Suppression of other soil microorganisms by mycelium of arbuscular mycorrhizal fungi in root-free soil

The influence of mycelium of two arbuscular mycorrhizal (AM) fungi, Glomus intraradices and Glomus mosseae, on other soil microorganisms, was examined in root-free soil with and without organic substrate amendment in terms of cellulose. The AM fungi were grown in symbiosis with cucumber in a compartmented growth system, which allowed AM fungal external mycelium to grow into root-free compartments. The fungicide Benomyl was applied to the root-free compartments to create an alternative non-mycorrhizal control treatment. Whole cell biomarker fatty acids were employed to quantify different groups of soil microorganisms including the two AM fungi. Abundance of most microbial groups were reduced by external mycelium of both AM fungi, though differential effects on the microbial community composition were observed between the two AM fungi as revealed from principal component analysis. Inhibition of other soil microorganisms was more pronounced in root-free soil with mycelium of G. mosseae than with mycelium of G. intraradices. In general, cellulose increased the amount of biomarker fatty acids of most groups of soil microorganisms, but cellulose did not affect the influence of AM fungi on other soil microorganisms. Benomyl suppressed growth of the external mycelium of the two AM fungi and had limited non-target effects on other microbial groups. In conclusion, our results show differential effects of external mycelium of AM fungi on other soil microbial communities, though both AM fungi included in the study overall inhibited most microbial groups as examined using whole cell biomarker fatty acids.     

EXPLOITATION OF PHOSPHORUS PATCHES WITH DIFFERENT PHOSPHORUS ENRICHMENT BY THREE ARBUSCULAR MYCORRHIZAL FUNGI

The effect of three arbuscular mycorrhizal (AM) fungi on phosphorus (P) nutrient activation and acquisition by maize from spatially heterogeneous sand was investigated using dual-mesh packages enriched with different P concentrations and compared with non-mycorrhizal cotrols. As would be expected the AM fungi significantly enhanced leaf photosynthetic rate and the biomass and P concentrations in shoots and roots. All three fungi (Glomus intraradices, Glomus mosseae and Glomus etunicatum) displayed the capacity to dissolve inorganic P and promoted P nutrient availability in the packages (P patches). G. etunicatum showed the largest effect comparing with Glomus intraradices and Glomus mosseae, particularly in packages with high concentrations of P. Possible mechanisms involved include the acidification of the P patches by the AM fungi, promotion of the dissolution of the P, and more marked effects of the three fungal isolates with increasing enrichment of P in the patches. Inoculation with G. etunicatum resulted in greater acidification compared to the other two fungi. We conclude that AM fungi can promote P availability by acidifying the soil and consequently exploiting the P in nutrient patches and by facilitating the growth and development of the host plants.     

The bacterial community of tomato rhizosphere is modified by inoculation with arbuscular mycorrhizal fungi but unaffected by soil enrichment with mycorrhizal root exudates or inoculation with Phytophthora nicotianae

Arbuscular mycorrhizal (AM) fungi have been shown to induce the biocontrol of soilborne diseases, to change the composition of root exudates and to modify the bacterial community structure of the rhizosphere, leading to the formation of the mycorrhizosphere. Tomato plants were grown in a compartmentalized soil system and were either submitted to direct mycorrhizal colonization or to enrichment of the soil with exudates collected from mycorrhizal tomato plants, with the corresponding negative controls. Three weeks after planting, the plants were inoculated or not with the soilborne pathogen Phytophthora nicotianae growing through a membrane from an adjacent infected compartment. At harvest, a PCR-Denaturing gradient gel electrophoresis analysis of 16S rRNA gene fragments amplified from the total DNA extracted from each plant rhizosphere was performed. Root colonization with the AM fungi Glomus intraradices or Glomus mosseae induced significant changes in the bacterial community structure of tomato rhizosphere, compared to non-mycorrhizal plants, while enrichment with root exudates collected from mycorrhizal or non-mycorrhizal plants had no effect. Our results support that the effect of AM fungi on rhizosphere bacteria would not be mediated by compounds present in root exudates of mycorrhizal plants but rather by physical or chemical factors associated with the mycelium, volatiles and/or root surface bound substrates. Moreover, infection of mycorrhizal or non-mycorrhizal plants with P. nicotianae did not significantly affect the bacterial community structure suggesting that rhizosphere bacteria would be less sensitive to the pathogen invasion than to mycorrhizal colonization. Of 96 unique sequences detected in the tomato rhizosphere, eight were specific to mycorrhizal fungi, including two Pseudomonas, a Bacillus simplex, an Herbaspirilium and an Acidobacterium. One Verrucomicrobium was common to rhizospheres of mycorrhizal plants and of plants watered with mycorrhizal root exudates.     

