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Studies have suggested that increased root hydraulic conductivity in mycorrhizal roots could be the result of increased cell‐to‐cell water flux via aquaporins. This study aimed to elucidate if the key effect of the regulation of maize aquaporins by the arbuscular mycorrhizal (AM) symbiosis is the enhancement of root cell water transport capacity. Thus, water permeability coefficient (Pf) and cell hydraulic conductivity (Lpc) were measured in root protoplast and intact cortex cells of AM and non‐AM plants subjected or not to water stress. Results showed that cells from droughted‐AM roots maintained Pf and Lpc values of nonstressed plants, whereas in non‐AM roots, these values declined drastically as a consequence of water deficit. Interestingly, the phosphorylation status of PIP2 aquaporins increased in AM plants subjected to water deficit, and Pf values higher than 12 μm s−1 were found only in protoplasts from AM roots, revealing the higher water permeability of AM root cells. In parallel, the AM symbiosis increased stomatal conductance, net photosynthesis, and related parameters, showing a higher photosynthetic capacity in these plants. This study demonstrates a better performance of AM root cells in water transport under water deficit, which is connected to the shoot physiological performance in terms of photosynthetic capacity. An enhancement of the root cell water permeability was measured under drought both in intact cortex cells and in protoplasts from AM maize plants, concomitantly with regulation of some plant aquaporins, increase of PIP2s phosphorylation status, and enhanced photosynthetic capacity. [ABSTRACT FROM AUTHOR]

Jasmonic acid (JA) and arbuscular mycorrhizal (AM) symbioses are known to protect plants against abiotic and biotic stresses, but are also involved in the regulation of root hydraulic conductance (L). The objective of this experiment was to elucidate the role of JA in the water relations and hormonal regulation of AM plants under drought by using tomato plants defective in the synthesis of JA (def-1). Our results showed that JA is involved in the uptake and transport of water through its effect on both physiological parameters (stomatal conductance and L) and molecular parameters, mainly by controlling the expression and abundance of aquaporins. We observed that def-1 plants increased the expression of seven plant aquaporin genes under well-watered conditions in the absence of AM fungus, which partly explain the increment of L by this mutation under well-watered conditions. In addition, the effects of the AM symbiosis on plants were modified by the def-1 mutation, with the expression of some aquaporins and plant hormone concentration being disturbed. On the other hand, methyl salicylate (MeSA) content was increased in non-mycorrhizal def-1 plants, suggesting that MeSA and JA can act together in the regulation of L. In a complementary experiment, it was found that exogenous MeSA increased L, confirming our hypothesis. Likewise, we confirmed that JA, ABA and SA are hormones involved in plant mechanisms to cope with stressful situations, their concentrations being controlled by the AM symbiosis. In conclusion, under wellwatered conditions, the def-1 mutation mimics the effects of AM symbiosis, but under drought conditions the def-1 mutation changed the effects of the AM symbiosis on plants. [ABSTRACT FROM AUTHOR]

The arbuscular mycorrhizal (AM) symbiosis has been shown to improve maize tolerance to different drought stress scenarios by regulating a wide range of host plants aquaporins. The objective of this study was to highlight the differences in aquaporin regulation by comparing the effects of the AM symbiosis on root aquaporin gene expression and plant physiology in two maize cultivars with contrasting drought sensitivity. This information would help to identify key aquaporin genes involved in the enhanced drought tolerance by the AM symbiosis. Results showed that when plants were subjected to drought stress the AM symbiosis induced a higher improvement of physiological parameters in drought-sensitive plants than in drought-tolerant plants. These include efficiency of photosystem II, membrane stability, accumulation of soluble sugars and plant biomass production. Thus, drought-sensitive plants obtained higher physiological benefit from the AM symbiosis. In addition, the genes ZmPIP1;1, ZmPIP1;3, ZmPIP1;4, ZmPIP1;6, ZmPIP2;2, ZmPIP2;4, ZmTIP1;1, and ZmTIP2;3 were down-regulated by the AM symbiosis in the drought-sensitive cultivar and only ZmTIP4;1 was up-regulated. In contrast, in the drought-tolerant cultivar only three of the studied aquaporin genes (ZmPIP1;6, ZmPIP2;2, and ZmTIP4;1) were regulated by the AM symbiosis, resulting induced. Results in the drought-sensitive cultivar are in line with the hypothesis that down-regulation of aquaporins under water deprivation could be a way to minimize water loss, and the AM symbiosis could be helping the plant in this regulation. Indeed, during drought stress episodes, water conservation is critical for plant survival and productivity, and is achieved by an efficient uptake and stringently regulated water loss, in which aquaporins participate. Moreover, the broader and contrasting regulation of these aquaporins by the AM symbiosis in the drought-sensitive than the drought-tolerant cultivar suggests a role of these aquaporins in water homeostasis or in the transport of other solutes of physiological importance in both cultivars under drought stress conditions, which may be important for the AM-induced tolerance to drought stress. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM PUBLISHER]

