Arbuscular Mycorrhizal Symbiosis Alleviates Salt Stress in Black Locust through Improved Photosynthesis, Water Status, and K+/Na+ Homeostasis

Soil salinization and the associated land degradation are major and growing ecological problems. Excess salt in soil impedes plant photosynthetic processes and root uptake of water and nutrients such as K+. Arbuscular mycorrhizal (AM) fungi can mitigate salt stress in host plants. Although, numerous studies demonstrate that photosynthesis and water status are improved by mycorrhizae, the molecular mechanisms involved have received little research attention. In the present study, we analyzed the effects of AM symbiosis and salt stress on photosynthesis, water status, concentrations of Na+ and K+, and the expression of several genes associated with photosynthesis ( ) and genes coding for aquaporins or membrane transport proteins involved in K+ and/or Na+ uptake, translocation, or compartmentalization homeostasis (RpSOS1, RpHKT1, RpNHX1, and RpSKOR) in black locust. The results showed that salinity reduced the net photosynthetic rate, stomatal conductance, and relative water content in both non-mycorrhizal (NM) and AM plants; the reductions of these three parameters were less in AM plants compared with NM plants. Under saline conditions, AM fungi significantly improved the net photosynthetic rate, quantum efficiency of photosystem II photochemistry, and K+ content in plants, but evidently reduced the Na+ content. AM plants also displayed a significant increase in the relative water content and an evident decrease in the shoot/root ratio of Na+ in the presence of 200 mM NaCl compared with NM plants. Additionally, mycorrhizal colonization upregulated the expression of three chloroplast genes  in leaves, and three genes (RpSOS1, RpHKT1, and RpSKOR) encoding membrane transport proteins involved in K+/Na+ homeostasis in roots. Expression of several aquaporin genes was regulated by AM symbiosis in both leaves and roots depending on soil salinity. This study suggests that the beneficial effects of AM symbiosis on the photosynthetic capacity, water status, and K+/Na+ homeostasis lead to the improved growth performance and salt tolerance of black locust exposed to salt stress.

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title: Arbuscular mycorrhizal fungi in alleviation of salt stress: a review.
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This review addresses the significance of arbuscular mycorrhiza in alleviation of salt stress and their beneficial effects on plant growth and productivity. It also focuses on recent progress in unravelling biochemical, physiological and molecular mechanisms in mycorrhizal plants to alleviate salt stress.



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title: Photosynthesis under drought and salt stress: regulation mechanisms from whole plant to cell.
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title: Effects of salinity stress on photosystem II function in cyanobacterial Spirulina platensis cells.
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title: Salt stress inhibits photosystems II and I in cyanobacteria
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title: Low threshold levels of ultraviolet-B in a background of photosynthetically active radiation trigger rapid degradation of the D2 protein of photosystem-II
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title: The effects of salt stress on photosynthetic electron transport and thylakoid membrane proteins in the cyanobacterium Spirulina platensis.
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title: Alteration of photochemistry and protein degradation of photosystem II from Chlamydomonas reinhardtii under high salt grown cells.
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title: Analysis of expression of aquaporins and Na+/H+ transporters in tomato colonized by arbuscular mycorrhizal fungi and affected by salt stress
abstract: Among the proteins functioning in salt tolerance of plants, Na + /H + transporters and aquaporins appear to be of paramount importance. The present study compares expression of Na + /H + transporter and aquaporin genes in tomato colonized by arbuscular mycorrhizal fungi (AMF) and in non-colonized controls under NaCl stress. As revealed by Northern analyses and in situ hybridizations, expression of two Na + /H + transporter genes is not significantly affected by salt stress or by colonization of the plants with AMF. In contrast, transcript levels of both a tonoplast and a plasmalemma aquaporin gene are reduced by salt stress, and this effect is distinctly enhanced by colonization of the tomato roots with AMF. In leaves, colonization of tomato by AMF results in a drastic increase of the mRNA of all three aquaporin genes assayed under salt stress. Aquaporins are known to significantly contribute to water movement in plants. The results presented here indicate that AMF controls aquaporin expression and thereby presumably regulates water flow in tomato under salt stress.

title: The Role of Tobacco Aquaporin1 in Improving Water Use Efficiency, Hydraulic Conductivity, and Yield Production Under Salt Stress1[C][W][OA]
abstract: Tobacco (Nicotiana tabacum; C3) plants increase their water use efficiency (WUE) under abiotic stress and are suggested to show characteristics of C4 photosynthesis in stems, petioles, and transmitting tract cells. The tobacco stress-induced Aquaporin1 (NtAQP1) functions as both water and CO2 channel. In tobacco plants, overexpression of NtAQP1 increases leaf net photosynthesis (AN), mesophyll CO2 conductance, and stomatal conductance, whereas its silencing reduces root hydraulic conductivity (Lp). Nevertheless, interaction between NtAQP1 leaf and root activities and its impact on plant WUE and productivity under normal and stress conditions have never been suggested. Thus, the aim of this study was to suggest a role for NtAQP1 in plant WUE, stress resistance, and productivity. Expressing NtAQP1 in tomato (Solanum lycopersicum) plants (TOM-NtAQP1) resulted in higher stomatal conductance, whole-plant transpiration, and AN under all conditions tested. In contrast to controls, where, under salt stress, Lp decreased more than 3-fold, TOM-NtAQP1 plants, similar to maize (Zea mays; C4) plants, did not reduce Lp dramatically (only by approximately 40%). Reciprocal grafting provided novel evidence for NtAQP1's role in preventing hydraulic failure and maintaining the whole-plant transpiration rate. Our results revealed independent, albeit closely related, NtAQP1 activities in roots and leaves. This dual activity, which increases the plant's water use and AN under optimal and stress conditions, resulted in improved WUE. Consequently, it contributed to the plant's stress resistance in terms of yield production under all tested conditions, as demonstrated in both tomato and Arabidopsis (Arabidopsis thaliana) plants constitutively expressing NtAQP1. The putative involvement of NtAQP1 in tobacco's C4-like photosynthesis characteristics is discussed.

title: Plant aquaporins: membrane channels with multiple integrated functions.
abstract: Aquaporins are channel proteins present in the plasma and intracellular membranes of plant cells, where they facilitate the transport of water and/or small neutral solutes (urea, boric acid, silicic acid) or gases (ammonia, carbon dioxide). Recent progress was made in understanding the molecular bases of aquaporin transport selectivity and gating. The present review examines how a wide range of selectivity profiles and regulation properties allows aquaporins to be integrated in numerous functions, throughout plant development, and during adaptations to variable living conditions. Although they play a central role in water relations of roots, leaves, seeds, and flowers, aquaporins have also been linked to plant mineral nutrition and carbon and nitrogen fixation.

