Arbuscular mycorrhizal fungi enhance aluminium resistance of broomsedge (Andropogon virginicus L.)

In the eastern United States, broomsedge (Andropogon virginicus L.) is found growing on abandoned coal-mined lands that have extremely acidic soils with high residual aluminium (Al) concentrations. Broomsedge may be inherently metal-resistant and nutrient-efficient or may rely on the arbuscular mycorrhizal (AM) fungal association to overcome limitations on such sites. Broomsedge plants were grown with and without an acidic ecotype AM fungal consortium and exposed to controlled levels of Al in two experiments. The AM fungal consortium conferred Al resistance to broomsedge. Arbuscular mycorrhizal fungi reduced Al uptake and translocation in host plants, potentially reflecting measured reductions in inorganic Al availability in the rhizosphere of mycorrhizal plants. Mycorrhizal plants exhibited lower shoot P concentrations, higher phosphorus use efficiency, and lower root acid phosphatase rates than non-mycorrhizal plants. Aluminium significantly reduced calcium (Ca) and magnesium (Mg) tissue concentrations in both mycorrhizal and non-mycorrhizal plants. However, plant response to any change in nutrient acquisition was substantially less pronounced in mycorrhizal plants. The exclusion of Al and greater stability of tissue biomass accretion-tissue nutrient relationships in mycorrhizal broomsedge plants exposed to Al may be important mechanisms that allow broomsedge to grow on unfavourable acidic soils.     

Arbuscular mycorrhizal fungi alter phosphorus relations of broomsedge (Andropogon virginicus L.) plants

Broomsedge (Andropogon virginicus L.) is a dominant grass revegetating many abandoned coal-mined lands in West Virginia, USA. Residual soils on such sites are often characterized by low pH, low nutrients, and high aluminium. Experiments were conducted to assess the resistance of broomsedge to limited phosphorus (Pi) availability and to investigate the role that arbuscular mycorrhizal (AM) fungi play in aiding plant growth under low Pi conditions. Pregerminated mycorrhizal and non-mycorrhizal seedlings were grown in a sand-culture system with nutrient solutions containing Pi concentrations ranging from 10 to 100 microM for 8 weeks. Non-mycorrhizal plants exhibited severe inhibition of growth under Pi limitation (<60 microM). Colonization by AM fungi (combined Glomus clarum Nicolson & Schenck and Gigaspora gigantea (Nicol. & Gerd.) Gerd. & Trappe) greatly enhanced host plant growth at low Pi concentrations, but did not benefit growth when Pi was readily available (100 microM). In comparison to non-mycorrhizal plants, mycorrhizal plants had higher phosphorus use efficiency at low Pi concentrations and maintained nearly constant tissue nutrient concentrations across the gradient of Pi concentrations investigated. Manganese (Mn) and sodium (Na) accumulated in shoots of non-mycorrhizal plants under Pi limitation. Mycorrhizal plants exhibited lower instantaneous Pi uptake rates and significantly lower C(min) values compared to non-mycorrhizal plants. These patterns suggest that the symbiotic association between broomsedge roots and AM fungi effectively maintains nutrient homeostasis through changes in physiological properties, including nutrient uptake, allocation and use. The mycorrhizal association is thus a major adaptation that allows broomsedge to become established on infertile mined lands.     

A comparison of nine species of arbuscular mycorrhizal fungi on the development and nutrition of micropropagated Rubus idaeus L. cv. Glen Prosen (Red Raspberry)

Growth, development and nutrient status of micropropagated Rubus idaeus cv. Glen Prosen in response to inoculation with nine species of arbuscular mycorrhizal (AM) fungi from three different genera was investigated. The nine species of AM fungi included, Glomus clarum, G. etunicatum, G. intraradices, Gigaspora rosea, Gi. gigantea, Gi. margarita, Scutellospora calospora, S. heterogama and S. persica. Plant responses to AM fungi varied from growth enhancement to growth depression. Depressive growth effects were specific to Gigaspora species. Furthermore, particular species of AM fungi had unique effects on the mineral status of the raspberry plants. Importance of isolate selection for inoculation of micropropagated raspberry plants is discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of mycorrhizal fungus isolates on mineral acquisition by Panicum virgatum in acidic soil

