Effect of onion (Allium cepa) root exudates on the hyphal growth of Gigaspora margarita

 The effect of root exudates from onions differing in P status on spore germination and hyphal growth of arbuscular mycorrhizal fungi was investigated. Onion (Allium cepa) was grown in solution culture at different phosphorus concentrations (0, 0.1, 1.0, 8.0 and 24.0 mg P l^–1) and root exudates were collected. When spores of the arbuscular mycorrhizal fungus, Gigaspora margarita were incubated with these root exudates, spore germination was only slightly affected but hyphal growth was greatly affected, particularly with exudates from P-deficient plants. This suggests that the P nutrition of host plants influences the composition of root exudates and thereby the hyphal growth of arbuscular mycorrhizal fungi. Accepted: 25 June 1995     

Effect of root exudates of mycorrhizal tomato plants on microconidia germination of Fusarium oxysporum f. sp. lycopersici

The effect of root exudates from mycorrhizal and non-mycorrhizal tomato plants on microconidia germination of the tomato pathogen Fusarium oxysporum f. sp. lycopersici was tested. Microconidia germination was enhanced in the presence of root exudates from mycorrhizal tomato plants. Tomato plants were colonised by the arbuscular mycorrhizal fungus Glomus fasciculatum, indicating that alterations of the exudation pattern depended on the degree of root AM colonisation. Testing the exudates from plants with a high and a low P level revealed that the alterations of the root exudates from mycorrhizal plants, resulting in a changed effect on microconidia germination, are not due to an improved P status of mycorrhizal plants.     

Root exudates of mycorrhizal tomato plants exhibit a different effect on microconidia germination of Fusarium oxysporum f. sp. lycopersici than root exudates from non-mycorrhizal tomato plants

The effect of root exudates from mycorrhizal and non-mycorrhizal tomato plants on microconidia germination of the tomato pathogen Fusarium oxysporum f. sp. lycopersici was tested. Microconidia germination was enhanced in the presence of root exudates from mycorrhizal tomato plants. The more tomato plants were colonized by the arbuscular mycorrhizal fungus Glomus mosseae, the more microconidia germination was increased, indicating that alterations of the exudation pattern depended on the degree of root AM colonization. Moreover, alterations of the exudation pattern of mycorrhizal plants are not only local, but also systemic. Testing the exudates from plants with a high and a low P level revealed that the alterations of the root exudates from mycorrhizal plants, resulting in a changed effect on microconidia germination, are not due to an improved P status of mycorrhizal plants.     

Phosphorus amendment inhibits hyphal branching of the VAM fungus Gigaspora margarita directly and indirectly through its effect on root exudation

 The effect of solution phosphorus (P) concentration upon growth of pregerminated spores of the vesicular-arbuscular mycorrhizal fungus Gigaspora margarita was examined in vitro. P at 1 mM significantly inhibited branching of the primary germ tube. The number of branches and the total hyphal length were both significantly inhibited at 10 mM P. In addition, germinated spores exposed to exudates produced by Ri T-DNA-transformed roots of Daucus carota L. grown in the presence of P showed significantly less hyphal branching than those exposed to exudates produced by P-stressed roots. These phenomena could contribute to the observed inhibition of mycorrhiza formation by high P. Accepted: 31 July 1996     

Effect of phosphate and the arbuscular mycorrhizal fungus Glomus intraradices on disease severity of root rot of peas (Pisum sativum) caused by Aphanomyces euteiches

 The effects of inorganic phosphate levels and the presence of arbuscular mycorrhiza on disease severity of Aphanomyces euteiches in pea roots were studied. Disease severity on roots and epicotyl as well as the oospore number within infected root tissue were correlated with the phosphorus (P) level in the growth medium. The arbuscular mycorrhizal fungus Glomus intraradices increased P uptake and the P concentration in the plant but reduced disease development in peas. Polyacrylamide gel electrophoresis followed by densitometry of glucose-6-phosphate dehydrogenase specific to A.euteiches was used to measure the activity of the pathogen in roots. The enzyme activity increased with disease severity and disease incidence, except in plants supplemented with P at the highest level, where a peak in activity was seen 12 days after inoculation with the pathogen, followed by a decrease in activity. The epicotyl of mycorrhizal plants showed a reduction in disease severity although this part of the plants was not mycorrhizal. Thus, an induced systemic factor may be responsible for increased resistance in mycorrhizal plants. Accepted: 21 August 1998     