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. [ABSTRACT FROM PUBLISHER]

Boron (B) is an essential micronutrient for higher plants, having structural roles in primary cell walls, but also other functions in cell division, membrane integrity, pollen germination or metabolism. Both high and low B levels negatively impact crop performance. Thus, plants need to maintain B concentration in their tissues within a narrow range by regulating transport processes. Both active transport and protein-facilitated diffusion through aquaporins have been demonstrated. This study aimed at elucidating the possible involvement of some plant aquaporins, which can potentially transport B and are regulated by the arbuscular mycorrhizal (AM) symbiosis in the plant B homeostasis. Thus, AM and non-AM plants were cultivated under 0, 25 or 100 μM B in the growing medium and subjected or not subjected to drought stress. The accumulation of B in plant tissues and the regulation of plant aquaporins and other B transporters were analyzed. The benefits of AM inoculation on plant growth (especially under drought stress) were similar under the three B concentrations assayed. The tissue B accumulation increased with B availability in the growing medium, especially under drought stress conditions. Several maize aquaporins were regulated under low or high B concentrations, mainly in non-AM plants. However, the general down-regulation of aquaporins and B transporters in AM plants suggests that, when the mycorrhizal fungus is present, other mechanisms contribute to B homeostasis, probably related to the enhancement of water transport, which would concomitantly increase the passive transport of this micronutrient. [ABSTRACT FROM AUTHOR]

This study investigates the possible involvement of maize aquaporins which are regulated by arbuscular mycorrhizae (AM) in the transport in planta of ammonium and/or urea under well-watered and drought stress conditions. The study also aims to better understand the implication of the AM symbiosis in the uptake of urea and ammonium and its effect on plant physiology and performance under drought stress conditions. AM and non-AM maize plants were cultivated under three levels of urea or ammonium fertilization (0, 3 µM or 10 mM) and subjected or not to drought stress. Plant aquaporins and physiological responses to these treatments were analyzed. AM increased plant biomass in absence of N fertilization or under low urea/ ammonium fertilization, but no effect of the AM symbiosis was observed under high N supply. This effect was associated with reduced oxidative damage to lipids and increased N accumulation in plant tissues. High N fertilization with either ammonium or urea enhanced net photosynthesis (AN) and stomatal conductance (gs) in plants maintained under well-watered conditions, but 14 days after drought stress imposition these parameters declined in AM plants fertilized with high N doses. The aquaporin ZmTIP1;1 was up-regulated by both urea and ammonium and could be transporting these two N forms in planta. The differential regulation of ZmTIP4;1 and ZmPIP2;4 with urea fertilization and of ZmPIP2;4 with NH4+ supply suggests that these two aquaporins may also play a role in N mobilization in planta. At the same time, these aquaporins were also differentially regulated by the AM symbiosis, suggesting a possible role in the AM-mediated plant N homeostasis that deserves future studies. [ABSTRACT FROM AUTHOR]

Abstract: In this study, we have determined whether the arbuscular mycorrhizal (AM) symbiosis is able to alter the pattern of Δ1-pyrroline-5-carboxylate synthetase (p5cs) gene expression under drought stress and whether such possible alteration functions in the protection of the host plants against drought. To achieve this, we cloned a P5CS-encoding gene from Glycine max (gmp5cs) and another from Lactuca sativa (lsp5cs) and analyzed their contribution to the response against drought in control and AM soybean and lettuce plants. The analysis of gmp5cs and lsp5cs gene expression showed that these genes were up-regulated by drought stress. The highest gene expression was found in non-inoculated plants subjected to drought. A contrasting result was obtained in soybean plants singly inoculated with Bradyrhizobium japonicum, where the gmp5cs gene showed little up-regulation in roots under drought stressed conditions. Moreover, both soybean and lettuce AM plants showed lower p5cs transcript accumulation under drought stress than non-inoculated plants. The present results indicate that the induction of p5cs gene is not a mechanism by which the AM symbiosis protects their host plant against drought. [Copyright &y& Elsevier]