 Plant ability to withstand acidic soil mineral deficiencies and toxicities can be enhanced by root-arbuscular mycorrhizal fungus (AMF) symbioses. The AMF benefits to plants may be attributed to enhanced plant acquisition of mineral nutrients essential to plant growth and restricted acquisition of toxic elements. Switchgrass (Panicum virgatum L.) was grown in pH_Ca (soil:10 mM CaCl₂, 1 : 1) 4 and 5 soil (Typic Hapludult) inoculated with Glomus clarum, G. diaphanum, G. etunicatum, G. intraradices, Gigaspora albida, Gi. margarita, Gi. rosea, and Acaulospora morrowiae to determine differences among AMF isolates for mineral acquisition. Shoots of mycorrhizal (AM) plants had 6.2-fold P concentration differences when grown in pH_Ca 4 soil and 2.9-fold in pH_Ca 5 soil. Acquisition trends for the other mineral nutrients essential for plant growth were similar for AM plants grown in pH_Ca 4 and 5 soil, and differences among AMF isolates were generally higher for plants grown in pH_Ca 4 than in pH_Ca 5 soil. Both declines and increases in shoot concentrations of N, S, K, Ca, Mg, Zn, Cu, and Mn relative to nonmycorrhizal (nonAM) plants were noted for many AM plants. Differences among AM plants for N and Mg concentrations were relatively small (<2-fold) and were large (2- to 9-fold) for the other minerals. Shoot concentrations of mineral nutrients did not relate well to dry matter produced or to percentage root colonization. Except for Mn and one AMF isolate, shoot concentrations of Mn, Fe, B, and Al in AM plants were lower than in nonAM plants, and differences among AM plants for these minerals ranged from a low of 1.8-fold for Fe to as high as 6.9-fold for Mn. Some AMF isolates were effective in overcoming acidic soil mineral deficiency and toxicity problems that commonly occur with plants grown in acidic soil. Accepted: 14 June 1999     

Field response of wheat to arbuscular mycorrhizal fungi and drought stress

Mycorrhizal plants often have greater tolerance to drought than nonmycorrhizal plants. This study was conducted to determine the effects of arbuscular mycorrhizal (AM) fungi inoculation on growth, grain yield and mineral acquisition of two winter wheat (Triticum aestivum L.) cultivars grown in the field under well-watered and water-stressed conditions. Wheat seeds were planted in furrows after treatment with or without the AM fungi Glomus mosseae or G. etunicatum. Roots were sampled at four growth stages (leaf, tillering, heading and grain-filling) to quantify AM fungi. There was negligible AM fungi colonization during winter months following seeding (leaf sampling in February), when soil temperature was low. During the spring, AM fungi colonization increased gradually. Mycorrhizal colonization was higher in well-watered plants colonized with AM fungi isolates than water-stressed plants. Plants inoculated with G. etunicatum generally had higher colonization than plants colonized with G. mosseae under both soil moisture conditions. Biomass and grain yields were higher in mycorrhizal than nonmycorrhizal plots irrespective of soil moisture, and G. etunicatum inoculated plants generally had higher biomass and grain yields than those colonized by G. mosseae under either soil moisture condition. The mycorrhizal plants had higher shoot P and Fe concentrations than nonmycorrhizal plants at all samplings regardless of soil moisture conditions. The improved growth, yield and nutrient uptake in wheat plants reported here demonstrate the potential of mycorrhizal inoculation to reduce the effects of drought stress on wheat grown under field conditions in semiarid areas of the world.     