Changes in soil aggregation and glomalin-related soil protein content as affected by the arbuscular mycorrhizal fungal species Glomus mosseae and Glomus intraradices

Arbuscular mycorrhizal (AM) fungi are key organisms of the soil/plant system, influencing soil fertility and plant nutrition, and contributing to soil aggregation and soil structure stability by the combined action of extraradical hyphae and of an insoluble, hydrophobic proteinaceous substance named glomalin-related soil protein (GRSP). Since the GRSP extraction procedures have recently revealed problems related to co-extracting substances, the relationship between GRSP and AM fungi still remains to be verified. In this work the hypothesis that GRSP concentration is positively correlated with the occurrence of AM fungi was tested by using Medicago sativa plants inoculated with different isolates of Glomus mosseae and Glomus intraradices in a microcosm experiment. Our results show that (i) mycorrhizal establishment produced an increase in GRSP concentration - compared to initial values - in contrast with non-mycorrhizal plants, which did not produce any change; (ii) aggregate stability, evaluated as mean weight diameter (MWD) of macroaggregates of 1-2 mm diameter, was significantly higher in mycorrhizal soils compared to non-mycorrhizal soil; (iii) GRSP concentration and soil aggregate stability were positively correlated with mycorrhizal root volume and weakly correlated with total root volume; (iv) MWD values of soil aggregates were positively correlated with values of total hyphal length and hyphal density of the AM fungi utilized.The different ability of AM fungal isolates to affect GRSP concentration and to form extensive and dense mycelial networks, which may directly affect soil aggregates stability by hyphal enmeshment of soil particles, suggests the possibility of selecting the most efficient isolates to be utilized for soil quality improvement and land restoration programs.     

Response of nitrogen-transforming microorganisms to arbuscular mycorrhizal fungi

 We studied fluctuations in the numbers of autotrophic ammonium oxidizers, ammonifying microorganisms and denitrifying microorganisms in pot cultures of mycorrhizal and non-mycorrhizal maize. The populations were enumerated after 0, 15, 30, 45 and 60 days of plant growth. Two arbuscular mycorrhizal (AM) fungi belonging to different Glomus species were investigated. Pot cultures with AM-infected maize had significant quantitative and qualitative changes in the root-associated population of N-transforming bacteria compared with the non-mycorrhizal controls. The occurrence of autotrophic ammonium oxidizers in pot cultures of the AM fungi Glomus mosseae and G. fasciculatum was significantly higher than in non-mycorrhizal cultures throughout maize growth. The occurrence of these bacteria was delayed by 15 days in non-mycorrhizal as opposed to Glomus-colonized soil. Ammonifying and denitrifying bacterial populations were significantly decreased in the pot cultures of AM plants compared with the control. The distribution patterns of the physiological groups of bacteria tested were similar for both AM treatments but different from that of the non-mycorrhizal controls. Activity measurements expressed on a per cell basis showed changes with respect to the form of N in the mycorrhizal soil. G. fasciculatum was more active than G. mosseae during the earlier stages of plant growth. Received: 8 July 1997     

Fertilizer Concentration Affects Growth Response and Leaf Color of Tradescantia virginiana L.