title: Aquaporins in a challenging environment: molecular gears for adjusting plant water status
abstract: Plants have to adjust their water balance in response to very challenging environmental conditions such as drought, salinity, and cold but also changes in light, nutrient deficiency or soil acidity. The molecular and functional characterization of aquaporins, a class of membrane proteins that facilitate water diffusion across cell membranes, has revealed the significance of their regulation in response to these environmental stimuli. The aim of this present review is to illustrate the variety of molecular and cellular mechanisms involved. These mechanisms include the control of aquaporin gene transcription and protein abundance, stimulus-induced aquaporin subcellular relocalization, and channel gating by reversible phosphorylation or by intracellular protons. The emergence of novel mechanisms of regulation by hetero-tetramer formation or through control by reactive oxygen species, and osmotic or hydrostatic pressure gradients is also discussed. These various mechanisms do not function individually and a challenge for future research will be to understand how plants respond to stresses by integrating these mechanisms in time and space, to constantly adjust the water transport and solute transport properties of their membranes. Genetic manipulation of aquaporin functions and in particular ectopic expression of deregulated aquaporins in transgenic plants provide promising strategies to address such questions.

title: Regulation of cation transporter genes by the arbuscular mycorrhizal symbiosis in rice plants subjected to salinity suggests improved salt tolerance due to reduced Na+ root-to-shoot distribution
abstract: Rice is a salt-sensitive crop whose productivity is strongly reduced by salinity around the world. Plants growing in saline soils are subjected to the toxicity of specific ions such as sodium, which damage cell organelles and disrupt metabolism. Plants have evolved biochemical and molecular mechanisms to cope with the negative effects of salinity. These include the regulation of genes with a role in the uptake, transport or compartmentation of Na+ and/or K+. Studies have shown that the arbuscular mycorrhizal (AM) symbiosis alleviates salt stress in several host plant species. However, despite the abundant literature showing mitigation of ionic imbalance by the AM symbiosis, the molecular mechanisms involved are barely explored. The objective of this study was to elucidate the effects of the AM symbiosis on the expression of several well-known rice transporters involved in Na+/K+ homeostasis and measure Na+ and K+ contents and their ratios in different plant tissues. Results showed that OsNHX3, OsSOS1, OsHKT2;1 and OsHKT1;5 genes were considerably upregulated in AM plants under saline conditions as compared to non-AM plants. Results suggest that the AM symbiosis favours Na+ extrusion from the cytoplasm, its sequestration into the vacuole, the unloading of Na+ from the xylem and its recirculation from photosynthetic organs to roots. As a result, there is a decrease of Na+ root-to-shoot distribution and an increase of Na+ accumulation in rice roots which seems to enhance the plant tolerance to salinity and allows AM rice plants to maintain their growing processes under salt conditions.

title: Potassium transport and plant salt tolerance.
abstract: Salinity is a major abiotic stress affecting approximately 7% of the world's total land area resulting in billion dollar losses in crop production around the globe. Recent progress in molecular genetics and plant electrophysiology suggests that the ability of a plant to maintain a high cytosolic K+/Na+ ratio appears to be critical to plant salt tolerance. So far, the major efforts of plant breeders have been aimed at improving this ratio by minimizing Na+ uptake and transport to shoot. In this paper, we discuss an alternative approach, reviewing the molecular and ionic mechanisms contributing to potassium homeostasis in salinized plant tissues and discussing prospects for breeding for salt tolerance by targeting this trait. Major K+ transporters and their functional expression under saline conditions are reviewed and the multiple modes of their control are evaluated, including ameliorative effects of compatible solutes, polyamines and supplemental calcium. Subsequently, the genetic aspects of inheritance of K+ transport 'markers' are discussed in the general context of salt tolerance as a polygenic trait. The molecular identity of 'salt tolerance' genes is analysed, and prospects for future research and breeding are examined.

title: Plant salt-tolerance mechanisms.
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title: Enhanced salt tolerance mediated by AtHKT1 transporter-induced Na unloading from xylem vessels to xylem parenchyma cells.
abstract: AtHKT1 is a sodium (Na+) transporter that functions in mediating tolerance to salt stress. To investigate the membrane targeting of AtHKT1 and its expression at the translational level, antibodies were generated against peptides corresponding to the first pore of AtHKT1. Immunoelectron microscopy studies using anti-AtHKT1 antibodies demonstrate that AtHKT1 is targeted to the plasma membrane in xylem parenchyma cells in leaves. AtHKT1 expression in xylem parenchyma cells was also confirmed by AtHKT1 promoter-GUS reporter gene analyses. Interestingly, AtHKT1 disruption alleles caused large increases in the Na+ content of the xylem sap and conversely reduced the Na+ content of the phloem sap. The athkt1 mutant alleles had a smaller and inverse influence on the potassium (K+) content compared with the Na+ content of the xylem, suggesting that K+ transport may be indirectly affected. The expression of AtHKT1 was modulated not only by the concentrations of Na+ and K+ but also by the osmolality of non-ionic compounds. These findings show that AtHKT1 selectively unloads sodium directly from xylem vessels to xylem parenchyma cells. AtHKT1 mediates osmolality balance between xylem vessels and xylem parenchyma cells under saline conditions. Thus AtHKT1 reduces the sodium content in xylem vessels and leaves, thereby playing a central role in protecting plant leaves from salinity stress.

title: The Na+ transporter AtHKT1;1 controls retrieval of Na+ from the xylem in Arabidopsis.
abstract: HKT-type transporters appear to play key roles in Na(+) accumulation and salt sensitivity in plants. In Arabidopsis HKT1;1 has been proposed to influx Na(+) into roots, recirculate Na(+) in the phloem and control root : shoot allocation of Na(+). We tested these hypotheses using (22)Na(+) flux measurements and ion accumulation assays in an hkt1;1 mutant and demonstrated that AtHKT1;1 contributes to the control of both root accumulation of Na(+) and retrieval of Na(+) from the xylem, but is not involved in root influx or recirculation in the phloem. Mathematical modelling indicated that the effects of the hkt1;1 mutation on root accumulation and xylem retrieval were independent. Although AtHKT1;1 has been implicated in regulation of K(+) transport and the hkt1;1 mutant showed altered net K(+) accumulation, (86)Rb(+) uptake was unaffected by the hkt1;1 mutation. The hkt1;1 mutation has been shown previously to rescue growth of the sos1 mutant on low K(+); however, HKT1;1 knockout did not alter K(+) or (86)Rb(+) accumulation in sos1.

title: Influence of arbuscular mycorrhizae on photosynthesis and water status of maize plants under salt stress
abstract: The influence of arbuscular mycorrhizal (AM) fungus Glomus mosseae on characteristics of the growth, water status, chlorophyll concentration, gas exchange, and chlorophyll fluorescence of maize plants under salt stress was studied in the greenhouse. Maize plants were grown in sand and soil mixture with five NaCl levels (0, 0.5, 1.0, 1.5, and 2.0 g/kg dry substrate) for 55 days, following 15 days of non-saline pretreatment. Under salt stress, mycorrhizal maize plants had higher dry weight of shoot and root, higher relative chlorophyll content, better water status (decreased water saturation deficit, increased water use efficiency, and relative water content), higher gas exchange capacity (increased photosynthetic rate, stomatal conductance and transpiration rate, and decreased intercellular CO2 concentration), higher non-photochemistry efficiency [increased non-photochemical quenching values (NPQ)], and higher photochemistry efficiency [increased the maximum quantum yield in the dark-adapted state (Fv/Fm), the maximum quantum yield in the light-adapted sate (Fv′/Fm′), the actual quantum yield in the light-adapted steady state (ϕPSII) and the photochemical quenching values (qP)], compared with non-mycorrhizal maize plants. In addition, AM symbiosis could trigger the regulation of the energy biturcation between photochemical and non-photochemical events reflected in the deexcitation rate constants (kN, kN′, kP, and kP′). All the results show that G. mosseae alleviates the deleterious effect of salt stress on plant growth, through improving plant water status, chlorophyll concentration, and photosynthetic capacity, while the influence of AM symbiosis on photosynthetic capacity of maize plants can be indirectly affected by soil salinity and mycorrhizae-mediated enhancement of water status, but not by the mycorrhizae-mediated enhancement of chlorophyll concentration and plant biomass.