Climate change is leading to the intensification of drought effects worldwide, which considerably reduce crop production. A better understanding of the drought-tolerance mechanisms would lead into a more productive agriculture. The arbuscular mycorrhizal (AM) symbiosis has been shown to improve plant tolerance to drought. Salicylic acid (SA) is a phenolic compound involved in many aspects of plant growth and development. Apart from its role in biotic interactions, it is also involved in the regulation of important plant physiological processes, including plant water relations under stressful conditions. However, despite the importance of SA in plant physiology and in AM colonization, little is known about its effect on regulation of root water transport. Thus, the aim of this work was to study the combined effect of AM symbiosis and SA on root hydraulic properties under drought stress, with special focus on how these factors can alter radial root water transport pathways through aquaporin regulation. Also, the crosstalk between SA and other phytohormones was taken into account. Results showed that the AM symbiosis modifies root hydraulic responses to drought episodes. Under these conditions, AM plants showed increased Lpr and Lo. Exogenous SA application decreased Lpr and Lo under drought. SA modulation of water conductivity could be due to a fine-regulation of root aquaporins (as ZmPIP2;4 or ZmTIP1;1 ). Furthermore, SA application differently modulated the percentage of water flowing by the apoplastic pathway, decreasing its contribution to total root water flow in AM plants and increasing it in non-AM plants. An intricate relationship between Lpr, aquaporins and endogenous levels of SA, ABA and jasmonic acid was observed. Future studies should explore more in detail the crosstalk mechanism between these hormones in the regulation of water transport in AM roots, in order to better understand the mechanism through which the AM symbiosis copes with drought stress. [ABSTRACT FROM AUTHOR]

At a world scale, tomato is an important horticultural crop, but its productivity is highly reduced by drought stress. Combining the application of beneficial microbial inoculants with breeding and grafting techniques may be key to cope with reduced tomato yield under drought. This study aimed to investigate the growth responses and physiological mechanisms involved in the performance under drought stress of four tomato recombinant inbred lines (RIL) after inoculation with the arbuscular mycorrhizal (AM) fungus Rhizophagus irregularis and the plant growth promoting rhizobacteria (PGPR) Variovorax paradoxus 5C-2. Results showed a variation in the efficiency of the different tomato RILs under drought stress and a differential effect of the microbial inoculants, depending on the RIL involved. The inoculants affected plant parameters such as net photosynthetic capacity, oxidative damage to lipids, osmolyte accumulation, root hydraulic conductivity or aquaporin abundance and phosphorylation status. RIL66 was the one obtaining maximum benefit from the microbial inoculants under drought stress conditions, due likely to improved CO 2 -fixation capacity and root hydraulic conductivity. We propose that RIL66 could be selected as a good plant material to be used as rootstock to improve tomato growth and productivity under water limiting conditions. Since RIL66 is highly responsive to microbial inoculants, this grafting strategy should be combined with inoculation of R. irregularis and V. paradoxus in order to improve plant yield under conditions of drought stress. [ABSTRACT FROM AUTHOR]

Rice is the most important food crop in the world and is a primary source of food for more than half of the world population. However, salinity is considered the most common abiotic stress reducing its productivity. Soil salinity inhibits photosynthetic processes, which can induce an over-reduction of the reaction centres in photosystem II (PSII), damaging the photosynthetic machinery. The arbuscular mycorrhizal (AM) symbiosis may improve host plant tolerance to salinity, but it is not clear how the AM symbiosis affects the plant photosynthetic capacity, particularly the efficiency of PSII. This study aimed at determining the influence of the AM symbiosis on the performance of PSII in rice plants subjected to salinity. Photosynthetic activity, plant gas-exchange parameters, accumulation of photosynthetic pigments and rubisco activity and gene expression were also measured in order to analyse comprehensively the response of the photosynthetic processes to AM symbiosis and salinity. Results showed that the AM symbiosis enhanced the actual quantum yield of PSII photochemistry and reduced the quantum yield of non-photochemical quenching in rice plants subjected to salinity. AM rice plants maintained higher net photosynthetic rate, stomatal conductance and transpiration rate than nonAM plants. Thus, we propose that AM rice plants had a higher photochemical efficiency for CO 2 fixation and solar energy utilization and this increases plant salt tolerance by preventing the injury to the photosystems reaction centres and by allowing a better utilization of light energy in photochemical processes. All these processes translated into higher photosynthetic and rubisco activities in AM rice plants and improved plant biomass production under salinity. [ABSTRACT FROM AUTHOR]