葡萄糖、根浸出液对丛枝菌根真菌吸收不同外源氮产生精氨酸的影响

以大葱(Allium fistulosum)为宿主植物, 接种丛枝菌根(Arbuscular mycorrhizal, AM)真菌Glomus intraradices, 采用三室隔离盆栽培养系统, 在菌丝室施加浓度为4 mmol/L的不同形态外源氮、1%葡萄糖及根浸出液, 通过测定根外菌丝(Extraradical mycelium, ERM)和菌根中精氨酸的含量, 探究葡萄糖、根浸出液对AM真菌吸收不同形式外源氮产生精氨酸的影响。结果表明, 不同外源氮对ERM中精氨酸含量的影响为尿素>Gln>NH₄NO₃>Arg/Gly>NH₄Cl>KNO₃, 对菌根中精氨酸含量的影响为Arg>Gln>尿素>NH₄NO₃>Gly>NH₄Cl>KNO₃; 施加葡萄糖和根浸出液在不同程度上提高ERM干重和菌丝室孢子数量, 但使ERM和菌根中的精氨酸含量降低。说明AM真菌吸收同化不同外源氮产生精氨酸的能力不同, 葡萄糖和根浸出液降低AM真菌吸收同化外源氮产精氨酸的能力。     

Colonization by Arbuscular Mycorrhizal Fungi of Sorghum Leads to Reduced Germination and Subsequent Attachment and Emergence of Striga hermonthica

Two sorghum cultivars: the Striga-tolerant S-35 and the Striga-sensitive CK60-B were grown with or without arbuscular mycorrhizal (AM) fungi, and with or without phosphorus addition. At 24 and 45 days after sowing (DAS) of sorghum, root exudates were collected and tested for effects on germination of preconditioned Striga hermonthica seeds. Root exudates from AM sorghum plants induced lower germination of S. hermonthica seeds than exudates from non-mycorrhizal sorghum. The magnitude of this effect depended on the cultivar and harvest time. A significantly (88–97%) lower germination of S. hermonthica seeds upon exposure to root exudates from AM S-35 plants was observed at both harvest times whereas for AM inoculated CK60-B plants a significantly (41%) lower germination was observed only at 45 DAS. The number of S. hermonthica seedlings attached to and emerged on both sorghum cultivars were also lower in mycorrhizal than in non-mycorrhizal plants. Again, this reduction was more pronounced with S-35 than with CK60-B plants. There was no effect of phosphorus addition on Striga seed germination, attachment or emergence. We hypothesize that the negative effect of mycorrhizal colonization on Striga germination and on subsequent attachment and emergence is mediated through the production of signaling molecules (strigolactones) for AM fungi and parasitic plants.Key Words: arbuscular mycorrhiza, root exudate, sorghum, striga, strigolactones, germination     

Effects of a vesicular-arbuscular mycorrhizal fungus and other soil microorganisms on growth,mineral nutrient acquisition and root exudation of soil-grown maize plants

Maize (Zea mays L. cv. Alize) plants were grown in a calcareous soil in pots divided by 30-m nylon nets into three compartments, the central one for root growth and the outer ones for hyphal growth. Sterle soil was inoculated with either (1) rhizosphere microorganisms other than vesicular-arbuscular mycorrhizal (VAM) fungi, (2) rhizosphere microorganisms together with a VAM fungus [Glomus mosseae (Nicol. and Gerd.) Gerdemann and Trappel], or (3) with a gamma-irradiated inoculum as control. Plants were grown under controlled-climate conditions and harvested after 3 or 6 weeks. VAM plants had higher shootroot ratios than non-VAM plants. After 6 weeks, the concentrations of P, Zn and Cu in roots and shoots had significantly increased with VAM colonization, whereas Mn concentrations had significantly decreased. Root exudates were collected on agar sheets placed on the interface between root and hyphal compartments. Six-week-old VAM and non-VAM plants had similar root exudate compositions of 72–73% reducing sugars, 17–18% phenolics, 7% organic acids and 3% amino acids. In another experiment in which root exudates were collected on agar sheets with or without antibiotics, the amounts of amino acids and carbohydrates recovered were similar in VAM and non-VAM plants. However, threeto sixfold higher amounts of carbohydrates, amino acids and phenolics were recovered when antibiotics were added to the agar sheets. Thus, the high microbial activity in the rhizosphere and on the rhizoplane limits the exudates recovered from roots.     

