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We investigated in two tobacco (Nicotiana tabacum) plant lines (wildtype or antisense mutant) whether impairment in expression of the plasma membrane aquaporin gene (NtAQP1) affects the arbuscular mycorrhizal (AM) fungal colonisation pattern or the symbiotic efficiency of AM fungi. These two objectives were investigated under well-watered and drought stress conditions. Both plant lines had a similar pattern of root colonisation under well-watered and drought stress conditions. In contrast, under drought stress, AM wildtype plants grew faster than mycorrhizal antisense plants. Plant gas exchange also appeared to depend on the expression of NtAQP1 and parallelled the determined growth increments. The implications of enhanced symplastic water transport via NtAQP1 for the efficiency of the AM symbiosis under drought stress conditions are further discussed.

In this study, it has been determined whether the arbuscular mycorrhizal (AM) symbiosis is able to alter the pattern of dehydrin (LEA D-11 group) transcript accumulation under drought stress, and whether such a possible alteration functions in the protection of the host plants against drought. Two dehydrin-encoding genes have been cloned from Glycine max (gmlea 8 and gmlea 10) and one from Lactuca sativa (lslea 1) and they have been analysed for their contribution to the response against drought in mycorrhizal soybean and lettuce plants. Results with soybean plants showed that most of the treatments did not show LEA gene expression under well-watered conditions. The higher gene expression was found in non-inoculated plants subjected to drought. Only plants singly inoculated with Bradyrhizobium japonicum showed an important level of LEA gene expression under well-watered conditions and a reduced level under drought-stress conditions. The same results were confirmed in subsequent experiments and at the latest stage of a time-course experiment. In lettuce, the lslea 1 gene was also induced by drought stress in all treatments. However, the level of induction was clearly higher in roots from non-inoculated plants than in roots from the two AM treatments assayed. The overall results demonstrated that the levels of lea transcript accumulation in mycorrhizal treatments subjected to drought were considerably lower than in the corresponding non-mycorrhizal plants, indicating that the accumulation of LEA proteins is not a mechanism by which the AM symbiosis protects their host plant.

This study investigated several aspects related to drought tolerance in arbuscular mycorrhizal (AM) soybean plants. The investigation included both shoot and root tissues in order to reveal the preferred target tissue for AM effects against drought stress. Non-AM and AM soybean plants were grown under well-watered or drought-stressed conditions, and leaf water status, solute accumulation, oxidative damage to lipids, and other parameters were determined. Results showed that AM plants were protected against drought, as shown by their significantly higher shoot-biomass production. The leaf water potential was also higher in stressed AM plants (-1.9 MPa) than in non-AM plants (-2.5 MPa). The AM roots had accumulated more proline than non-AM roots, while the opposite was observed in shoots. Lipid peroxides were 55% lower in shoots of droughted AM plants than in droughted non-AM plants. Since there was no correlation between the lower oxidative damage to lipids in AM plants and the activity of antioxidant enzymes, it seems that first the AM symbiosis enhanced osmotic adjustment in roots, which could contribute to maintaining a water potential gradient favourable to the water entrance from soil into the roots. This enabled higher leaf water potential in AM plants during drought and kept the plants protected against oxidative stress, and these cumulative effects increased the plant tolerance to drought.

Water deficit is considered one of the most important abiotic factors limiting plant growth and yield in many areas on earth. Several eco-physiological studies have demonstrated that the arbuscular mycorrhizal (AM) symbiosis often results in altered rates of water movement into, through and out of the host plants, with consequent effects on tissue hydration and plant physiology. It is now accepted that the contribution of AM symbiosis to plant drought tolerance is the result of accumulative physical, nutritional, physiological and cellular effects. This review considers several aspects that should be investigated at a molecular level in order to gain a whole understanding of the different mechanisms by which the AM symbiosis protects the host plants against the detrimental effects of water deficit.