Rice (Oryza sativa L.) is the most important crop for human consumption is grown conventionally in flooded conditions throughout their cycle. However, approximately half the surface of rice in the world does not have sufficient water to maintain optimal growth conditions. One possible way to increase rice production to meet demand is to improve the efficient use of water by arbuscular mycorrhizal fungi (AMF). The research was conducted at the Experimental Station of Zaidín, Granada Spain, with mycorrhizal and non-mycorrhizal rice plant under controlled conditions and with three water management, without stress (25 mL), moderate stress (10 mL) and severe stress (5 mL) for 15 days, in order to evaluate the effect of AMF inoculation of rice plants subjected to hydric stress and after the plants recovered. The results show that rice can benefit from the symbiosis MA and improve long-term development after a period of water stress. In fact, at each level of irrigation, the mycorrhizal plants showed a 50 % increase in aerial fresh mass and 40 % photosynthetic efficiency compared to non-mycorrhizal plants. Besides, the proline content increased with hydric stress in mycorrizal and non-mycorrizal plants. [ABSTRACT FROM AUTHOR]

Abstract: The response of rice plants to inoculation with an arbuscular mycorrhizal (AM) fungus, Azospirillum brasilense, or combination of both microorganisms, was assayed under well-watered or drought stress conditions. Water deficit treatment was imposed by reducing the amount of water added, but AM plants, with a significantly higher biomass, received the same amount of water as non-AM plants, with a poor biomass. Thus, the water stress treatment was more severe for AM plants than for non-AM plants. The results showed that AM colonization significantly enhanced rice growth under both water conditions, although the greatest rice development was reached in plants dually inoculated under well-watered conditions. Water level did not affect the efficiency of photosystem II, but both AM and A. brasilense inoculations increased this value. AM colonization increased stomatal conductance, particularly when associated with A. brasilense, which enhanced this parameter by 80% under drought conditions and by 35% under well-watered conditions as compared to single AM plants. Exposure of AM rice to drought stress decreased the high levels of glutathione that AM plants exhibited under well-watered conditions, while drought had no effect on the ascorbate content. The decrease of glutathione content in AM plants under drought stress conditions led to enhance lipid peroxidation. On the other hand, inoculation with the AM fungus itself increased ascorbate and proline as protective compounds to cope with the harmful effects of water limitation. Inoculation with A. brasilense also enhanced ascorbate accumulation, reaching a similar level as in AM plants. These results showed that, in spite of the fact that drought stress imposed by AM treatments was considerably more severe than non-AM treatments, rice plants benefited not only from the AM symbiosis but also from A. brasilense root colonization, regardless of the watering level. However, the beneficial effects of A. brasilense on most of the physiological and biochemical traits of rice plants were only clearly visible when the plants were mycorrhized. This microbial consortium was effective for rice plants as an acceptable and ecofriendly technology to improve plant performance and development. [Copyright &y& Elsevier]

Abstract: Rice (Oryza sativa) is the most important crop for human consumption, providing staple food for more than half of the world’s population. Rice is conventionally grown under flooded conditions for most of its growing cycle. However, about half of the rice area in the world does not have sufficient water to maintain optimal growing conditions and yield is reduced by drought. One possible way to increase rice production in order to meet the rice demand is to improve its drought tolerance by means of the arbuscular mycorrhizal (AM) symbiosis. Thus, AM and non-AM rice plants were maintained under well-watered conditions or were subjected to moderate and severe drought stress for 15d. After that, half of the plants from each treatment were harvested, while the other half were allowed to recover from drought for additional 25d. The results showed that rice can benefit from the AM symbiosis and improve their long-term development after a drought stress period. In fact, at each watering level, AM plants showed about 50% enhanced shoot fresh weight as compared to non-AM plants. The AM symbiosis enhanced the plant photosynthetic efficiency under stress over 40%, induced the accumulation of the antioxidant molecule glutathione and reduced the accumulation of hydrogen peroxide and the oxidative damage to lipids in these plants. Thus, these combined effects enhanced the plant performance after a drought stress period. [Copyright &y& Elsevier]

Abstract: The plant growth, nutrient acquisition, metal translocation and antioxidant activities [ascorbate peroxidase (APX), glutatione reductase (GR), superoxide dismutase (SOD) and catalase (CAT)] were measured in plants growing in a heavy-metal (HM) multicontaminated soil inoculated with selected autochthonous microorganisms [arbuscular mycorrhizal (AM) fungus and/or plant growth promoting bacteria (PGPB)] and/or amended with an Aspergillus niger-treated agrowaste. The treated agrowaste on its own increased root growth by 296% and shoot growth by 504% compared with non-treated control plants. Both chemical and biological treatments, particularly when combined, enhanced plant shoot and root development. The stimulation effect on plant biomass was concomitant with increased AM colonization, P and K assimilation, and reduced metal translocation from soil to plant shoot. The treated residue, particularly through interactions with AM inoculation, produced the expected bioremediation effect, leading to enhanced plant development and successful rehabilitation of contaminated soil. The enhancement of CAT, APX and GR activities caused by AM inoculation suggests that AM colonization helped plants to limit oxidative damage to biomolecules in response to metal stress. The response of the plant''s antioxidant activities to the amendment appears to be related to enhanced plant biomass production. The application of amendments and/or microbial inoculations to enhance plant growth and reduce metal translocation in multicontaminated soil could be a promising strategy for remediating HM pollution. [Copyright &y& Elsevier]