Systemic inhibition of arbuscular mycorrhiza development by root exudates of cucumber plants colonized by<Emphasis Type="Italic"> Glomus mosseae</Emphasis>

The arbuscular mycorrhizal (AM) non-host plants mustard, sugar beet, lupin and the AM host plant cucumber were used as test plants. Cucumber plants were grown either in the absence of the AM fungus (AMF) Glomus mosseae or in a split-root system, with one side mycorrhizal and one side non-mycorrhizal. Root exudates of the AM non-host plants, the non-mycorrhizal cucumber plants and the mycorrhizal and the non-mycorrhizal side of the split-root system of mycorrhizal cucumber plants were collected and applied to cucumber plants inoculated with the AMF. Root exudates of non-mycorrhizal cucumber plants showed a significant stimulatory effect on root colonization, whereas root exudates from the mycorrhizal and the non-mycorrhizal sides of a split-root system of a mycorrhizal cucumber plant did not show this stimulatory effect and were even slightly inhibitory. Root exudates of the two AM non-host plants mustard and sugar beet significantly reduced root colonization in cucumber plants, whereas no such effect was observed when root exudates of the AM non-host plant lupin were applied.     

Positive effects of an arbuscular mycorrhizal fungus on aphid life history traits

Two generations of two aphid species (Myzus ascalonicus and M. persicae) were reared on Plantago lanceolata plants, with and without root colonization by the arbuscular mycorrhizal fungus, Glomus intraradices. Life history traits of the aphids measured were nymphal development time, teneral adult weight, growth rate, total fecundity, adult longevity and duration of post-reproductive life. For both aphids in both generations, mycorrhizal colonization increased aphid weight and fecundity, while other traits were unaffected. The increases were consistent between generations. In a second experiment, M. persicae was reared on plants with and without the fungus, under varying N and P regimes. The results of N addition were inconclusive because there was high aphid mortality. However, under P supplementation, positive effects of the mycorrhiza on aphid growth were seen at low and medium P levels, while at high P levels these effects disappeared. The positive effects of mycorrhizal colonization reported here are contrary to the majority of previous studies with chewing insects, which have reported negative effects. A number of possible mechanisms for this apparent discrepancy are discussed. Received: 1 February 1999 / Accepted: 22 March 1999     

Improved Tolerance of <Emphasis Type="Italic">Acacia nilotica</Emphasis> to Salt Stress by Arbuscular Mycorrhiza, <Emphasis Type="Italic">Glomus fasciculatum</Emphasis> may be Partly Related to Elevated K/Na Ratios in Root and Shoot Tissues

A pot experiment was conducted to examine the effect of arbuscular mycorrhizal fungus, Glomus fasciculatum, and salinity on the growth of Acacia nilotica. Plants were grown in soil under different salinity levels (1.2, 4.0, 6.5, and 9.5 dS m⁻¹). In saline soil, mycorrhizal colonization was higher at 1.2, 4.0, and 6.5 dS m⁻¹ salinity levels in AM-inoculated plants, which decreased as salinity levels further increased (9.5 dS m⁻¹). Mycorrhizal plants maintained greater root and shoot biomass at all salinity levels compared to nonmycorrhizal plants. AM-inoculated plants had higher P, Zn, and Cu concentrations than uninoculated plants. In mycorrhizal plants, nutrient concentrations decreased with the increasing levels of salinity, but were higher than those of the nonmycorrhizal plants. Mycorrhizal plants had greater Na concentration at low salinity levels (1.2, 4.0 dS m⁻¹), which lowered as salinity levels increased (6.5, 9.5 dS m⁻¹), whereas Na concentration increased in control plants. Mycorrhizal plants accumulated a higher concentration of K at all salinity levels. Unlike Na, the uptake of K increased in shoot tissues of mycorrhizal plants with the increasing levels of salinity. Our results indicate that mycorrhizal fungus alleviates deleterious effects of saline soils on plant growth that could be primarily related to improved P nutrition. The improved K/Na ratios in root and shoot tissues of mycorrhizal plants may help in protecting disruption of K-mediated enzymatic processes under salt stress conditions.     