We isolated two bacterial strains from an experimentally lead (Pb)-polluted soil in Hungary, 10 years after soil contamination. These strains represented the two most abundant cultivable bacterial groups in such soil, and we tested their influence on Trifolium pratense L. growth and on the functioning of native mycorrhizal fungi under Pb toxicity in a second Pb-spiked soil. Our results showed that bacterial strain A enhanced plant growth, nitrogen and phosphorus accumulations, nodule formation, and mycorrhizal infection, demonstrating its plant-growth-promoting activity. In addition, strain A decreased the amount of Pb absorbed by plants, when expressed on a root weight basis, because of increased root biomass due to the production of indoleacetic acid. The positive effect of strain A was not only evident after a single inoculation but also in dual inoculation with arbuscular mycorrhizal fungi. Strain A also exhibited higher tolerance than strain B when cultivated under increasing Pb levels in the spiked soil. Molecular identification unambiguously placed strain A within the genus Brevibacillus. We showed that it is important to select the most tolerant and efficient bacterial strain for co-inoculation with arbuscular mycorrhizal fungi to promote effective symbiosis and thus stimulate plant growth under adverse environmental conditions, such as heavy-metal contamination.

The effects of bacterial inoculation (Bacillus sp.) on the development and physiology of the symbiosis between lettuce and the arbuscular mycorrhizal (AM) fungi Glomus mosseae (Nicol. and Gerd.) Gerd. and Trappe and Glomus intraradices (Schenck and Smith) were investigated. Plant growth, mineral nutrition and gas-exchange values in response to bacterial inoculation after PEG-induced drought stress were also evaluated. In AM plants, inoculation with Bacillus sp. enhanced fungal development and metabolism, measured as succinate dehydrogenase (SDH) and alkaline phosphatase (ALP) activities, more than plant growth. Under non-stressed conditions, G. intraradices colonization increased all plant physiological values to a higher extent when in dual inoculation with the bacterium. Under stress conditions, the bacterium had an important stimulatory effect on G. intraradices development. Under such conditions, the effects of the bacterium on photosynthetic rate, water use efficiency (WUE) and stomatal conductance of lettuce plants differed with the fungus species. Plant-gas exchange was enhanced in G. intraradices- and reduced in G. mosseae-colonized plants when co-inoculated with Bacillus sp. Thus, the effects of each fungus on plant physiology were modulated by the bacterium. Stress was detrimental, particularly in G. intraradices-colonized plants without the bacterium, reducing intra and extraradical mycelium growth and vitality (SDH), as well as plant-gas exchange. Nevertheless, Bacillus sp. inoculation improved all these plant and fungal parameters to the same level as in non-stressed plants. The highest amount of alive and active AM mycelium for both fungi was obtained after co-inoculation with Bacillus sp. These results suggest that selected free-living bacteria and AM fungi should be co-inoculated to optimize the formation and functioning of the AM symbiosis in both normal and adverse environments.

•  The mechanisms by which the mycorrhizal symbiosis protects soybean ( Glycine max ) plants against premature nodule senescence induced by drought stress is investigated here by evaluating the activity of a set of antioxidant enzymes in relation to nodule senescence. •  Superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and glutathione reductase (GR) activity was determined in well watered or drought-stressed soybean plants inoculated with Bradyrrhizobium japonicum alone or in combination with Glomus mosseae . •  In roots, only GR activity was higher in mycorrhizal than in non-mycorrhizal plants. The other antioxidant activities were similar, or lower (APX), in droughted, mycorrhizal plants than in the corresponding nonmycorrhizal ones. Similarly, in nodules, SOD, CAT and APX activities were lower in droughted, mycorrhizal plants than in nonmycorrhizal plants whereas, again, GR activity was higher in nodules from mycorrhizal plants. •  We propose that the consistently higher GR activity in roots and nodules of mycorrhizal plants might have contributed to decreased oxidative damage to biomolecules, which are involved in premature nodule senescence. Additional drought-avoidance mechanisms induced by the AM symbiosis might also contribute to the lower oxidative stress in mycorrhizal plants.

A cDNA library was constructed with RNA from Glomus intraradices-colonized lettuce roots and used for differential screening. This allowed the identification of a cDNA (Gi-1) that was expressed only in mycorrhizal roots and was of fungal origin. The function of the gene product is unknown, because Gi-1 contained a complete open reading frame that was predicted to encode a protein of 157 amino acids which only showed little homology with glutamine synthetase from Helicobacter pylori. The time-course analysis of gene expression during the fungal life cycle showed that Gi-1 was expressed only during the mycorrhizal symbiosis and was not detected in dormant or germinating spores of G. intraradices. P fertilization did not significantly change the pattern of Gi-1 expression compared with that in the unfertilized treatment, whereas N fertilization (alone or in combination with P) considerably enhanced the Gi-1 transcript accumulation. This increase in gene expression correlated with plant N status and growth under such conditions. The possible role of the Gi-1 gene product in intermediary N metabolism of arbuscular mycorrhizal symbiosis is further discussed.