title: Colonization with arbuscular mycorrhizal fungi improves salinity tolerance of tomato (Solanum lycopersicum L.) plants
abstract: The purpose of this study was to investigate the mechanisms underlying alleviation of salt stress by mycorrhization. Solanum lycopersicum L. cultivars Behta and Piazar with different salinity tolerance were cultivated in soil without salt (EC = 0.63 dSm−1), with low (EC = 5 dSm−1), or high (EC = 10 dSm−1) salinity. Plants inoculated with the arbuscular mycorrhizal fungi Glomus intraradices (+AMF) were compared to non-inoculated plants (−AMF). Under salinity, AMF-mediated growth stimulation was higher in more salt tolerant Piazar than in sensitive Behta. Mycorrhization alleviated salt-induced reduction of P, Ca, and K uptake. Ca/Na and K/Na ratios were also better in +AMF. However, growth improvement by AMF was independent from plant P nutrition under high salinity. Mycorrhization improved the net assimilation rates through both elevating stomatal conductance and protecting photochemical processes of PSII against salinity. Higher activity of ROS scavenging enzymes was concomitant with lowering of H2O2, less lipid peroxidation, and higher proline in +AMF. Cultivar differences in growth responses to salinity and mycorrhization could be well explained by differences in ion balance, photochemistry, and gas exchange of leaves. Function of antioxidant defenses seemed responsible for different AMF-responsiveness of cultivars under salinity. In conclusion, AMF may protect plants against salinity by alleviating the salt-induced oxidative stress.

title: How does arbuscular mycorrhizal symbiosis regulate root hydraulic properties and plasma membrane aquaporins in Phaseolus vulgaris under drought, cold or salinity stresses?
abstract: Here, we evaluated how the arbuscular mycorrhizal (AM) symbiosis regulates root hydraulic properties and root plasma membrane aquaporins (PIP) under different stresses sharing a common osmotic component. Phaseolus vulgaris plants were inoculated or not with the AM fungus Glomus intraradices, and subjected to drought, cold or salinity. Stress effects on root hydraulic conductance (L), PIP gene expression and protein abundance were evaluated. Under control conditions, L in AM plants was about half that in nonAM plants. However, L was decreased as a result of the three stresses in nonAM plants, while it was almost unchanged in AM plants. At the same time, PIP2 protein abundance and phosphorylation state presented the same trend as L. Finally, the expression of each PIP gene responded differently to each stress and was dependent on the AM fungal presence. Differential expression of the PIP genes studied under each stress depending on the AM fungal presence may indicate a specific function and regulation by the AM symbiosis of each gene under the specific conditions of each stress tested.

title: Influence of arbuscular mycorrhiza on photosynthesis and water status of Populus cathayana Rehder males and females under salt stress
abstract: The influence of arbuscular mycorrhizal fungi (AMF) Rhizophagus irregularis on biomass, root morphological characteristics, water status, gas exchange and ion contents of Populus cathayana Rehder males and females under salt stress was studied in a greenhouse experiment. Sex had no influence on AMF colonization. Salt stress decreased the biomass, modified root characteristics, inhibited photosynthesis, relative water content (RWC), while increased the water use efficiency (WUEi), the ratio of root/shoot (RSR) and salt ion contents within the plants. Males and females differed in response to the salt stress and AMF inoculation. AMF inoculation increased the biomass of root, the biomass of shoot, and decreased RSR, especially in males. The root length, root volume (RV), root surface area, and root tip numbers (RTN) were increased with AMF inoculation in males, while RV and RTN were decreased with AMF inoculation in females. AMF had positive effects on RWC and WUEi in both sexes. Mycorrhizal cuttings also had higher gas exchange capacity, lower salt ion contents than non-mycorrhizal cuttings. The principal component analysis showed that males exhibited a better salt tolerance than females, and AMF inoculation had positive effects on both sexes, especially in males.

title: Influence of Salinity on the In Vitro Development of Glomus intraradices and on the In Vivo Physiological and Molecular Responses of Mycorrhizal Lettuce Plants
abstract: Increased salinization of arable land is expected to have devastating global effects in the coming years. Arbuscular mycorrhizal fungi (AMF) have been shown to improve plant tolerance to abiotic environmental factors such as salinity, but they can be themselves negatively affected by salinity. In this study, the first in vitro experiment analyzed the effects of 0, 50, or 100 mM NaCl on the development and sporulation of Glomus intraradices. In the second experiment, the effects of mycorrhization on the expression of key plant genes expected to be affected by salinity was evaluated. Results showed that the assayed isolate G. intraradices DAOM 197198 can be regarded as a moderately salt-tolerant AMF because it did not significantly decrease hyphal development or formation of branching absorbing structures at 50 mM NaCl. Results also showed that plants colonized by G. intraradices grew more than nonmycorrhizal plants. This effect was concomitant with a higher relative water content in AM plants, lower proline content, and expression of Lsp5cs gene (mainly at 50 mM NaCl), lower expression of the stress marker gene Lslea gene, and lower content of abscisic acid in roots of mycorrhizal plants as compared to nonmycorrhizal plants, which suggest that the AM fungus decreased salt stress injury. In addition, under salinity, AM symbiosis enhanced the expression of LsPIP1. Such enhanced gene expression could contribute to regulating root water permeability to better tolerate the osmotic stress generated by salinity.

title: An Expression Analysis of a Gene Family Encoding Plasma Membrane Aquaporins in Response to Abiotic Stresses in Arabidopsis Thaliana
abstract: Aquaporin belongs to a highly conserved group of membrane proteins called major intrinsic proteins that facilitate water transport across biological membranes. The genome of Arabidopsis encodes 35 aquaporin genes with 13 homologs in the plasma membrane intrinsic protein (PIP) subgroup. However, the function of each individual aquaporin isoform and the integrated function of plant aquaporins under various physiological conditions remain unclear. As a step toward understanding the aquaporin function in plants under various environmental stimuli, the expressions of a gene family encoding 13 PIPs in Arabidopsis thalianaunder various abiotic stress conditions including drought, cold, and high salinity, or abscisic acid (ABA) treatment were investigated by a quantitative real-time reverse transcription-PCR analysis. Several PIPgenes were predominantly expressed either in the roots or in the flowers. The expressions of both the highly expressed aquaporins including PIP1;1,PIP1;2,and PIP2;7and the weakly expressed aquaporins such as PIP1;4,PIP2;1,PIP2;4, and PIP2;5were modulated by external stimuli. The analyses of our data revealed that only the PIP2;5was up-regulated by cold treatment, and most of the PIPgenes were down-regulated by cold stress. Marked up- or down-regulation in PIPexpression was observed by drought stress, whereas PIPgenes were less-severely modulated by high salinity. The responsiveness of each aquaporin to ABA were different, implying that the regulation of aquaporin expression involves both ABA-dependent and ABA-independent signaling pathways. Together, our comprehensive expression profile of the 13 members of the PIP gene family provides novel basis to allocate the stress-related biological function to each PIP gene.

title: Transcription factors and regulation of photosynthetic and related metabolism under environmental stresses.
abstract: Environmental conditions, such as water supply, temperature and salinity, strongly affect plant growth and development. Extremes of these conditions (abiotic stresses) adversely affect many different mechanisms associated with plant responses and adaptation to stress: photosynthetic mechanisms, e.g. stomatal control of CO(2) diffusion, photosystem II repair, ribulose bisphosphate carboxylase/oxygenase (Rubisco) activity and scavenging of reactive oxygen species (ROS), are susceptible to damage, and photosynthetic efficiency can be greatly decreased. Responses and adaptations require differential gene expression, which is regulated by specific transcription factors (TFs).