The effect of soil phosphorus on the external mycelium growth of arbuscular mycorrhizal fungi during the early stages of mycorrhiza formation

The effects of soil P amendments and time of application on the formation of external mycelium by different arbuscular mycorrhizal (AM) fungi were studied. In the first experiment the external mycelium produced in the soil by the AM fungus Glomus etunicatum Beck. and Gerd., during the early stages of root colonization (7 and 14 days after inoculation), was quantified by the soil-agar film technique. A Brazilian Oxisol was used with three different phosphate levels, varying from deficient to supra-optimal for the plant. Significant differences were observed in the phosphate and inoculation treatments for plant dry weight, P content in the tissue, root length and root colonization, at fourteen days after planting. At 7 days, mycelium growth, root colonization and their relationship were reduced at supra-optimal P concentrations. Applications of P one week after planting reduced mycelium growth and root colonization more than when applied to the soil before planting. In a second experiment the arbuscular mycorrhizal (AM) fungi, Scutellospora heterogama (Nicol. and Gerd.) Walker and Sanders and E3 were tested and compared with Glomus etunicatum. For the species studied, the length of external hyphae per unit of colonized root length was affected by small P additions but no further significant differences were observed at high P levels. The three AM endophytes showed marked differences in their response to P in the soil: Scutellospora heterogama, although producing external mycelium more profusely than the Glomus spp., showed a higher sensitivity to soil P supply.     

The role of arbuscular mycorrhizas in decreasing aluminium phytotoxicity in acidic soils: a review

Soil acidity is an impediment to agricultural production on a significant portion of arable land worldwide. Low productivity of these soils is mainly due to nutrient limitation and the presence of high levels of aluminium (Al), which causes deleterious effects on plant physiology and growth. In response to acidic soil stress, plants have evolved various mechanisms to tolerate high concentrations of Al in the soil solution. These strategies for Al detoxification include mechanisms that reduce the activity of Al³⁺ and its toxicity, either externally through exudation of Al-chelating compounds such as organic acids into the rhizosphere or internally through the accumulation of Al–organic acid complexes sequestered within plant cells. Additionally, root colonization by symbiotic arbuscular mycorrhizal (AM) fungi increases plant resistance to acidity and phytotoxic levels of Al in the soil environment. In this review, the role of the AM symbiosis in increasing the Al resistance of plants in natural and agricultural ecosystems under phytotoxic conditions of Al is discussed. Mechanisms of Al resistance induced by AM fungi in host plants and variation in resistance among AM fungi that contribute to detoxifying Al in the rhizosphere environment are considered with respect to altering Al bioavailability.     

Arbuscular mycorrhizal adaptation,spore germination,root colonization,and host plant growth and mineral acquisition at low pH

Arbuscular mycorrhizal (AM) fungi colonize plant roots and often enhance host plant growth and mineral acquisition, particularly for plants grown under low nutrient and mineral stress conditions. Information about AM fungi and mycorrhizal ( +AM) host plant responses at low pH ( < 5) is limited. Acaulospora are widely reported in acid soil, and Gigaspora sp. appear to be more common in acid soils than Glomus sp. Spores of some AM fungi are more tolerant to acid conditions and high Al than others; t Acaulospora sp., Gigaspora sp., and Glomus manihotis are particularly tolerant. Root colonization is generally less in low than in high pH soils. Percentage root colonization is generally not related to dry matter (DM) produced. Maximum enhancement of plant growth in acid soil varies with AM fungal isolate and soil pH, indicating adaptation of AM isolates to edaphic conditions. Acquisition of many mineral nutrients other than P and Zn is enhanced by +AM plants in acid soil, and the minerals whose concentration is enhanced are those commonly deficient in acid soils (Ca, Mg, and K). Some AM fungal isolates are effective in overcoming soil acidity factors, especially Al toxicity, that restrict plant growth at low pH.     

Diversity of Arbuscular Mycorrhizal Fungi Associated with Plants Growing in Fly Ash Pond and Their Potential Role in Ecological Restoration

Root colonization and diversity of arbuscular mycorrhizal fungi (AMF) were analyzed in plants growing in fly ash pond. Eight species could be separated morphologically, while phylogenetic analyses after PCR amplification of the ITS region followed by RFLP and sequencing revealed seven different AM fungal sequence types. Phylogenetic analysis showed that these sequences cluster into four discrete groups, belonging to the genus Glomus and Archaeospora. Inoculation of plants with spores of AM fungal consortia (Glomus etunicatum, Glomus heterogama, Glomus maculosum, Glomus magnicaule, Glomus multicaule, Glomus rosea, Scutellospora heterogama, and Scutellospora nigra) along with colonized root pieces increased the growth (84.9%), chlorophyll (54%), and total P content (44.3%) of Eucalyptus tereticornis seedlings grown on fly ash compared to non-inoculated seedlings. The growth improvement was the consequence of increased P nutrition and decreased Al, Fe, Zn, and Cu accumulations. These observations suggested that the inoculation of tree seedlings with stress adapted AM fungi aid in the reclamation of fly ash ponds.     