ABSTRACT

A pot experiment was conducted out to investigate the yield and pungency of spring onion (Allium fistulosum L.) as affected by inoculation with arbuscular mycorrhizal (AM) fungi and addition of nitrogen (N) and sulfur (S) fertilizers. Plants were inoculated with either Glomus mosseae or Glomus intraradices or grown as uninoculated controls. Two levels of N and S were applied to the soil in factorial combinations of 50 and 250 mg N kg^?1 soil and 0 and 60 mg S kg^?1 soil. Plants were grown in a greenhouse for 25 weeks and then harvested. Mycorrhizal colonization resulted in increased shoot dry weight, shoot-to-root ratio, shoot length, sheath diameter, and phosphorus (P) concentrations. Shoot dry-matter yield was significantly affected by added N, but not by S. Shoot dry weight increased with increasing N supply (except for non-mycorrhizal controls without additional S fertilizer). Shoot total S concentration (TSC), enzyme-produced pyruvate (EPY), and organic sulfur concentration (OSC) in plants inoculated with Glomus mosseae were significantly lower than those of non-mycorrhizal controls, while these parameters in plants inoculated with Glomus intraradices were comparable to or higher than in the controls. Neither N nor S supply affected shoot EPY or OSC, whereas shoot TSC (except in plants inoculated with Glomus mosseae) and SO₄ ^2? concentrations were usually significantly increased by S supply. In soil of high S and low P availability, mycorrhizal colonization had a profound influence on both the yield and the pungency of spring onion.     

Water status and stomatal behaviour of cowpea,Vigna unguiculata (L.) Walp,plants inoculated with two Glomus species at low soil moisture levels

The effects of arbuscular mycorrhizal (AM) fungi on water status and stomatal behaviour of cowpea, Vigna unguiculata (L.) Walp. cv. B89-504, under water-stressed conditions in the greenhouse were studied. The 3 × 2 experimental design included two levels of mycorrhizal colonisation (Glomus mosseae, Glomus versiforme) and non-mycorrhizal control treatment and two soil moisture levels (well-watered pots and pots allowed to dry). Relative water content and leaf water potential values were higher in well-watered mycorrhizal and non-mycorrhizal plants than in water-stressed mycorrhizal and non-mycorrhizal plants. AM species had no significant effect on leaf osmotic potential, stomatal conductance and leaf transpiration in both well watered and water-stressed plants. The values of stomatal conductance and leaf transpiration were high during the vegetative stage and low during the flowering stage. These responses which can be related to the age of the plant suggest that mycorrhizal colonisation did not affect stomatal closure of cowpea plants during water stress. The decrease in plant growth and dry matter production in both mycorrhizal and non-mycorrhizal plants shows that drought resistance in cowpea was unaffected by mycorrhiza in the vegetative phase.     

红三叶草丛枝菌根对有机磷的吸收

The capacities of two arbuscular mycorrhizal (AM) fungi, Glomus mosseae and Glomus versderme, tomineralize added organic P were studied in a sterilized calcareous soil. Mycorrhizal (inoculated with either of the AM fungi) and non-mycorrhizal red clover (Trghlium pmtense L.) plants were grown for eight weeksin pots with upper root, central hyphal and lower soil compartments. The hyphal and soil compartmentsreceived either organic P (as Na-phytate) or inorganic P (as KH₂PO₄) at tbe rate of 50 mg P kg^-1. No P wasadded to the root compartments. Control pots received no added P. Yields were higher in mycorrhizal than innon-mycorrhizal clover. Mycorrhizal inoculation doubled shoot P concentration and more than doubled total P uptake of plaflts in P-amended soil, irrespective of the form of applied P. The mycorrhizal contributionto inorganic P uptake was 80% or 76% in plants inoculated with G. mosseae or G. verefforme, respectively. Corresponding values were 74% and 82% when Na-phytate was applied. In the root compartments of the mycorrhizal treatments, the proportion of root length infected, hyphal length density and phosphatase activity were all higher when organic P was applied than when inorganic P was added.     