The role and regulation of several TFs involved in abiotic stress response pathways are considered, with emphasis on new findings regarding expression of genes related to both stomatal and non-stomatal limitations to CO(2) photosynthetic assimilation.



Many TFs, belonging to different families (e.g. MYB, bZIP and DREB), have been related to abiotic stress responses; however, only a few are known to regulate the expression of photosynthesis-related genes in response to stress. Several TFs belonging to the MYB family play an important role in both stomatal and non-stomatal responses by regulation of stomatal numbers and sizes, and metabolic components, respectively. To obtain more insight into this area of potentially large agronomic impact, it is essential to identify and functionally characterize new TFs that mediate the stress responses regulating the expression of genes associated with photosynthesis and related metabolism.

title: The action of aquaporins in cell elongation, salt stress and photosynthesis.
abstract: Aquaporin belongs to a highly conserved group of membrane proteins called major intrinsic proteins (MIPs) that facilitate water transport across biological membranes. Aquaporins are membrane water channels that play critical roles in controlling the water content of cells and tissues. We focused on GhPIP1;2 which belongs to the PIP subfamily and GhgammaTIP1 which belongs to the gammaTIP group of the TIP subfamily. Northern blot analysis with gene-specific probes and real-time PCR demonstrated that GhPIP1;2 and GhgammaTIP1 are predominantly expressed during cotton fiber elongation, with the highest expression levels at 5 days post anthesis. The high and preferential expression of GhPIP1;2 and GhgammaTIP1 suggests that they may play important roles in supporting the rapid influx of water into vacuoles during cotton fiber cell expansion. Also, the effects of Ca2+ on aquaporins in salinity-stressed plants were studied. Researchers treated the protoplasts and plasma membrane with NaCl or CaCl2, alone or in combination. Under saline conditions, osmotic water permeability (Pf) values decreased in protoplasts and plasma membrane vesicles, and the same reduction was observed in the PIP1 aquaporin abundance, indicating inhibitory effects of NaCl on aquaporin functionality and protein abundance. Two different actions of Ca2+ were observed. Increase in free cytosolic calcium concentrations associated with stress perception may lead to aquaporin closure, however, the extra-calcium would lead to an upregulation of aquaporins. Meanwhile, experiments have demonstrated HvPIP2;1, one of barley aquaporins, has a higher water and CO2 transport activity. The goal of our plant aquaporin research is to determine the key aquaporin species responsible for water and CO2 transport, and to improve plant water relations, stress tolerance, CO2 uptake or assimilation, and plant productivity.

title: Contribution of Glomus intraradices inoculation to nutrient acquisition and mitigation of ionic imbalance in NaCl-stressed Trigonella foenum-graecum
abstract: The study aimed to investigate the effects of an AM fungus (Glomus intraradices Schenck and Smith) on mineral acquisition in fenugreek (Trigonella foenum-graecum) plants under different levels of salinity. Mycorrhizal (M) and non-mycorrhizal (NM) fenugreek plants were subjected to four levels of NaCl salinity (0, 50, 100, and 200 mM NaCl). Plant tissues were analyzed for different mineral nutrients. Leaf senescence (chlorophyll concentration and membrane permeability) and lipid peroxidation were also assessed. Under salt stress, M plants showed better growth, lower leaf senescence, and decreased lipid peroxidation as compared to NM plants. Salt stress adversely affected root nodulation and uptake of NPK. This effect was attenuated in mycorrhizal plants. Presence of the AM fungus prevented excess uptake of Na+ with increase in NaCl in the soil. It also imparted a regulatory effect on the translocation of Na+ ions to shoots thereby maintaining lower Na+ shoot:root ratios as compared to NM plants. Mycorrhizal colonization helped the host plant to overcome Na+-induced Ca2+ and K+ deficiencies. M plants maintained favorable K+:Na+, Ca2+:Na+, and Ca2+:Mg2+ ratios in their tissues. Concentrations of Cu, Fe, and Zn2+ decreased with increase in intensity of salinity stress. However, at each NaCl level, M plants had higher concentration of Cu, Fe, Mn2+, and Zn2+ as compared to NM plants. M plants showed reduced electrolyte leakage in leaves as compared to NM plants. The study suggests that AM fungi contribute to alleviation of salt stress by mitigation of NaCl-induced ionic imbalance thus maintaining a favorable nutrient profile and integrity of the plasma membrane.

title: Physiological Responses and Gene Co-Expression Network of Mycorrhizal Roots under K+ Deprivation1[OPEN]
abstract: Arbuscular mycorrhizal symbiosis compensates the transcriptional response of M. truncatula roots at low potassium level and activates specific mechanisms to tolerate long-term potassium deprivation. Arbuscular mycorrhizal (AM) associations enhance the phosphorous and nitrogen nutrition of host plants, but little is known about their role in potassium (K+) nutrition. Medicago truncatula plants were cocultured with the AM fungus Rhizophagus irregularis under high and low K+ regimes for 6 weeks. We determined how K+ deprivation affects plant development and mineral acquisition and how these negative effects are tempered by the AM colonization. The transcriptional response of AM roots under K+ deficiency was analyzed by whole-genome RNA sequencing. K+ deprivation decreased root biomass and external K+ uptake and modulated oxidative stress gene expression in M. truncatula roots. AM colonization induced specific transcriptional responses to K+ deprivation that seem to temper these negative effects. A gene network analysis revealed putative key regulators of these responses. This study confirmed that AM associations provide some tolerance to K+ deprivation to host plants, revealed that AM symbiosis modulates the expression of specific root genes to cope with this nutrient stress, and identified putative regulators participating in these tolerance mechanisms.