The contribution of the arbuscular mycorrhizal (AM) symbiosis to plant drought tolerance results from a combination of physical, nutritional and cellular effects. However, the exact mechanisms involved in such enhanced tolerance are still a matter of debate. In this study a BiP-encoding gene from the AM fungus Glomus intraradices has been identified after differential hybridization of a cDNA library constructed from the fungus growing in vitro and subjected to drought stress by addition of 25% PEG 6000. Results show that its expression was up-regulated by drought stress not only during in vitro conditions (AM monoxenic cultures) but also ex vitro, when forming natural symbioses with plants. The identification of GiBiP gene provides new evidences that the contribution of AM fungi to the enhanced drought tolerance of the host plant can be mediated by proteins with chaperone-like activity, such as that of BiP. [Copyright &y& Elsevier]

• IAA affected maize root hydraulic, mainly decreasing root hydraulic conductivity (Lo). • Aquaporins are likely involved in the IAA-dependent inhibition of Lo. • IAA differentially regulated apoplastic water flow in AM and non-AM plants. • Other phytohormones contributed to the IAA-dependent effects on root hydraulics. Drought stress is one of the most devastating abiotic stresses, compromising crop growth, reproductive success and yield. The arbuscular mycorrhizal (AM) symbiosis has been demonstrated to be beneficial in helping the plant to bear with water deficit. In plants, development and stress responses are largely regulated by a complex hormonal crosstalk. Auxins play significant roles in plant growth and development, in responses to different abiotic stresses or in the establishment and functioning of the AM symbiosis. Despite these important functions, the role of indole-3acetic acid (IAA) as a regulator of root water transport and stress response is not well understood. In this study, the effect of exogenous application of IAA on the regulation of root radial water transport in AM plants was analyzed under well-watered and drought stress conditions. Exogenous IAA application affected root hydraulic parameters, mainly osmotic root hydraulic conductivity (Lo), which was decreased in both AM and non-AM plants under water deficit conditions. Under drought, the relative apoplastic water flow was differentially regulated by IAA application in non-AM and AM plants. The effect of IAA on the internal cell component of root water conductivity suggests that aquaporins are involved in the IAA-dependent inhibition of this water pathway. [ABSTRACT FROM AUTHOR]

• The AM symbiosis significantly modified the radial root water transport in maize. • Lpr, Lo and apoplastic water flow raised in AM plants as compared to non-AM ones. • Sodium azide inhibited Lo in non-AM plants but not in AM plants. • This seems to be due to posttranslational regulation of aquaporins activity. In roots, water flows radially through three parallel pathways: apoplastic, symplastic and transcellular (the last two referred as the cell-to-cell), with a different contribution depending on the environmental conditions. Thus, during drought, the cell-to-cell pathway, which is largely regulated by aquaporins, dominates. While it is accepted that water can flow across roots following the apoplastic, symplastic and transcellular pathways, the relative contribution of these pathways to whole root hydraulic conductivity is not well stablished. In addition, the symbiosis with arbuscular mycorrhizal (AM) fungi was reported to modify root water transport in host plants. This study aims to understand if the AM symbiosis alters radial root water transport in the host plant and whether this modification is due to alteration of plant aquaporins activity or amounts and/or changes in apoplastic barriers. Hence, the combined effect of mycorrhizal fungus, water deficit and application of the aquaporin inhibitor sodium azide (NaN 3) on radial root water transport of maize plants was analyzed. The development of Casparian bands in these roots was also assessed. NaN 3 clearly inhibited osmotic root hydraulic conductivity (Lo). However, the inhibitory effect of sodium azide on Lo was lower in AM plants than in non-AM plants, which together with their higher relative apoplastic water flow values suggests a compensatory mechanism for aquaporin activity inhibition in AM plants, leading to a higher hydrostatic root hydraulic conductivity (Lpr) compared to non-AM plants. This effect seems to be related to the mycorrhizal regulation of aquaporins activity through posttranslational modifications. The development of Casparian bands increased with drought and AM colonization, although this did not decrease water flow values in AM plants. The work provides new clues on the differential mycorrhizal regulation of root water transport. [ABSTRACT FROM AUTHOR]