The significant contribution of mycorrhizal fungi and earthworms to maize protection and phytoremediation in Cd-polluted soils

Mycorrhizal fungi and earthworms can individually or interactively influence plant growth and heavy metal uptake. The influence of earthworms and arbuscular mycorrhizal (AM) fungi either alone or in combination on maize (Zea mays L.) growth and cadmium (Cd) uptake was investigated in a calcareous soil artificially spiked with Cd. Soils were contaminated with Cd (10 and 20 mg Cd kg⁻¹), inoculated or un-inoculated with the epigeic earthworm Lumbricus rubellus and two AM fungal species (Rhizophagus irregularis and Funneliformis mosseae) for two months of growth under greenhouse conditions. Generally, earthworms alone increased both shoot P uptake and biomass but decreased shoot Cd concentration and root Cd uptake. AM fungi individually often increased total maize P uptake, declined shoot Cd concentration, and consequently produced higher total biomass. However, R. irregularis enhanced shoot Cd uptake at low Cd level and root Cd uptake at high Cd level. In plants inoculated with F. mosseae species, earthworms increased shoot biomass and Cd uptake, decreased root biomass and Cd uptake at all Cd levels, and increased shoot Cd concentration at low Cd level. In plants colonized by R. irregularis species, however, earthworm addition decreased maize biomass only at high Cd level and root Cd concentration and total maize Cd uptake at both Cd levels. Earthworm activity decreased Cd transfer from the soil to maize roots at low Cd level, but this was counterbalanced in the presence of F. mosseae. Mycorrhizal symbiosis significantly reduced the transfer of Cd from roots to shoots, independence of earthworm effect. Overall, it is concluded that L. rubellus and AM fungi, in particular F. mosseae isolate, improved maize tolerance to Cd toxicity both individually and interactively by increasing plant growth and P nutrition, and restricting Cd transfer to the aboveground biomass. Consequently, the single and interactive effects of the two soil organisms might potentially be important not only in protecting maize plants against Cd toxicity, but also in Cd phytostabilization in soils polluted by this highly toxic metal.     

Contribution of arbuscular mycorrhizal fungi to utilization of organic sources of phosphorus by red clover in a calcareous soil

A glasshouse pot experiment investigated the uptake by arbuscular mycorrhizal (AM) fungi associated with red clover of three organic sources of P added to a sterilized calcareous soil of low P availability. Each pot was separated into a central compartment for plant growth and two outer compartments for external mycelium using 30-μm nylon mesh to restrict the roots but allow hyphal penetration. Plants in the central compartments were inoculated with the AM fungus Glomus versiforme and uninoculated controls were included. Plants were harvested on three occasions: 5, 7 and 10 weeks after sowing. Application of each of the three organic P sources (lecithin, RNA and sodium phytate) or inorganic P (KH₂PO₄) at 50 mg P kg⁻¹ to the outer compartments of mycorrhizal and uninoculated pots increased the yield, P concentration and total P uptake of red clover compared with pots to which no P was applied, with no differences among P sources in non-mycorrhizal plants but differences observed in mycorrhizal plants both 7 and 10 weeks after sowing suggesting differences in availability of the four P sources to AM mycelium. The contribution of external mycelium to plant uptake of applied P increased with time. The three organic P sources made smaller contributions to plant P nutrition than KH₂PO₄ at the first and second harvests. At the third harvest, the contribution from KH₂PO₄ was 23%, while those from lecithin, RNA and sodium phytate were 23, 17 and 31%, respectively. This suggests that with the mediation of AM fungi, soil organic P sources can make a contribution to host plant P nutrition comparable to that of soluble orthophosphate.     