Effect of the saprophytic fungus Fusarium oxysporum on arbuscular mycorrhizal colonization and growth of plants in greenhouse and field trials

Effects of the saprophytic fungus Fusarium oxysporum on arbuscular mycorrhizal (AM) colonization and plant dry matter were studied in greenhouse and field experiments. Host plants: maize (Zea mays L.), sorghum (Sorghum vulgare L.), lettuce (Lactuca sativa L.), tomato (Lycopersicum esculentum L.), wheat (Triticum vulgare L), lentil (Ervum lens L.) and pea (Pisum sativum L.), the AM fungi: Glomus mosseae, G. fasciculatum, G. intraradices, G. clarum, and G. deserticola and the carriers for F. oxysporum inoculum: aqueous solution, thin agar slices, and pellets of agar and alginate were tested under greenhouse conditions. Greatest plant growth and AM colonization responses in sterilized and unsterilized soils were observed with pea, Glomus deserticola and sodium alginate pellets as the carrier for F. oxysporum inoculum. Under field conditions, adding F. oxysporum increased the survival of transplanted pea, possibly through a beneficial effect on AM fungi. Application of F. oxysporum increased shoot dry matter, N and P concentrations of pea and sorghum plants, and the level of AM colonization attained by indigenous or introduced AM fungi. These parameters were similar in plants inoculated with either G. deserticola or with the indigenous AM fungi. Application of the saprophytic fungus increased the number of propagules of AM fungi in field plots in which pea was grown, but this increase was not sufficient to increase AM colonization of sorghum after the pea crop. This revised version was published online in June 2006 with corrections to the Cover Date.     

Arbuscular mycorrhizal fungi increased early growth of two nontimber forest product species Dyera polyphylla and Aquilaria filaria under greenhouse conditions

Nontimber forest products (NTFPs) represent an important source of income to millions of people in tropical forest regions, but some NTFP species have decreased in number and become endangered due to overexploitation. There is increasing concern that the planting stocks of Dyera polyphylla and Aquilaria filaria are not sufficient to sustain the yield of NTFPs and promote forest conservation. The objective of this study was to determine the effect of two arbuscular mycorrhizal (AM) fungi, Glomus clarum and Gigaspora decipiens, on the early growth of two NTFP species, D. polyphylla and A. filaria, under greenhouse conditions. The seedlings of both species were inoculated with G. clarum or G. decipiens, or uninoculated (control) under greenhouse conditions. Percentage of AM colonization, plant growth, survival rate, and nitrogen (N) and phosphorus (P) concentrations were measured after 180 days of growth. The percentage of AM colonization of D. polyphylla and A. filaria ranged from 87 to 93% and from 22 to 39%, respectively. Colonization by G. clarum and G. decipiens increased plant height, diameter, and shoot and root dry weights. Shoot N and P concentrations of the seedlings were increased by AM colonization by as much as 70–153% and 135–360%, respectively. Survival rates were higher in the AM-colonized seedlings at 180 days after transplantation than in the control seedlings. The results suggest that AM fungi can accelerate the establishment of the planting stocks of D. polyphylla and A. filaria, thereby promoting their conservation ecologically and sustaining the production of these NTFPs economically.     