Mycorrhization alleviates benzo[a]pyrene-induced oxidative stress in an in vitro chicory root model

Among chemicals that are widely spread both in terrestrial and aquatic ecosystems, benzo[a]pyrene is a major source of concern. However, little is known about its adverse effects on plants, as well as about the role of mycorrhization in protection of plant grown in benzo[a]pyrene-polluted conditions. Hence, to contribute to a better understanding of the adverse effects of polycyclic aromatic hydrocarbons on the partners of mycorrhizal symbiotic association, benzo[a]pyrene-induced oxidative stress was studied in transformed Cichorium intybus roots grown in vitro and colonized or not by Glomus intraradices. The arbuscular mycorrhizal fungus development (colonization, extraradical hyphae length, and spore formation) was significantly reduced in response to increasing concentrations of benzo[a]pyrene (35–280 μM). The higher length of arbuscular mycorrhizal roots, compared to non-arbuscular mycorrhizal roots following benzo[a]pyrene exposure, pointed out a lower toxicity of benzo[a]pyrene in arbuscular mycorrhizal roots, thereby suggesting protection of the roots by mycorrhization. Accordingly, in benzo[a]pyrene-exposed arbuscular mycorrhizal roots, statistically significant decreases were observed in malondialdehyde concentration and 8-hydroxy-2′-desoxyguanosine formation. The higher superoxide dismutase activity detected in mycorrhizal chicory roots could explain the benzo[a]pyrene tolerance of the colonized roots. Taken together, these results support an essential role of mycorrhizal fungi in protecting plants submitted to polycyclic aromatic hydrocarbon, notably by reducing polycyclic aromatic hydrocarbon-induced oxidative stress damage.     

Arbuscular mycorrhiza enhances preference of ovipositing predatory mites for direct prey‐related cues

Most terrestrial plants are associated with arbuscular mycorrhizal fungi but research on the effects of arbuscular mycorrhizal symbiosis on aboveground plant‐associated organisms is scarcely expanded to tri‐trophic systems. The arbuscular mycorrhizal fungus Glomus mosseae Nicol. & Gerd. enhances fitness of the two‐spotted spider mite Tetranychus urticae Koch and its natural enemy, the predatory mite Phytoseiulus persimilis Athias‐Henriot, via changes in host plant and prey quality, respectively. In the present study, it is hypothesized that gravid P. persimilis are able to recognize arbuscular mycorrhiza‐enhanced prey quality and behave accordingly. In two experiments, on leaf arenas and in cages, P. persimilis is given a choice between prey patches deriving from mycorrhizal and non‐mycorrhizal bean plants (Phaseolus vulgaris L.) as feeding and oviposition sites. The use of cages allows the manipulation of distinct patch components acting as possible cues to guide predator foraging and oviposition behaviours, such as eggs produced and traces (webbing and faeces) left by the spider mite females. Both experiments show that P. persimilis preferentially resides close to prey fed on mycorrhizal plants. The cage experiment reveals that P. persimilis uses direct prey‐related cues, mainly derived from eggs, to discern prey quality and preferentially oviposits close to prey from mycorrhizal plants. This is the first study to document that predators recognize arbuscular mycorrhiza‐induced changes in herbivorous prey quality via direct prey‐related cues.     

Accumulation of secondary compounds in barley and wheat roots in response to inoculation with an arbuscular mycorrhizal fungus and co-inoculation with rhizosphere bacteria