Brassinosteroid (BR) and arbuscular mycorrhizal (AM) fungi alleviate salinity in wheat

This study was conducted to assess the impacts of brassinosteroide (BR), arbuscular mycorrhizal (AM) fungi, Glomus mosseae and their interactions on salt stress tolerance in Triticum aestivum L. After foliar spraying of mycorrhizal and non-mycorrhizal plants by 5 µM epibrassinolide, they were subjected to 0 and 150 mM sodium chloride (NaCl) for 2 weeks. The experiment was conducted in a randomized complete block design, replicated 4 times. Our results showed a probable potential of BR and/or AM fungi in improving salt tolerance of plants. Total phenol and proline content increased in BR and/ or AM treatments. AM fungi promoted plant growth, including leaf area, shoot and root dry weights, and lengths under saline condition. Moreover, BR improved growth parameters except root dry weights and lengths. This study indicated that BR and/or AM fungi may contribute to improve salt tolerance of the plant.     

Mycorrhizae,biocides, and biocontrol. 4. Response of a mixed culture of arbuscular mycorrhizal fungi and host plant to three fungicides

The effects of three commonly used fungicides on the colonization and sporulation by a mixture of three arbuscular mycorrhizal (AM) fungi consisting of Glomus etunicatum (Becker & Gerd.), Glomus mosseae (Nicol. & Gerd.) Gerd. & Trappe, and Gigaspora rosea (Nicol. & Schenck) in symbiosis with pea plants and the resulting response of the host-plant were examined. Benomyl, PCNB, and captan were applied as soil drenches at a rate of 20 mg active ingredient kg^-1 soil 2 weeks after transplanting pea seedlings in a silty clay-loam soil containing the mixed inocula of AM fungi (AM plants). Effects of fungicides were compared to untreated plants that were inoculated with fungi (AM control). The effect of mycorrhizal inoculation on plant growth was also examined by including nonmycorrhizal, non-fungicide-treated plants (non-AM control). Fungicides or inoculation with AM fungi had only a small effect on the final shoot weights of pea plants, but had greater effects on root length and seed yield. AM control plants had higher seed yields and lower root lengths than the corresponding non-AM plants, and the fungicide-treated AM plants had intermediate yields and root lengths. Seed N and P contents were likewise highest in AM control plants, lowest in non-AM plants, and intermediate in fungicide-treated AM plants. All three fungicides depressed the proportion (%) of root length colonized by AM fungi, but these differences did not translate to reductions in the total root length that was colonized, since roots were longer in the fungicide-treated AM plants. Pea plants apparently compensated for the reduction in AM-fungal metabolism due to fungicides by increasing root growth. Fungicides affected the population of the three fungi as determined by sporulation at the final harvest. Captan significantly reduced the number, relative abundance, and relative volume of G. rosea spores in the final population relative to the controls. The relative volume of G. etunicatum spores was greater in all the fungicide-treated soils, while G. mosseae relative volumes were only greater in the captan-treated soil. These findings show that fungicides can alter the species composition of an AM-fungal community. The results also show that AM fungi can increase seed yield without enhancing the vegetative shoot growth of host plants.     

Synergistic effect of Glomus mosseae and nitrogen fixing Bacillus subtilis strain Daz26 on artemisinin content in Artemisia annua L.

Artemisia annua L. (Asteraceae) is an important medicinal plant whose secondary metabolite artemisinin is used for the treatment of cerebral malaria. A study was undertaken to determine the effect of arbuscular mycorrhizal (AM) fungi, Glomus mosseae, Glomus aggregatum, Glomus fasciculatum, Glomus intraradices and two free living nitrogen fixing bacteria (NFB) (identified as Bacillus subtilis and Stenotrophomonas spp. through 16S rRNA gene sequence analysis), inoculated alone or in combinations on the biomass, nutrient uptake, and content of artemisinin in A. annua under glass house conditions. Various parameters like plant height, total plant biomass, NPK content, leaf yield, mycorrhizal colonization, bacterial population in rhizosphere, phosphatase activity and artemisinin content were determined and found to vary with different treatments. Among all the treatments, plants inoculated with G. mosseae + B. subtilis performed better than any other treatment or uninoculated control plants. The results of the experiment clearly indicated the compatibility and synergy between G. mosseae and B. subtilis and suggested the use of this microbial consortium in A. annua for enhancing growth, biomass yield, and the content and yield of artemisinin.     