title: Arbuscular mycorrhizal fungi native from a Mediterranean saline area enhance maize tolerance to salinity through improved ion homeostasis.
abstract: Soil salinity restricts plant growth and productivity. Na(+) represents the major ion causing toxicity because it competes with K(+) for binding sites at the plasma membrane. Inoculation with arbuscular mycorrhizal fungi (AMF) can alleviate salt stress in the host plant through several mechanisms. These may include ion selection during the fungal uptake of nutrients from the soil or during transfer to the host plant. AM benefits could be enhanced when native AMF isolates are used. Thus, we investigated whether native AMF isolated from an area with problems of salinity and desertification can help maize plants to overcome the negative effects of salinity stress better than non-AM plants or plants inoculated with non-native AMF. Results showed that plants inoculated with two out the three native AMF had the highest shoot dry biomass at all salinity levels. Plants inoculated with the three native AMF showed significant increase of K(+) and reduced Na(+) accumulation as compared to non-mycorrhizal plants, concomitantly with higher K(+) /Na(+) ratios in their tissues. For the first time, these effects have been correlated with regulation of ZmAKT2, ZmSOS1 and ZmSKOR genes expression in the roots of maize, contributing to K(+) and Na(+) homeostasis in plants colonized by native AMF.

title: Effect of inoculation with ecto- and arbuscular mycorrhizae and Rhizobium on the growth and nitrogen fixation by black locust, Robinia pseudoacacia
abstract: Experiments were made to determine the effect of inoculation withRhizobium, the ectomycorrhizal fungus Hebelomamesophasem and the arbuscular mycorrhizal fungus Glomuscaledonium on the growth and nitrogen fixation of black locust(Robinia pseudoacacia Linn.) seedlings grown invitro and in pot culture. The results showed that both mycorrhizalfungi and Rhizobium stimulated the growth and nitrogenfixing ability of inoculated seedlings. Inoculation with all three microbestogether produced the most beneficial effects on nitrogen fixation, mycorrhizaldevelopment and seedling growth.

title: Does ectomycorrhizal fungal community structure vary along a Japanese black pine (Pinus thunbergii) to black locust (Robinia pseudoacacia) gradient?
abstract: In this study we examined the role of the nitrogen-fixing tree, Robinia pseudoacacia (black locust), in ectomycorrhizal (ECM) formation and ECM community of Pinus thunbergii (Japanese black pine) seedlings. Two 200 m(2) experimental plots were established at the border between a Japanese black pine- and a black locust-dominated area in a coastal forest. The ECM fungal community of pine seedlings was examined by PCR-RFLP and sequence analysis. We analyzed the relationship between ECM formation, ECM community, growth, and nutrient status of pine seedlings and environmental conditions using the Mantel test and structural equation model. Percentages of ECM root tips, the number of ECM fungal species and ECM diversity on pine seedlings decreased in the black locust-dominated area. Cenococcum geophilum and Russula spp. were dominant in the Japanese black pine-dominated area, whereas Tomentella spp. were dominant in the black locust-dominated area. Nitrogen (N) concentration in soils or pine seedlings strongly influenced the percentage of ECM root tips, the number of ECM fungal species and ECM fungal similarity. These results imply the long-term eutrophication caused by N-fixing trees can change ECM formation and ECM community structure.

title: Assessing adaptability of planted trees using leaf traits: A case study with Robinia pseudoacacia L. in the Loess Plateau, China
abstract: Leaf trait patterns and their variations with climate are interpreted as an adaptive adjustment to environment. This study assessed the adaptability of planted black locust (Robinia pseudoacacia L.) based on the analysis of leaf traits and the comparison of its leaf traits with inter-specific ones existing in the same area. We measured some water and N use related leaf traits: leaf dry mass per unit area (LMA) and N, P and K concentrations based on both leaf area (Narea, Parea and Karea) and leaf mass (Nmass, Pmass and Kmass) of R. pseudoacacia at 31 sites along a water stress gradient in North Shaanxi Province, China. The results show that leaves of R. pseudoacacia have high Nmass and low LMA in the study area. High Nmass and low LMA are usually representative of luxurious resource use, and will advance plant resource competitiveness in high-resource conditions. As a whole, LMA-nutrient relationships of R. pseudoacacia display patterns that are fairly similar to the inter-specific relationships in both direction and intensity. The tendency for LMA and Narea to increase with decreasing water availability and the positive correlation between LMA and Narea reflect the trend for R. pseudoacacia to enhance water use efficiency (WUE) at the expense of down-regulated photosynthetic N use efficiency (PNUE) and high construction cost in dry conditions. However, the positive relationship between LMA and Narea in high mean annual precipitation (MAP) area is either unremarkable or reversed with decreasing water availability. This implies a lower photosynthetic capacity and a higher construction cost for high-LMA leaves. The inter-specific relationship between LMA and Narea is positive and does not change with water availability. This difference between inter-species and intra-species may be due to more diversified anatomies and more specialised structures for inter-species than intra-species. The failure of R. pseudoacacia adaption to dry conditions reflected by LMA-Narea relationship may be partially responsible for the emergence of rampike and dwarf forms found frequently in dry conditions. Incorporating intrinsic characteristics of planted trees into vegetation restoration project will be instructive and meaningful for species selection.

title: Aquaporin gene expression and physiological responses of Robinia pseudoacacia L. to the mycorrhizal fungus Rhizophagus irregularis and drought stress
abstract: The influence of arbuscular mycorrhiza (AM) and drought stress on aquaporin (AQP) gene expression, water status, and photosynthesis was investigated in black locust (Robinia pseudoacacia L.). Seedlings were grown in potted soil inoculated without or with the AM fungus Rhizophagus irregularis, under well-watered and drought stress conditions. Six full-length AQP complementary DNAs (cDNAs) were isolated from Robinia pseudoacacia, named RpTIP1;1, RpTIP1;3, RpTIP2;1, RpPIP1;1, RpPIP1;3, and RpPIP2;1. A phylogenetic analysis of deduced amino acid sequences demonstrated that putative proteins coded by these RpAQP genes belong to the water channel protein family. Expression analysis revealed higher RpPIP expression in roots while RpTIP expression was higher in leaves, except for RpTIP1;3. AM symbiosis regulated host plant AQPs, and the expression of RpAQP genes in mycorrhizal plants depended on soil water condition and plant tissue. Positive effects were observed for plant physiological parameters in AM plants, which had higher dry mass and lower water saturation deficit and electrolyte leakage than non-AM plants. Rhizophagus irregularis inoculation also slightly increased leaf net photosynthetic rate and stomatal conductance under well-watered and drought stress conditions. These findings suggest that AM symbiosis can enhance the drought tolerance in Robinia pseudoacacia plants by regulating the expression of RpAQP genes, and by improving plant biomass, tissue water status, and leaf photosynthesis in host seedlings.