Effectiveness of mycorrhizal fungi on globe artichoke (Cynara cardunculus L. var. scolymus) micropropagation

The effectiveness of two arbuscular mycorrhizal (AM) fungal isolates (Glomus intraradices and Glomus viscosum) in sustaining plant growth and the physiological activities of the micropropagated globe artichoke (Cynara cardunculus L. var. scolymus (L.) Fiori) were investigated during acclimatization and 90 days after plant establishment. All the mycorrhizal microplants survived transplant shock thus confirming the positive role of AM fungi colonization on ex vitro establishment. The growth increased in mycorrhizal plants, especially in plants inoculated with Glomus viscosum. Mycorrhizal plantlets showed higher stomatal conductance, which is probably necessary to supply the carbon needs of fungal symbionts. The SPAD (soil plant analysis development) data could be useful for plant management as a predictor for tissue nitrogen levels. The higher SPAD values in mycorrhizal plants are strictly related to a higher photosynthetic potential, and consequently to their better nitrogen nutrient status due to the symbiotic relationship. Regardless of the mycorrhizal performance in the host–fungus combination, the most efficient fungus for the artichoke microplants was Glomus viscosum.     

Exogenous systemin has a contrasting effect on disease resistance in mycorrhizal tomato (<Emphasis Type="Italic">Solanum lycopersicum</Emphasis>) plants infected with necrotrophic or hemibiotrophic pathogens

A study was performed to determine the effect of the systemin polypeptide on the bio-protective effect of arbuscular mycorrhizal fungi (AMF) in tomato plants infected with Alternaria solani, Phytophthora infestans or P. parasitica. Before infection, tomato plants were colonized with two different AMF, Glomus fasciculatum or G. clarum. In addition, a group of inoculated plants was treated with systemin, just after emergence. The exogenous application of systemin marginally suppressed the resistance against A. solani leaf blight observed in G. fasciculatum mycorrhizal plants but significantly enhanced it in plants colonized with G. clarum. Systemin induced resistance to P. parasitica in leaves of G. fasciculatum mycorrhizal plants, in which AMF colonization alone was shown to have no protective effect. Conversely, none of the treatments led to resistance to root or stem rots caused by P. infestans or P. parasitica. The above effects did not correlate with changes in the activity levels of β-1,3-glucanase (BG), chitinase (CHI), peroxidase (PRX), and phenylalanine ammonium lyase (PAL) in leaves of infected plants. However, they corroborated previous reports showing that colonization by AMF can lead to a systemic resistance response against A. solani. Systemic resistance to A. solani was similarly observed in non-mycorrhizal systemin-treated plants, which, in contrast, showed increased susceptibility to P. infestans and P. parasitica. The results indicated that the pattern of systemic disease resistance conferred by mycorrhizal colonization was dependent on the AMF employed and could be altered by the exogenous application of systemin, by means of a still undefined mechanism.     

Ecological roles of arbuscular mycorrhizal fungi in two wild legume plants

Two wild legume plants,Glycine soja andCassia mimosoides var.nomame, and a cultivated plant, soybean (Glycine max), were employed for a study of triple symbiosis with an inoculum ofScutellispora heterogama harvested from natural soils and an inoculum of their own rhizobial cells. The dry weight, colonization of arbuscular mycorrhizal fungus, nodule formation and N₂-fixation activity were estimated as the parameters of triple symbiosis. The two wild legume plants showed greater growth with colonization of arbuscular mycorrhizae than with nodulation, whereas the cultivated legume showed more nodulation than colonization of arbuscular mycorrhizae. Moreover,S. heterogama appeared to stimulate the triple symbiosis for the wild legume plants. The results suggested that spores ofS. heterogama are important in disturbed soils in Korea.     

Mycorrhizal colonization status of plant species established in an exposed area following the 2000 eruption of Mt. Usu, Hokkaido, Japan

We investigated the ectomycorrhizal (ECM) and arbuscular mycorrhizal (AM) colonization status of plant seedlings that established in areas devastated by the eruption in March 2000 of Mt. Usu, Hokkaido, Japan. In 2005, we estimated the mycorrhizal colonization ratios and frequencies in seedlings of twelve herbaceous and seven woody plant species. Although arbuscular mycorrhizas were found to colonize Equisetum arvense and Polygonum sachalinense, they were presented at very low frequencies and colonization ratios. Other herbaceous plants exhibited higher frequencies of AM associations and either AM and/or ECM fungal associations were observed in all of the woody plant seedlings investigated. The dominant woody plant species (Populus maximowiczii, Salix sachalinensis and Salix hultenii var. angustifolia) associated mainly with ECM fungi and exhibited variable associations with AM fungi. Mycorrhizal associations were common and significant events for plant establishment, even in the early stages of the revegetation process.     