Response of Two Maize Inbred Lines with Contrasting Phosphorus Efficiency and Root Morphology to Mycorrhizal Colonization at Different Soil Phosphorus Supply Levels

ABSTRACT

A pot experiment investigated the response of two maize inbred lines with contrasting root morphology and phosphorus (P) efficiency to inoculation with Glomus mosseae or Glomus etunicatum compared with non-mycorrhizal controls. Soil phosphorus was supplied at rates of 10, 50, and 100 mg P kg ^?1 soil. Root length, specific root length, and specific phosphorus uptake of maize line 178 (P-efficient) were significantly higher than of line Hc (P-inefficient). Percentage of root length colonized showed the opposite trend regardless of soil P supply level. The two maize lines did not differ significantly in growth response to mycorrhizal colonization. Root colonization rate decreased with increasing soil phosphorus supply. The beneficial effect of the two AM fungi on plant growth and P uptake was greatest at low soil P level and the responses were negative at high P supply. Mycorrhizal responsiveness also decreased with increasing P supply and differed between the two mycorrhizal fungal isolates.     

Arbuscular mycorrhizal colonisation increases copper binding capacity of root cell walls of Oryza sativa L. and reduces copper uptake

There is evidence that colonisation by mycorrhizal fungi can protect host plants from toxic concentrations of heavy metals. The mechanism by which protection is provided by the fungus for any particular metal is poorly understood. Rice (Oryza sativa L.) plants were inoculated with Glomus mosseae and grown for 4 weeks to ensure strong colonisation. The plants were then exposed to low to toxic concentrations of copper (Cu) and the uptake and distribution were examined. The effect of mycorrhizal colonisation on the cell wall composition and Cu binding capacity of roots was also investigated. Mycorrhizal plants showed moderate reductions in Cu concentrations in roots but large reductions in shoots. In roots, mycorrhizal plants accumulated more Cu in cell walls but much less in the symplasm compared to non-mycorrhizal plants. The differences in cell wall binding of Cu could be partly explained by changes in the composition of the cell wall. The mechanistic basis for the reduced Cu accumulation and the potential beneficial consequences of mycorrhizal associations on plant growth in Cu toxic soil are discussed.     

Effect of indigenous arbuscular mycorrhizal fungi combined with manure on the change in concentration some mineral elements in cotton (Gossypium hirsutum L.)

A pot experiment was conducted to evaluate the effect of indigenous arbuscular mycorrhizal fungi (AMF) and the synergy of indigenous AMF and sheep manure (SM) on potassium (K), calcium (Ca) and some micronutrient concentrations in cotton plant. Indigenous AMF were a mixture of Glomus viscosum, Glomus mosseae and Glomus intraradices initially isolated from a cotton field. Cotton was grown for 12 weeks and the elements of shoot were determined at three stages of plant growth. Inoculated cotton plants with AMF had higher concentrations of K, Ca, manganese (Mn), iron (Fe), copper (Cu) and zinc (Zn) than non-mycorrhizal plants. Shoot concentrations of these elements increased significantly when SM was added to mycorrhizal plants. Maximum plant micronutrient uptake was found in the treatment of AMF inoculation with SM. Mn, Fe, Cu and Zn uptake increased significantly by 457%, 282%, 272% and 295%, respectively, over control. Indigenous AMF combined with SM resulted in better plant growth and micronutrient uptake.     