title: Improved procedures for clearing roots and staining parasitic and vesicular-arbuscular mycorrhizal fungi for rapid assessment of infection.
abstract: During studies on the effect of vesicular-arbuscular (VA) mycorrhiza, caused by species of Endogene, on plant growth, we needed to estimate the amount of mycorrhizal infection in the root cortex. Such infections in gramineous roots stain fairly clearly with standard procedures such as simmering in o· I % trypan blue in lactophenol (Hayman, 1970), or cotton blue and Sudan IV in lactophenol (Nicolson, 1959), and in some dicotyledonous roots by boiling for 10 min in 0'0 I % acid fuchsin in saturated chloral hydrate (Gerdemann, 1955), or standing cold overnight in any of these stains. Infection in onion, our main test plant (Hayman & Mosse, 1970), is difficult to estimate using these methods because, although external hyphae and entry points are strongly stained, internal hyphae, arbuscules and vesicles are usually only well defined near the cut ends of the root segments. To improve stain penetration and clearing in whole mycorrhizal roots of onion and other host plants, and in roots infected by other fungi, we developed the following two procedures, which give deeply stained fungal structures which show distinctly against the outlines of the cells in the cortex of intact roots. The first procedure was used successfully for non-pigmented roots such as those of onion. Root segments or whole roots, fresh or fixed in FAA ( I 3 ml formalin, 5 ml glacial acetic acid, 200 ml 50 % ethanol), were heated at 90°C for about I h in 10 % KOH. This removed the host cytoplasm and most of the nuclei, and the roots became very clear with

title: AN EVALUATION OF TECHNIQUES FOR MEASURING VESICULAR ARBUSCULAR MYCORRHIZAL INFECTION IN ROOTS
abstract: Assessment of infection is an essential part of many studies involving VA mycorrhiza. A summary is given of the range of techniques that have been used. We calculated the standard error of four methods of assessment based on observations of stained root samples either randomly arranged in a petri dish or mounted on microscope slides. The methods are based on presence or absence of infection at root/grid intersect points, on a visual estimate of percentage cortex occupied by fungus or on estimates of length, or presence or absence of infection in root pieces mounted on slides. The number of replicate observations required for a given standard error % infection can be read from the curves provided. The advantages of the different methods of assessment are discussed and reasons given why they all probably overestimate the true values.

title: Chlorophyll fluorescence: a probe of photosynthesis in vivo.
abstract: The use of chlorophyll fluorescence to monitor photosynthetic performance in algae and plants is now widespread. This review examines how fluorescence parameters can be used to evaluate changes in photosystem II (PSII) photochemistry, linear electron flux, and CO(2) assimilation in vivo, and outlines the theoretical bases for the use of specific fluorescence parameters. Although fluorescence parameters can be measured easily, many potential problems may arise when they are applied to predict changes in photosynthetic performance. In particular, consideration is given to problems associated with accurate estimation of the PSII operating efficiency measured by fluorescence and its relationship with the rates of linear electron flux and CO(2) assimilation. The roles of photochemical and nonphotochemical quenching in the determination of changes in PSII operating efficiency are examined. Finally, applications of fluorescence imaging to studies of photosynthetic heterogeneity and the rapid screening of large numbers of plants for perturbations in photosynthesis and associated metabolism are considered.

title: MEGA5: molecular evolutionary genetics analysis using maximum likelihood, evolutionary distance, and maximum parsimony methods.
abstract: Comparative analysis of molecular sequence data is essential for reconstructing the evolutionary histories of species and inferring the nature and extent of selective forces shaping the evolution of genes and species. Here, we announce the release of Molecular Evolutionary Genetics Analysis version 5 (MEGA5), which is a user-friendly software for mining online databases, building sequence alignments and phylogenetic trees, and using methods of evolutionary bioinformatics in basic biology, biomedicine, and evolution. The newest addition in MEGA5 is a collection of maximum likelihood (ML) analyses for inferring evolutionary trees, selecting best-fit substitution models (nucleotide or amino acid), inferring ancestral states and sequences (along with probabilities), and estimating evolutionary rates site-by-site. In computer simulation analyses, ML tree inference algorithms in MEGA5 compared favorably with other software packages in terms of computational efficiency and the accuracy of the estimates of phylogenetic trees, substitution parameters, and rate variation among sites. The MEGA user interface has now been enhanced to be activity driven to make it easier for the use of both beginners and experienced scientists. This version of MEGA is intended for the Windows platform, and it has been configured for effective use on Mac OS X and Linux desktops. It is available free of charge from http://www.megasoftware.net.

title: Soil salinity delays germination and limits growth of hyphae from propagules of arbuscular mycorrhizal fungi
abstract: Colonisation of plant roots by some arbuscular mycorrhizal (AM) fungi is reduced in the presence of sodium chloride (NaCl), probably due to a direct effect of NaCl on the fungi. However, there appear to be differences between the fungi in their ability to colonise plants in the presence of NaCl. This experiment tested the hypothesis that propagules of different isolates and species of AM fungi from saline and nonsaline soils would differ in their ability to germinate and grow in the presence of NaCl in the soil solution. Spores or pieces of root colonised by a range of AM fungi were incubated between filters buried in soil to which NaCl had been added at concentrations of 0, 150 or 300 mM in the soil solution. At regular intervals, filters were removed from the soil and both the percentage of propagules which had germinated and the length of proliferating hyphae were determined. Germination of spores of AM fungi studied was delayed in the presence of NaCl, but the fungi differed in the extent to which germination was inhibited. Two isolates of Scutellospora calospora reached maximum germination in 300 mM NaCl, but neither of two isolates of Acaulospora laevis germinated in the presence of NaCl. Germination of spores of the other fungi, including some isolated from saline soil, fell between these extremes. For some fungi, the specific rate of hyphal extension was reduced by NaCl. For others, the specific rate of growth was similar in the presence of NaCl to that in the control treatment, but overall production of hyphae was reduced in the NaCl treatments because germination was reduced.

title: Arbuscular mycorrhizal symbiosis modulates antioxidant response in salt-stressed Trigonella foenum-graecum plants
abstract: An experiment was conducted to evaluate the influence of Glomus intraradices colonization on the activity of antioxidant enzymes [superoxide dismutase (SOD), catalase (CAT), peroxidase (PX), ascorbate peroxidase (APX), and glutathione reductase (GR)] and the accumulation of nonenzymatic antioxidants (ascorbic acid, α-tocopherol, glutathione, and carotenoids) in roots and leaves of fenugreek plants subjected to varying degrees of salinity (0, 50, 100, and 200 mM NaCl) at two time intervals (1 and 14 days after saline treatment, DAT). The antioxidative capacity was correlated with oxidative damage in the same tissue. Under salt stress, lipid peroxidation and H2O2 concentration increased with increasing severity and duration of salt stress (DoS). However, the extent of oxidative damage in mycorrhizal plants was less compared to nonmycorrhizal plants. The study reveals that mycorrhiza-mediated attenuation of oxidative stress in fenugreek plants is due to enhanced activity of antioxidant enzymes and higher concentrations of antioxidant molecules. However, the significant effect of G. intraradices colonization on individual antioxidant molecules and enzymes varied with plant tissue, salinity level, and DoS. The significant effect of G. intraradices colonization on antioxidative enzymes was more evident at 1DAT in both leaves and roots, while the concentrations of antioxidant molecules were significantly influenced at 14DAT. It is proposed that AM symbiosis can improve antioxidative defense systems of plants through higher SOD activity in M plants, facilitating rapid dismutation of O2- to H2O2, and subsequent prevention of H2O2 build-up by higher activities of CAT, APX, and PX. The potential of G. intraradices to ameliorate oxidative stress generated in fenugreek plants by salinity was more evident at higher intensities of salt stress.