 Colonization of Hordeum vulgare L. cv. Salome (barley)and Triticum aestivum L. cv. Caprimus (wheat) roots by the arbuscular mycorrhizal fungus Glomus intraradices Schenck & Smith leads to de novo synthesis of isoprenoid cyclohexenone derivatives with blumenin [9-O-(2′-O-β-glucuronosyl)-β-glucopyranoside of 6-(3-hydroxybutyl)-1,1,5-trimethyl-4-cyclohexen-3-one] as the major constituent and to transient accumulation of hydroxycinnamate amides (4-coumaroylagmatine and -putrescine). Accumulation of these compounds in mycorrhizal wheat roots started 2 weeks after sowing together with the onset of arbuscule formation and proceeded with mycorrhizal progression. Highest levels were reached in 3- to 4-week-old secondary roots (root branches of first and higher order) characterized by the formation of vesicles. In the final developmental stages, the fungus produced massive amounts of spores, enclosing the stele of older root parts (older than 5 weeks) characterized by cortical death. In these root parts, the secondary compounds were detected in trace amounts only, indicating that they were located in the cortical tissues. Some rhizosphere bacteria tested, i.e. Agrobacterium rhizogenes, Pseudomonas fluorescens, and Rhizobium leguminosarum, markedly stimulated both fungal root colonization and blumenin accumulation, thus, acting as mycorrhiza-helper bacteria (MHB). Application of blumenin itself strongly inhibited fungal colonization and arbuscule formation at early stages of mycorrhiza development. This was associated with a markedly reduced accumulation of the hydroxycinnamate amides 4-coumaroylputrescine and -agmatine. The results suggest that both the isoprenoid and the phenylpropanoid metabolism are closely linked to the developmental stage and the extent of fungal colonization. Their possible involvement in the regulation of mycorrhiza development is discussed. Accepted: 18 September 1998     

Acquisition of Cu, Zn, Mn and Fe by mycorrhizal maize (Zea mays L.) grown in soil at different P and micronutrient levels

Sustainability of soil-plant systems requires, among other things, good development and function of mycorrhizal symbioses. The effects of P and micronutrient levels on development of an arbuscular mycorrhizal fungus (AMF) and uptake of Zn, Cu, Mn and Fe by maize (Zea mays L.) were studied. A pot experiment with maize either inoculated or not with Glomus intraradices was conducted in a sand:soil (3 :1) mix (pH 6.5) in a greenhouse. Our goal was to evaluate the contribution of mycorrhizae to uptake of Cu, Zn, Mn and Fe by maize as influenced by soil P and micronutrient levels. Two levels of P (10 and 40 mg kg⁻¹ soil) and three levels of a micronutrient mixture: 0, 1X and 2X (1X contained, in mg kg⁻¹ soil, 4.2 Fe, 1.2 Mn, 0.24 Zn, 0.06 Cu, 0.78 B and 0.036 Mo), were applied to pots. There were more extraradical hyphae at the low P level than at the high P level when no micronutrients were added to the soil. Root inoculation with mycorrhiza and application of micronutrients increased shoot biomass. Total Zn content in shoots was higher in mycorrhizal than non-mycorrhizal plants grown in soils with low P and low or no micronutrient addition. Total Cu content in shoots was increased by mycorrhizal colonization when no micronutrients were added. Mycorrhizal plants had lower Mn contents than non-mycorrhizal plants only at the highest soil micronutrient level. AMF increased total shoot Fe content when no micronutrients were added, but decreased shoot Fe when plants were grown at the high level of micronutrient addition. The effects of G. intraradices on Zn, Cu, Mn, and Fe uptake varied with micronutrient and P levels added to soil. Accepted: 27 December 1999     

Growth and arsenic uptake by Chinese brake fern inoculated with an arbuscular mycorrhizal fungus

A split-root experiment investigated the effects of inoculation with the arbuscular mycorrhizal fungus Glomus mosseae and arsenic (As) addition on As uptake by Pteris vittata L. Either part or all of the root system was inoculated with G. mosseae or exposed to As addition (50 ml 1000 μmol L⁻¹ As 1 week before harvest). Mycorrhizal colonization substantially increased frond and root dry weight and P and As contents irrespective of As addition. Frond As contents in mycorrhizal plants were highest when the whole root system was exposed to As. Frond As concentrations and contents were higher when inoculation and As addition were in the same parts of the root system than when spatially separate. There were positive effects of arbuscular mycorrhiza inoculation on plant growth and As uptake, and inoculation of part of the roots seemed to be as effective as inoculation of the whole root system.     