In the present study three cDNA fragments were cloned using degenerate primers for Mn-sod genes and PCR: two showed a high degree of identity with Mn-sods from plants and the third with Fe-sod. Arbuscular mycorrhizal (AM) symbiosis down-regulated their expression pattern under well-watered conditions. In contrast, AM symbiosis in combination with drought stress considerably increased the expression of the Mn-sod II gene and this correlated well with plant tolerance to drought. These results would suggest that mycorrhizal protection against oxidative stress caused by drought may be an important mechanism by which AM fungi protect the host plant against drought.

The arbuscular mycorrhizal (AM) fungus Gigaspora margarita harbors a resident population of endosymbiontic Burkholderia in its cytoplasm. Nothing is known about the acquisition of such bacteria and about the molecular bases which allow colonization of the fungus. We wondered whether the intracellular Burkholderia strain possesses genetic determinants involved in colonization of a eukaryotic cell. Using degenerated oligonucleotide primers for vacB, a gene involved in host cell colonization by pathogenic bacteria, an 842 bp DNA fragment was cloned, sequenced, and identified as a part of the vacB gene in Burkholderia sp. The insert was used as a probe to screen a fungal library that, because of the presence of intracellular Burkholderia cells, was also representative of the bacterial genome. The complete nucleotide sequence of vacB and flanking genes was determined. The bacterial origin of this genomic region was established by PCR, using specific vacB primers on DNA from Gigasporaceae that did or did not contain cytoplasmic Burkholderia, as well as on DNA from other bacteria, including free-living Burkholderia. We hypothesize that the vacB gene is part of a new genetic region acquired by a rhizospheric Burkholderia strain, which became able to establish a symbiotic interaction with the AM fungus G. margarita.

This article reports the identification of a putative P-transporter operon in the genome of a Burkholderia sp. living in the cytoplasm of the arbuscular mycorrhizal fungus Gigaspora margarita. Its presence suggests that Burkholderia sp. has the potential for P uptake from this environment. This finding raises new questions concerning the importance of intracellular bacteria for mycorrhizal symbiosis.

The tolerance of lettuce plants (Lactuca sativa L. cv. Romana) to drought stress differed with the arbuscular-mycorrhizal fungal isolate with which the plants were associated. Seven fungal species belonging to the genus Glomus were studied for their ability to enhance the drought tolerance of lettuce plants. These fungi had different traits that affected the drought resistance of host plants. The ranking of arbuscular-mycorrhizal fungal effects on drought tolerance, based on the relative decreases in shoot dry weight, was as follows: Glomus deserticola > Glomus fasciculatum > Glomus mosseae > Glomus etunicatum > Glomus intraradices > Glomus caledonium > Glomus occultum. In this comparative study specific mycorrhizal fungi had consistent effects on plant growth, mineral uptake, the CO(inf2) exchange rate, water use efficiency, transpiration, stomatal conductance, photosynthetic phosphorus use efficiency, and proline accumulation under either well-watered or drought-stressed conditions. The ability of the isolates to maintain plant growth effectively under water stress conditions was related to higher transpiration rates, levels of leaf conductance, and proline, N, and P contents. Differences in proline accumulation in leaves among the fungal symbioses suggested that the fungi were able to induce different degrees of osmotic adjustment. The detrimental effects of drought were not related to decreases in photosynthesis or water use efficiency. Neither of these parameters was related to P nutrition. The differences in P and K acquisition, transpiration, and stomatal conductance were related to the mycorrhizal efficiencies of the different fungi. Our observations revealed the propensities of different Glomus species to assert their protective effects during plant water stress. The greater effectiveness of G. deserticola in improving water deficit tolerance was associated with the lowest level of growth reduction (9%) under stress conditions. The growth of plants colonized by G. occultum was reduced by 70% after a progressive drought stress period. In general, the different protective effects of the mycorrhizal isolates were not associated with colonizing ability. Nevertheless, G. deserticola was the most efficient fungus and exhibited the highest levels of mycorrhizal colonization, as well as the greatest stimulation of physiological parameters.