title: Regulation by arbuscular mycorrhizae of the integrated physiological response to salinity in plants: new challenges in physiological and molecular studies.
abstract: Excessive salt accumulation in soils is a major ecological and agronomical problem, in particular in arid and semi-arid areas. Excessive soil salinity affects the establishment, development, and growth of plants, resulting in important losses in productivity. Plants have evolved biochemical and molecular mechanisms that may act in a concerted manner and constitute the integrated physiological response to soil salinity. These include the synthesis and accumulation of compatible solutes to avoid cell dehydration and maintain root water uptake, the regulation of ion homeostasis to control ion uptake by roots, compartmentation and transport into shoots, the fine regulation of water uptake and distribution to plant tissues by the action of aquaporins, the reduction of oxidative damage through improved antioxidant capacity and the maintenance of photosynthesis at values adequate for plant growth. Arbuscular mycorrhizal (AM) symbiosis can help the host plants to cope with the detrimental effects of high soil salinity. There is evidence that AM symbiosis affects and regulates several of the above mentioned mechanisms, but the molecular bases of such effects are almost completely unknown. This review summarizes current knowledge about the effects of AM symbiosis on these physiological mechanisms, emphasizing new perspectives and challenges in physiological and molecular studies on salt-stress alleviation by AM symbiosis.

title: Influence of arbuscular mycorrhiza on organic solutes in maize leaves under salt stress
abstract: A pot experiment was conducted to examine the effect of the arbuscular mycorrhizal (AM) fungus, Glomus mosseae, on plant biomass and organic solute accumulation in maize leaves. Maize plants were grown in sand and soil mixture with three NaCl levels (0, 0.5, and 1.0 g kg−1 dry substrate) for 55 days, after 15 days of establishment under non-saline conditions. At all salinity levels, mycorrhizal plants had higher biomass and higher accumulation of organic solutes in leaves, which were dominated by soluble sugars, reducing sugars, soluble protein, and organic acids in both mycorrhizal and non-mycorrhizal plants. The relative abundance of free amino acids and proline in total organic solutes was lower in mycorrhizal than in non-mycorrhizal plants, while that of reducing sugars was higher. In addition, the AM symbiosis raised the concentrations of soluble sugars, reducing sugars, soluble protein, total organic acids, oxalic acid, fumaric acid, acetic acid, malic acid, and citric acid and decreased the concentrations of total free amino acids, proline, formic acid, and succinic acid in maize leaves. In mycorrhizal plants, the dominant organic acid was oxalic acid, while in non-mycorrhizal plants, the dominant organic acid was succinic acid. All the results presented here indicate that the accumulation of organic solutes in leaves is a specific physiological response of maize plants to the AM symbiosis, which could mitigate the negative impact of soil salinity on plant productivity.

title: Arbuscular mycorrhizal symbiosis influences strigolactone production under salinity and alleviates salt stress in lettuce plants.
abstract: Arbuscular mycorrhizal (AM) symbiosis can alleviate salt stress in plants. However the intimate mechanisms involved, as well as the effect of salinity on the production of signalling molecules associated to the host plant-AM fungus interaction remains largely unknown. In the present work, we have investigated the effects of salinity on lettuce plant performance and production of strigolactones, and assessed its influence on mycorrhizal root colonization. Three different salt concentrations were applied to mycorrhizal and non-mycorrhizal plants, and their effects, over time, analyzed. Plant biomass, stomatal conductance, efficiency of photosystem II, as well as ABA content and strigolactone production were assessed. The expression of ABA biosynthesis genes was also analyzed. AM plants showed improved growth rates and a better performance of physiological parameters such as stomatal conductance and efficiency of photosystem II than non-mycorrhizal plants under salt stress since very early stages - 3 weeks - of plant colonization. Moreover, ABA levels were lower in those plants, suggesting that they were less stressed than non-colonized plants. On the other hand, we show that both AM symbiosis and salinity influence strigolactone production, although in a different way in AM and non-AM plants. The results suggest that AM symbiosis alleviates salt stress by altering the hormonal profiles and affecting plant physiology in the host plant. Moreover, a correlation between strigolactone production, ABA content, AM root colonization and salinity level is shown. We propose here that under these unfavourable conditions, plants increase strigolactone production in order to promote symbiosis establishment to cope with salt stress.

title: Effects of salt stress on basic processes of photosynthesis
abstract: Salt stress causes decrease in plant growth and productivity by disrupting physiological processes, especially photosynthesis. The accumulation of intracellular sodium ions at salt stress changes the ratio of K : Na, which seems to affect the bioenergetic processes of photosynthesis. Both multiple inhibitory effects of salt stress on photosynthesis and possible salt stress tolerance mechanisms in cyanobacteria and plants are reviewed.

title: Arbuscular mycorrhizas influence Lycium barbarum tolerance of water stress in a hot environment
abstract: Arbuscular mycorrhizal (AM) fungi can assist their hosts to cope with water stress and other abiotic stresses in different ways. In order to test whether AM plants have a greater capacity than control plants to cope with water stress, we investigated the water status and photosynthetic capacity of Lycium barbarum colonized or not by the AM fungus Rhizophagus irregularis under three water conditions during a hot summer. Sugar levels and transcriptional responses of both plant and AM fungus aquaporin genes in roots were analyzed. Compared with control plants, AM plants increased transpiration rate and stomatal conductance but decreased leaf relative water content under moderate water stress. Severe water stress, however, did not inhibit the quantum yield of PSII photochemistry in AM plants versus control plants. AM plants had higher expression levels of plasma membrane intrinsic proteins or tonoplast intrinsic proteins and Rir-AQP2 and lower leaf temperature than control plants under dry-hot stress. Additionally, AM plant sugar levels under normal water conditions were similar to those of control plants under moderate water stress, but sugar levels of AM plants especially increased with severe water stress. When these aspects of performance of AM and control plants under different water conditions are compared overall, AM plants displayed an obvious superiority over control plants at coping with moderate water stress in the hot environment; AM plants maintained normal photochemical processes under severe water stress, while sugar levels were affected strongly.

title: New insights into the regulation of aquaporins by the arbuscular mycorrhizal symbiosis in maize plants under drought stress and possible implications for plant performance.
abstract: The relationship between modulation by arbuscular mycorrhizae (AM) of aquaporin expression in the host plant and changes in root hydraulic conductance, plant water status, and performance under stressful conditions is not well known. This investigation aimed to elucidate how the AM symbiosis modulates the expression of the whole set of aquaporin genes in maize plants under different growing and drought stress conditions, as well as to characterize some of these aquaporins in order to shed further light on the molecules that may be involved in the mycorrhizal responses to drought. The AM symbiosis regulated a wide number of aquaporins in the host plant, comprising members of the different aquaporin subfamilies. The regulation of these genes depends on the watering conditions and the severity of the drought stress imposed. Some of these aquaporins can transport water and also other molecules which are of physiological importance for plant performance. AM plants grew and developed better than non-AM plants under the different conditions assayed. Thus, for the first time, this study relates the well-known better performance of AM plants under drought stress to not only the water movement in their tissues but also the mobilization of N compounds, glycerol, signaling molecules, or metalloids with a role in abiotic stress tolerance. Future studies should elucidate the specific function of each aquaporin isoform regulated by the AM symbiosis in order to shed further light on how the symbiosis alters the plant fitness under stressful conditions.

