was not added, but did not increase the shoot dry weight at either P level. The rhizosphere pH was lower in the inoculated plants compared to the noninoculated plants in the absence of added P, while the Eh did not change. The decrease in pH in the rhizosphere of inoculated plants could be responsible for the increased P and Zn uptake observed with inoculation. In contrast, Mn uptake was decreased by inoculation. The resin-adsorbed P was increased by inoculation, which, along with the bioavailability index data, may indicate that mycorrhizal roots were able to increase the solubility of soil P.     

Studies on the diversity of arbuscular mycorrhizal fungi and the efficacy of two native isolates in a highly alkaline anthropogenic sediment

A field survey of the arbuscular mycorrhizal status of herbaceous plant species was conducted in a highly alkaline anthropogenic sediment resulting from the disposal of waste from an acetylene and polyvinyl chloride factory. Most plant species found at the site were mycorrhizal and the dominant mycotrophic plant species was Conyza bilbaoana. Fungal species richness was assessed by identification of spores extracted from the sediment and from continuously propagated trap pot cultures. All of the six species of arbuscular mycorrhizal fungi (AMF) found were from the genus Glomus. Glomus intraradices and G. mosseae were found in field-collected sediment samples and also occurred most frequently in trap cultures. To test the symbiotic effectiveness of these two fungi, seedlings of C. bilbaoana were inoculated with either native G. intraradices BEG163 or G. mosseae BEG198 and non-native G. intraradices BEG75 or G. mosseae BEG25 isolates in sterile and non-sterile sediment collected from the study site. All four isolates were able to colonise C. bilbaoana. However, AMF native to the target sediments were generally more effective than the non-native fungi in promoting plant establishment and growth under highly alkaline conditions. The non-native G. intraradices was, however, more effective than the non-native G. mosseae. The results of this study suggest the use of adapted AMF as inoculants for phytorestoration of alkaline anthropogenic-stressed sediments.     

Suppression of the Biocontrol Agent Trichoderma harzianum by Mycelium of the Arbuscular Mycorrhizal Fungus Glomus intraradices in Root-Free Soil

Trichoderma harzianum is an effective biocontrol agent against several fungal soilborne plant pathogens. However, possible adverse effects of this fungus on arbuscular mycorrhizal fungi might be a drawback in its use in plant protection. The objective of the present work was to examine the interaction between Glomus intraradices and T. harzianum in soil. The use of a compartmented growth system with root-free soil compartments enabled us to study fungal interactions without the interfering effects of roots. Growth of the fungi was monitored by measuring hyphal length and population densities, while specific fatty acid signatures were used as indicators of living fungal biomass. Hyphal ³³P transport and β-glucuronidase (GUS) activity were used to monitor activity of G. intraradices and a GUS-transformed strain of T. harzianum, respectively. As growth and metabolism of T. harzianum are requirements for antagonism, the impact of wheat bran, added as an organic nutrient source for T. harzianum, was investigated. The presence of T. harzianum in root-free soil reduced root colonization by G. intraradices. The external hyphal length density of G. intraradices was reduced by the presence of T. harzianum in combination with wheat bran, but the living hyphal biomass, measured as the content of a membrane fatty acid, was not reduced. Hyphal ³³P transport by G. intraradices also was not affected by T. harzianum. This suggests that T. harzianum exploited the dead mycelium but not the living biomass of G. intraradices. The presence of external mycelium of G. intraradices suppressed T. harzianum population development and GUS activity. Stimulation of the hyphal biomass of G. intraradices by organic amendment suggests that nutrient competition is a likely means of interaction. In conclusion, it seemed that growth of and phosphorus uptake by the external mycelium of G. intraradices were not affected by the antagonistic fungus T. harzianum; in contrast, T. harzianum was adversely affected by G. intraradices.     

Mycorrhizal symbiosis and phosphorus fertilization effects on Zea mays growth and heavy metals uptake

Arbuscular mycorrhizal fungi (AMF) can promote plant growth and reduce plant uptake of heavy metals. Phosphorus (P) fertilization can affect this relationship. We investigated maize (Zea mays L.) uptake of heavy metals after soil AMF inoculation and P fertilization. Maize biomass, glomaline and chlorophyll contents and uptake of Fe, Mn, Zn, Cu, Cd and Pb have been determined in a soil inoculated with AMF (Glomus aggregatum, or Glomus intraradices) and treated with 30 or 60 µg P-K₂HPO₄ g^?1 soil. Consistent variations were found between the two mycorrhizal species with respect to the colonization and glomalin content. Shoot dry weight and chlorophyll content were higher with G. intraradices than with G. aggregatum inoculation. The biomass was highest with 30 µg P g^?1 soil. Shoot concentrations of Cd, Pb and Zn decreased with G. aggregatum inoculation, but that of Cd and Pb increased with G. intraradices inoculation. Addition of P fertilizers decreased Cd and Zn concentrations in the shoot. AMF with P fertilization greatly reduced maize content of heavy metals. The results provide that native AMF with a moderate application rate of P fertilizers can be exploited in polluted soils to minimize the heavy metals uptake and to increase maize growth.     

The bacterium Paenibacillus validus stimulates growth of the arbuscular mycorrhizal fungus Glomus intraradices up to the formation of fertile spores

Two isolates of Paenibacillus validus (DSM ID617 and ID618) stimulated growth of the arbuscular mycorrhizal fungus Glomus intraradices Sy167 up to the formation of fertile spores, which recolonize carrot roots. Thus, the fungus was capable of completing its life cycle in the absence of plant roots, but relied instead on the simultaneous growth of bacteria. The supernatant of a mixed batch culture of the two P. validus isolates contained raffinose and another, unidentified trisaccharide. Among the oligosaccharides tested, raffinose was most effective in stimulating hyphal mass formation on plates but could not promote growth to produce fertile spores. A suppressive subtractive hybridization library followed by reverse Northern analyses indicated that several genes with products involved in signal transduction are differentially expressed in G. intraradices SY 167 when grown in coculture with P. validus (DSM 3037). The present investigation, while likely representing a significant step forward in understanding the arbuscular mycorrhizal fungus symbioses, also confirms that its optimal establishing and functioning might rely on many, as yet unidentified factors.