Mycorrhizal fungi and earthworms reduce antioxidant enzyme activities in maize and sunflower plants grown in Cd-polluted soils

Changes in plant antioxidant enzymes (AOEs) in response to cadmium (Cd) pollution are an important mechanism for plant growth and tolerance to Cd-induced stress. The main objective of this greenhouse study was to determine the combined influence of earthworm and arbuscular mycorrhiza (AM) fungal inoculation and their interactions with Cd on AOEs and proline accumulation in leaves of two major crops under Cd stress. Maize (Zea mays L.) and sunflower (Helianthus annuus L.) plants were exposed to Cd stress (10 and 20 mg kg⁻¹ soil), inoculated with either earthworm (Lumbricus rubellus L.) or AM fungi (Glomus intraradices and Glomus mosseae species) in a pot experiment for three months. Exposure to Cd decreased shoot dry weights, increased shoot Cd and P concentrations, leaf proline accumulation and the activity of superoxide dismutase (SOD), catalase (CAT), peroxidase (POD) and polyphenol oxidase (PPO) in both mycorrhizal and non-mycorrhizal plants and both in the presence and absence of earthworms. Inoculation of both model plants with earthworms and AM fungi decreased shoot Cd concentrations and the activity of all AOEs, except PPO. Although earthworm activity enhanced the proline content of sunflower in Cd-polluted soils, the proline level of both plants remained unaffected by AM fungi. AM fungi and earthworms may decrease the activity of AOEs through a decline in shoot Cd toxicity and concentration, confirming that plant inoculation with these soil organisms improves maize and sunflower tolerance and protection against Cd toxicity. Generally, the effect of AM fungal inoculation on plant responses to Cd addition was greater than that of earthworm activity. Nonetheless, the interactive effect of AM fungus and earthworm is of minor importance for most of the plant AOEs in Cd-polluted soils.     

Improved zinc tolerance in Eucalyptus globulus inoculated with Glomus deserticola and Trametes versicolor or Coriolopsis rigida

The potential of interactions between saprophytic and arbuscular mycorrhizal (AM) fungi to improve Eucalyptus globulus grown in soil contaminated with Zn were investigated. The presence of 100 mg kg ⁻¹ Zn decreased the shoot and root dry weight of E. globulus colonized with Glomus deserticola less than in plants not colonized with AM. Zn also decreased the extent of root length colonization by AM and the AM fungus metabolic activity, measured as succinate dehydrogenase (SDH) activity of the fungal mycelium inside the E. globulus root. The saprophytic fungi Trametes versicolor and Coriolopsis rigida increased the shoot dry weight and the tolerance of E. globulus to Zn when these plants were AM-colonized. Both saprophytic fungi increased the percentage of AM root length colonization and elevated G. deserticola SDH activity in the presence of all Zn concentrations applied to the soil. In the presence of 500 and 1000 mg kg⁻¹ Zn, there were higher metal concentrations in roots and shoots of AM than in non-AM plants; furthermore, both saprophytic fungi increased Zn uptake by E. globulus colonized by G. deserticola. The higher root to shoot metal ratio observed in mycorrhizal E. globulus plants indicates that G. deserticola enhanced Zn uptake and accumulation in the root system, playing a filtering/sequestering role in the presence of Zn. However, saprophytic fungi did not increase the root to shoot Zn ratio in mycorrhizal E. globulus plants. The effect of the saprophytic fungi on the tolerance and the accumulation of Zn in E. globulus was mediated by its effect on the colonization and metabolic activity of the AM fungi.     