title: The tobacco aquaporin NtAQP1 is a membrane CO2 pore with physiological functions
abstract: Aquaporins, found in virtually all living organisms, are membrane-intrinsic proteins that form water-permeable complexes. The mammalian aquaporin AQP1 has also shown CO2 permeability when expressed heterologously in Xenopus oocytes, although whether this is a biochemical curiosity or of physiological significance is a matter of debate. Here we report that, in the same expression system, a CO2 permeability comparable to that of the human AQP1 is observed for the tobacco plasma membrane aquaporin NtAQP1. NtAQP1 facilitates CO2 membrane transport in the homologous plant system at the cellular level, and has a significant function in photosynthesis and in stomatal opening. NtAQP1 overexpression heightens membrane permeability for CO2 and water, and increases leaf growth. The results indicate that NtAQP1-related CO2 permeability is of physiological importance under conditions where the CO2 gradient across a membrane is small, as is the case between the atmosphere and the inside of a plant cell.

title: Effects of HgCl(2) on CO(2) dependence of leaf photosynthesis: evidence indicating involvement of aquaporins in CO(2) diffusion across the plasma membrane.
abstract: Experiments were conducted to examine whether mercury-sensitive aquaporins facilitate photosynthetic CO(2) diffusion across the plasma membrane of leaf mesophyll cells. Discs without abaxial epidermes from Vicia faba leaflets were treated with HgCl(2), an inhibitor of aquaporins. Hydraulic conductivity of the plasma membrane of these discs, measured as the weight loss of the discs in the 1 M sorbitol solution, was inhibited by sub-mM concentrations of HgCl(2) by 70 to 80%. Photosynthetic CO(2) fixation was also inhibited by the HgCl(2) treatment in a similar concentration range. When 0.3 mM HgCl(2) solution was fed to the V. faba leaflets with intact epidermes via the transpiration stream, the rate of photosynthesis on leaf area basis (A) measured at photosynthetically active photon flux density of 700 micromol m(-2) s(-1) and at leaf temperature of 25 degrees C, decreased by about 20 to 30% at any CO(2) concentration in the intercellular spaces (C(i)). However, when CO(2) concentration in the chloroplast stroma (C(c)) was calculated from fluorescence and gas exchange data and A was plotted against C(c), A at low C(c) concentrations did not differ before and after the treatment. The conductance for CO(2) diffusion from the intercellular spaces to the chloroplast stroma (g(i)) decreased to 40 and 30% of the control value, when the leaflets were fed with 0.3 mM and 1.2 mM HgCl(2), respectively. Similar results were obtained with leaves of Phaseolus vulgaris. Although effects of HgCl(2) were not specific, the present results showed that HgCl(2) consistently lowered g(i). It is, thus, probable that the photosynthetic CO(2) uptake across the plasma membrane of the mesophyll cells is facilitated by mercury-sensitive aquaporins.

title: Influence of arbuscular mycorrhizae on yield, nutrients, organic solutes, and antioxidant enzymes of two wheat cultivars under salt stress
abstract: The efficacy of arbuscular mycorrhizae (AM) on nutrients, organic solutes, and antioxidant enzymes of wheat under salt stress was investigated and related to root colonization and plant productivity. The mycorrhizal inoculation increased N, P, K, Ca, and Mg uptake, soluble sugars, free amino acids, and proline accumulation, as well as peroxidase and catalase activities under saline conditions as compared to nonmycorrhizal plants. On the other hand, Na concentration was lower in mycorrhizal than in nonmycorrhizal plants grown under saline conditions. Arbuscular mycorrhizae protected wheat against the detrimental effects of salinity and stimulated its productivity. Hence, mycorrhizal colonization can play a vital role in the mitigation of the adverse effects of salinity by improving the wheat osmotic adjustment response, enhancing its defense system, and alleviating oxidative damage to cells. Arbuscular mycorrhizae are able to alter plant physiology in a way that empowers the plant to grow more efficiently on salt-affected lands.

title: Elemental composition of arbuscular mycorrhizal fungi at high salinity
abstract: We investigated the elemental composition of spores and hyphae of arbuscular mycorrhizal fungi (AMF) collected from two saline sites at the desert border in Tunisia, and of Glomus intraradices grown in vitro with or without addition of NaCl to the medium, by proton-induced X-ray emission. We compared the elemental composition of the field AMF to those of the soil and the associated plants. The spores and hyphae from the saline soils showed strongly elevated levels of Ca, Cl, Mg, Fe, Si, and K compared to their growth environment. In contrast, the spores of both the field-derived AMF and the in vitro grown G. intraradices contained lower or not elevated Na levels compared to their growth environment. This resulted in higher K:Na and Ca:Na ratios in spores than in soil, but lower than in the associated plants for the field AMF. Na and Ca:Na ratios of G. intraradices grown in monoxenic cultures were also in the same range as those of the field AMF and did not change even when those ratios in the growth medium were lowered several orders of magnitude by adding NaCl. These results indicate that AMF can selectively take up elements such as K and Ca, which act as osmotic equivalents while they avoid uptake of toxic Na. This could make them important in the alleviation of salinity stress in their plant hosts.

title: Potassium transport and signaling in higher plants.
abstract: As one of the most important mineral nutrient elements, potassium (K(+)) participates in many plant physiological processes and determines the yield and quality of crop production. In this review, we summarize K(+) signaling processes and K(+) transport regulation in higher plants, especially in plant responses to K(+)-deficiency stress. Plants perceive external K(+) fluctuations and generate the initial K(+) signal in root cells. This signal is transduced into the cytoplasm and encoded as Ca(2+) and reactive oxygen species signaling. K(+)-deficiency-induced signals are subsequently decoded by cytoplasmic sensors, which regulate the downstream transcriptional and posttranslational responses. Eventually, plants produce a series of adaptive events in both physiological and morphological alterations that help them survive K(+) deficiency.

title: The Putative Plasma Membrane Na+/H+ Antiporter SOS1 Controls Long-Distance Na+ Transport in Plants Article, publication date, and citation information can be found at www.plantcell.org/cgi/doi/10.1105/tpc.010371.
abstract: The salt tolerance locus SOS1 from Arabidopsis has been shown to encode a putative plasma membrane Na+/H+ antiporter. In this study, we examined the tissue-specific pattern of gene expression as well as the Na+ transport activity and subcellular localization of SOS1. When expressed in a yeast mutant deficient in endogenous Na+ transporters, SOS1 was able to reduce Na+ accumulation and improve salt tolerance of the mutant cells. Confocal imaging of a SOS1–green fluorescent protein fusion protein in transgenic Arabidopsis plants indicated that SOS1 is localized in the plasma membrane. Analysis of SOS1 promoter–β-glucuronidase transgenic Arabidopsis plants revealed preferential expression of SOS1 in epidermal cells at the root tip and in parenchyma cells at the xylem/symplast boundary of roots, stems, and leaves. Under mild salt stress (25 mM NaCl), sos1 mutant shoot accumulated less Na+ than did the wild-type shoot. However, under severe salt stress (100 mM NaCl), sos1 mutant plants accumulated more Na+ than did the wild type. There also was greater Na+ content in the xylem sap of sos1 mutant plants exposed to 100 mM NaCl. These results suggest that SOS1 is critical for controlling long-distance Na+ transport from root to shoot. We present a model in which SOS1 functions in retrieving Na+ from the xylem stream under severe salt stress, whereas under mild salt stress it may function in loading Na+ into the xylem.