Studies of iron transport by arbuscular mycorrhizal hyphae from soil to peanut and sorghum plants

 The influence of an arbuscular mycorrhizal (AM) fungus on phosphorus (P) and iron (Fe) uptake of peanut (Arachis hypogea L.) and sorghum (Sorghum bicolor L.) plants was studied in a pot experiment under controlled environmental conditions. The plants were grown for 10 weeks in pots containing sterilised calcareous soil with two levels of Fe supply. The soil was inoculated with rhizosphere microorganisms only or with rhizosphere microorganisms together with an AM fungus (Glomus mosseae [Nicol. & Gerd.] Gerdemann & Trappe). An additional small soil compartment accessible to hyphae but not roots was added to each pot after 6 weeks of plant growth. Radiolabelled P and Fe were supplied to the hyphae compartment 2 weeks after addition of this compartment. After a further 2 weeks, plants were harvested and shoots were analysed for radiolabelled elements. In both plant species, P uptake from the labelled soil increased significantly more in shoots of mycorrhizal plants than non-mycorrhizal plants, thus confirming the well-known activity of the fungus in P uptake. Mycorrhizal inoculation had no significant influence on the concentration of labelled Fe in shoots of peanut plants. In contrast, ⁵⁹Fe increased in shoots of mycorrhizal sorghum plants. The uptake of Fe from labelled soil by sorghum was particularly high under conditions producing a low Fe nutritional status of the plants. These results are preliminary evidence that hyphae of an arbuscular mycorrhizal fungus can mobilise and/or take up Fe from soil and translocate it to the plant. Accepted: 6 March 1998     

Influence of arbuscular mycorrhiza on clover and ryegrass grown together in a soil spiked with polycyclic aromatic hydrocarbons

 The effect of arbuscular mycorrhiza (AM) on white clover and ryegrass grown together in a soil spiked with polycyclic aromatic hydrocarbons (PAH) was assessed in a pot experiment. The soil was spiked with 500 mg kg^–1 anthracene, 500 mg kg^–1 chrysene and 50 mg kg^–1 dibenz(a,h)anthracene, representing common PAH compounds with three, four and five aromatic rings, respectively. Three treatments and two harvest times (8 and 16 weeks) were imposed on plants grown in spiked soil: no mycorrhizal inoculation, mycorrhizal inoculation (Glomus mosseae P2, BEG 69) and mycorrhizal inoculation and surfactant addition (Triton X-100). Pots without PAH were also included as a control of plant growth and mycorrhizal colonization as affected by PAH additions. The competitive ability of clover vis-à-vis ryegrass regarding shoot and root growth was enhanced by AM, but reduced by PAH and the added surfactant. This was reflected by mycorrhizal root colonization which was moderate for clover (20–40% of total root length) and very low for ryegrass (0.5–5% of total root length). Colonization of either plant was similar in spiked soil with and without the added surfactant, but the PAH reduced colonization of clover to half that in non-spiked soil. P uptake was maintained in mycorrhizal clover when PAH were added, but was reduced in non-mycorrhizal clover and in mycorrhizal clover that received surfactant. Similar effects were not observed on ryegrass. These results are discussed in the context of the natural attenuation of organic pollutants in soils. Accepted: 12 June 2000     

Response of 11 eucalyptus species to inoculation with three arbuscular mycorrhizal fungi

 Numerous publications have reported growth stimulation of Eucalyptus following ectomycorrhizal inoculation in nursery or field conditions. Although Eucalyptus species can also form arbuscular mycorrhiza, their dependency on this type of mycorrhiza is still debatable. This paper presents information on the effect of inoculation of arbuscular mycorrhizal fungi on eucalypt growth. Twenty weeks after mycorrhizal inoculation, Eucalyptus seedlings' stem dry weight could be increased up to 49% compared to non-inoculated control plants. Intensity of root colonization by a given fungus depended on the host species, but it was not related to a plant growth response. Leaf phosphorus concentration of non-inoculated Eucalyptus seedlings varied greatly between species. Increases in leaf phosphorus concentration following mycorrhizal infection were not necessarily associated with plant growth stimulation. The most mycorrhiza-dependent Eucalyptus species tended to be those having the highest leaf phosphorus concentration in the absence of a fungal symbiont. These mycorrhiza-dependent Eucalyptus species seem to have greater phosphorus requirements and consequently to rely more on the symbiotic association. Accepted: 1 September 1995