Arbuscular mycorrhizal fungi enhance root cadmium and copper accumulation in the roots of the salt marsh plant Aster tripolium L.

It is known that vegetation plays an important role in the retention of heavy metals in salt marshes by taking up and accumulating the metals. In this study, we investigated whether arbuscular mycorrhizal fungi (AMF) increase Cd and Cu uptake and accumulation in the root system of the salt marsh species Aster tripolium L., and whether indigenous AMF isolated from polluted salt marshes have higher capacity to resist and alleviate metal stress in A. tripolium than isolates of the same species originated from non-polluted sites. Plants inoculated with Glomus geosporum, either isolated from a polluted salt marsh site (PL isolate) or from a non-polluted site (NP isolate), and non-mycorrhizal (NM) plants were compared in a pot experiment at four different Cd and Cu concentrations. Cd had no effect in root colonization, whereas high concentrations of Cu decreased colonization level in plants inoculated with the NP isolate. AM colonization did not increase plant dry weight or P concentration but influenced root Cd and Cu concentrations. Inoculation with PL and NP isolates enhanced root Cd and Cu concentrations, especially at highest metal addition levels, as compared to NM plants, without increasing shoot Cd and Cu concentrations. There was no evidence of intraspecific variation in the effects between AMF isolated from polluted and non-polluted sites, since there were no differences between plants inoculated with PL or NP isolate in any of the tested plant variables. The results of this study showed that AMF enhance metal accumulation in the root system of A. tripolium, suggesting a contribution of AMF to the sink of metals within vegetation in the salt marshes.     

Effect of two AMF life strategies on the tripartite symbiosis with Bradyrhizobium japonicum and soybean

This study is the first in assessing the effect of soil disturbance on the contribution of arbuscular mycorrhizal fungi (AMF) with different life-history strategies to the tripartite symbiosis with soybeans and Bradyrhizobium japonicum (Kirchner) Jordan. We hypothesized that Gigaspora margarita Becker and Hall would be more affected by soil disturbance than Glomus clarum Nicol. and Schenck, and consequently, the tripartite symbiosis would develop more rapidly and lead to greater N₂ fixation in the presence of the latter. Soil pasteurization allowed the establishment of treatments with individual AMF species and soil disturbance enabled the development of contrasting root colonization potentials. In contrast, the colonization potential of B. japonicum was kept the same in all treatments. Soil disturbance significantly reduced root colonization by both AMF, with Gi. margarita being considerably more affected than G. clarum. Furthermore, the tripartite symbiosis progressed faster with G. clarum, and at 10 days after plant emergence, there was 30% more nodules when G. clarum was present compared to that when the bacterial symbiont alone was present. At flowering, the absence of soil disturbance stimulated N₂ fixation by 17% in mycorrhizal plants. However, this response was similar for both AMF.     

Availability and function of arbuscular mycorrhizal and ectomycorrhizal fungi during revegetation of dewatered reservoirs left after dam removal

Revegetation following dam removal projects may depend on recovery of arbuscular mycorrhizal (AM) and ectomycorrhizal (EM) fungal communities, which perform valuable ecosystem functions. This study assessed the availability and function of AM and EM fungi for plants colonizing dewatered reservoirs following a dam removal project on the Elwha River, Olympic Peninsula, Washington, United States. Availability was assessed via AM fungal spore density in soils and EM root tip colonization of Salix sitchensis (Sitka willow) in an observational field study. The effect of mycorrhizal fungi from 4 sources (reservoir soils, commercial inoculum, and 2 mature plant community soils) on growth and nutrient status of S. sitchensis was quantified in a greenhouse study. AM fungal spores and EM root tips were present in all field samples. In the greenhouse, plants receiving reservoir soil inoculum had only incipient mantle formation, while plants receiving inoculum from mature plant communities had fully formed EM root tips. EM formation corresponded with alleviation of phosphorus stress in plants (lower shoot nitrogen:phosphorus). Thus, revegetating plants have access to AM and EM fungi following dam removal, and EM formation may be especially important for plant P uptake in reservoir soils. However, availability of mycorrhizal fungi declines with distance from established plant communities. Furthermore, EM fungal communities in recently dewatered reservoirs may not be as effective at forming beneficial mycorrhizae as those from mature plant communities. Whole soil inoculum from mature plant communities may be important for the success of revegetating plants and recovery of mycorrhizal fungal communities.     