Impact of soil nitrogen concentration on Glomus spp.-Sinorhizobium interactions as affecting growth, nitrate reductase activity and protein content of Medicago sativa

Our objective was to evaluate how increasing levels of N in the medium (0, 4, 8 and 16 mmol N added kg^-1 soil) affect the interaction between Sinorhizobium and arbuscular mycorrhiza (AM) fungi in the tripartite symbiosis with Medicago sativa. Growth response, nutrient acquisition, protein content, and nitrate reductase (NR) activity were measured both in plant shoots and roots. Results showed that N levels in soil did not affect mycorrhizal colonization but they strongly influenced nodulation, particularly of mycorrhizal plants. Mycorrhizal colonization was required for a proper nodulation when no N was applied to soil. In contrast, the addition of 4 mmol N kg^-1 soil reduced nodulation only in mycorrhizal plants and 8 mmol N added kg^-1 soil allowed nodule formation only in non-mycorrhizal plants. Nodulation was totally inhibited in all treatments with the addition of 16 mmol N added kg^-1 soil. N addition enhanced NR activity in all the treatments, while AM colonization increased the proportion of NR allocated to roots. This effect was more pronounced under the lowest N levels in the medium. The two AM fungal species showed different distribution pattern of enzymatic activities in plant tissues indicating specific physiological traits. Protein content as well as the relative proportion of protein in roots were greatly increased after mycorrhizal colonization. Glomus intraradices-colonized plants had the highest protein content in shoot and root. Mycorrhizal effects on growth, N acquisition and biochemical variables cannot be interpreted as an indirect P-mediated effect since P content was lower in mycorrhizal plants than in those which were P fertilized. Mycorrhizal colonization increased the N content in plants irrespective of the N level, but the effectiveness of AM fungi on plant N acquisition depended on the AM fungus involved, G. intraradices being the most effective, particularly at the highest N rate. N₂ fixation, enhanced by AM colonization, contributed to N acquisition when a moderate N quantity was available in the soil. Nevertheless, under a high N amount the nodulating process and/or fixing capacity by Sinorhizobium was reduced in AM plants. In contrast, the AM fungal mycelium from a particular mycorrhizal fungus may continue to contribute efficiently to the N uptake from the soil even at high N levels. These results demonstrate the particular sensitivity of AM fungal species in terms of their growth and/or function to increasing N amounts in the medium. A selection of AM fungi used to address specific environmental conditions, such as N fertilization regimes comparable to those used in agronomic practices, is required for a better use of N applied to soil.     

Differential effects of soil disturbance and plant residue retention on function of arbuscular mycorrhizal (AM) symbiosis are not reflected in colonization of roots or hyphal development in soil

The effects of soil disturbance and residue retention on the functionality of the symbiosis between medic (Medicago truncatula L.) and arbuscular mycorrhizal fungi (AMF) were assessed in a two-stage experiment simulating a crop rotation of wheat (Triticum aestivum L.) followed by medic. Plants were inoculated or not with the AMF, Glomus intraradices and Gigaspora margarita, separately or together. The contribution of the arbuscular mycorrhizal (AM) pathway for P uptake was determined using ³²P-labeled soil in a small hyphal compartment accessible only to hyphae of AMF. In general AM colonization was not affected by soil disturbance or residue application and disturbance did not affect hyphal length densities (HLDs) in soil. At 4 weeks disturbance had a negative effect on growth and phosphorus (P) uptake of plants inoculated with G. margarita, but not G. intraradices. By 7 weeks disturbance reduced growth of plants inoculated with G. margarita or AMF mix and total P uptake in all inoculated plants. With the exception of plants inoculated with G. margarita in disturbed soil at 4 weeks, the AM pathway made a significant contribution to P uptake in all AM plants at both harvests. Inoculation with both AMF together eliminated the negative effects of disturbance on AM P uptake and growth, showing that a fungus insensitive to disturbance can compensate for loss of contribution of a sensitive one. Application of residue increased growth and total P uptake of plants but decreased ³²P in plants inoculated with the AMF mix in disturbed soil, compared with plants receiving no residue. The AMF responded differently to disturbance and G. intraradices, which was insensitive to disturbance, compensated for lack of contribution by the sensitive G. margarita when they were inoculated together. Colonization of roots and HLDs in soil were not good predictors of the outcomes of AM symbioses on plant growth, P uptake or P delivery via the AM pathway.