Effects of arbuscular mycorrhiza (AM) on health ofLinum usitatissimum L. infected by fungal pathogens

Effects of arbuscular mycorrhizal (AM) symbiosis on health ofLinum usitatissimum infected by fungal pathogens were investigated exemplarily. Physiological and biochemical analyses were done to explain the mechanisms underlying the AM effects. AM plants showed increased resistance against the wilt pathogen (Fusarium oxysporum f. sp.lini), the level of this effects depended on the plant cultivars which all showed the same level of root colonization by arbuscular mycorrhizal fungi (AMF). In contrary to that, AM plants were highly susceptible against the shoot pathogenOidium lini, but they suffered less than non-AM plants in terms of shoot fresh weight, CO₂ assimilation and content of sucrose in shoot apex. This indicates that AM not only activates resistance mechanisms but also can induce tolerance against pathogens. The concentration of phytohormones such as auxin- and gibberellin-like substances were increased in shoots of AM plants. In roots the ethylene production was increased, too. Furthermore the content and composition of free sterols were highly altered in leaves of AM plants. Root infection by AMF caused an increased respiratory activity and a reduced degree of DNA methylation, but both modifications only occurred in infected root parts indicating an increasing gene activity. The presented results suggest that nearly all parts of a plant are influenced by AM but not in the same manner. In the case of mildewed linseed the effect of AM on plant health was impressing, it indicates that AM has an ability to induce tolerance.     

Arbuscular mycorrhizal contribution to heavy metal uptake by maize (Zea mays L.) in pot culture with contaminated soil

In two pot-culture experiments with maize in a silty loam (P2 soil) contaminated by atmospheric deposition from a metal smelter, root colonization with indigenous or introduced arbuscular mycorrhizal (AM) fungi and their influence on plant metal uptake (Cd, Zn, Cu, Pb, Mn) were investigated. Soil was -irradiated for the nonmycorrhizal control. In experiment 1, nonirradiated soil provided the mycorrhizal treatment, whereas in experiment 2 the irradiated soil was inoculated with spores of a fungal culture from P2 soil or a laboratory reference culture, Glomus mosseae. Light intensity was considerably higher in experiment 2 and resulted in a fourfold higher shoot and tenfold higher root biomass. Under the conditions of experiment 1, biomass was significantly higher and Cd, Cu, Zn and Mn concentrations significantly lower in the mycorrhizal plants than in the nonmycorrhizal plants, suggesting a protection against metal toxicity. In contrast, in experiment 2, biomass did not differ between treatments and only Cu root concentration was decreased with G. mosseae-inoculated plants, whereas Cu shoot concentration was significantly increased with the indigenous P2 fungal culture. The latter achieved a significantly higher root colonization than G. mosseae (31.7 and 19.1%, respectively) suggesting its higher metal tolerance. Zn shoot concentration was higher in both mycorrhizal treatments and Pb concentrations, particularly in the roots, also tended to increase with mycorrhizal colonization. Cd concentrations were not altered between treatments. Cu and Zn, but not Pb and Cd root-shoot translocation increased with mycorrhizal colonization. The results show that the influence of AM on plant metal uptake depends on plant growth conditions, on the fungal partner and on the metal, and cannot be generalized. It is suggested that metal-tolerant mycorrhizal inoculants might be considered for soil reclamation, since under adverse conditions AM may be more important for plant metal resistance. Under the optimized conditions of normal agricultural practice, however, AM colonization even may increase plant metal absorption from polluted soils.