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Salt cress (Eutrema salsugineum) presents relatively high phosphate (Pi) use efficiency cy in its natural habitat. Phosphate Transporters (PHTs) play critical roles in Pi acquisition and homeostasis. Here, a comparative study of PHT families between salt cress and Arabidopsis was performed. A total of 27 putative PHT genes were identified in E. salsugineum genome. Notably, seven tandem genes encoding PHT1;3 were found, and function analysis in Arabidopsis indicated at least six EsPHT1;3s participated in Pi uptake. Meanwhile, different expression profiles of PHT genes between the two species under Pi limitation and salt stress were documented. Most PHT1 genes were down-regulated in Arabidopsis while up-regulated in salt cress under salinity, among which EsPHT1;9 was further characterized. EsPHT1;9 was involved in root-to-shoot Pi translocation. Particularly, the promoter of EsPHT1;9 outperformed that of AtPHT1;9 in promoting Pi translocation, K+ /Na+ ratio, thereby salt tolerance. Through cis-element analysis, we identified a bZIP transcription factor EsABF5 negatively regulating EsPHT1;9 and plant tolerance to low-Pi and salt stress. Altogether, more copies and divergent transcriptional regulation of PHT genes contribute to salt cress adaptation to the co-occurrence of salinity and Pi limitation, which add our knowledge on the evolutionary and molecular component of multistress- tolerance of this species.

Salinity-induced lipid alterations have been reported in many plant species; however, how lipid biosynthesis and metabolism are regulated and how lipids work in plant salt tolerance are much less studied. Here, a constitutively much higher phosphatidylserine (PS) content in the plasma membrane (PM) was found in the euhalophyte Salicornia europaea than in Arabidopsis. A gene encoding PS synthase (PSS) was subsequently isolated from S. europaea, named SePSS, which was induced by salinity. Multiple alignments and phylogenetic analysis suggested that SePSS belongs to a base exchange-type PSS, which localises to the endoplasmic reticulum. Knockdown of SePSS in S. europaea suspension cells resulted in reduced PS content, decreased cell survival rate, and increased PM depolarization and K+ efflux under 400 or 800 mM NaCl. By contrast, the upregulation of SePSS leads to increased PS and phosphatidylethanolamine levels and enhanced salt tolerance in Arabidopsis, along with a lower accumulation of reactive oxygen species, less membrane injury, less PM depolarization and higher K+/Na+ in the transgenic lines than in wild-type (WT). These results suggest a positive correlation between PS levels and plant salt tolerance, and that SePSS participates in plant salt tolerance by regulating PS levels, hence PM potential and permeability, which help maintain ion homeostasis. Our work provides a potential strategy for improving plant growth under multiple stresses.

Key miRNAs including sbi-miR169p/q, sbi-miR171g/j, sbi-miR172a/c/d, sbi-miR172e, sbi-miR319a/b, sbi-miR396a/b, miR408, sbi-miR5384, sbi-miR5565e and nov_23 were identified to function in the regulation of Cd accumulation and tolerance. As an energy plant, sweet sorghum shows great potential in the phytoremediation of Cd-contaminated soils. However, few studies have focused on the regulatory roles of miRNAs and their targets under Cd stress. In this study, comparative analysis of sRNAs, degradome and transcriptomics was conducted in high-Cd accumulation (H18) and low-Cd accumulation (L69) genotypes of sweet sorghum. A total of 38 conserved and 23 novel miRNAs with differential expressions were identified under Cd stress or between H18 and L69, and 114 target genes of 41 miRNAs were validated. Furthermore, 25 miRNA–mRNA pairs exhibited negatively correlated expression profiles and sbi-miR172e together with its target might participate in the distinct Cd tolerance between H18 and L69 as well as sbi-miR172a/c/d. Additionally, two groups of them: miR169p/q-nov_23 and miR408 were focused through the co-expression analysis, which might be involved in Cd uptake and tolerance by regulating their targets associated with transmembrane transportation, cytoskeleton activity, cell wall construction and ROS (reactive oxygen species) homeostasis. Further experiments exhibited that cell wall components of H18 and L69 were different when exposed to cadmium, which might be regulated by miR169p/q, miR171g/j, miR319a/b, miR396a/b, miR5384 and miR5565e through their targets. Through this study, we aim to reveal the potential miRNAs involved in sweet sorghum in response to Cd stress and provide references for developing high-Cd accumulation or high Cd-resistant germplasm of sweet sorghum that can be used in phytoremediation.

Summary Cadmium (Cd) is a widespread soil contaminant threatening human health. As an ideal energy plant, sweet sorghum (Sorghum bicolor (L.) Moench) has great potential in phytoremediation of Cd‐polluted soils, although the molecular mechanisms are largely unknown. In this study, key factors responsible for differential Cd accumulation between two contrasting sweet sorghum genotypes (high‐Cd accumulation one H18, and low‐Cd accumulation one L69) were investigated. H18 exhibited a much higher ability of Cd uptake and translocation than L69. Furthermore, Cd uptake through symplasmic pathway and Cd concentrations in xylem sap were both higher in H18 than those in L69. Root anatomy observation found the endodermal apoplasmic barriers were much stronger in L69, which may restrict the Cd loading into xylem. The molecular mechanisms underlying these morpho‐physiological traits were further dissected by comparative transcriptome analysis. Many genes involved in cell wall modification and heavy metal transport were found to be Cd‐responsive DEGs and/or DEGs between these two genotypes. KEGG pathway analysis found phenylpropanoid biosynthesis pathway was over‐represented, indicating this pathway may play important roles in differential Cd accumulation between two genotypes. Based on these results, a schematic representation of main processes involved in differential Cd uptake and translocation in H18 and L69 is proposed, which suggests that higher Cd accumulation in H18 depends on a multilevel coordination of efficient Cd uptake and transport, including efficient root uptake and xylem loading, less root cell wall binding, and weaker endodermal apoplasmic barriers.

Cadmium (Cd) contamination is a worldwide environmental problem, and remediation of Cd pollution is of great significance for food production as well as human health. Here, the responses of sweet sorghum cv. 'M-81E' to cadmium stress were studied for its potential as an energy plant in restoring soils contaminated by cadmium. In hydroponic experiments, the biomass of 'M-81E' showed no obvious change under 10 μM cadmium treatment. Cadmium concentration was the highest in roots of seedlings as well as mature plants, but in agricultural practice, the valuable and harvested parts of sweet sorghum are shoots, so promoting the translocation of cadmium to shoots is of great importance in order to improve its phytoremediation capacity. Further histochemical assays with dithizone staining revealed that cadmium was mainly concentrated in the stele of roots and scattered in intercellular space of caulicles. Moreover, the correlation analysis showed that Cd had a negative relationship with iron (Fe), zinc (Zn), and manganese (Mn) in caulicles and leaves and a positive relationship with Fe in roots. These results implied that cadmium might compete with Fe, Zn, and Mn for the transport binding sites and further prevent their translocation to shoots. In addition, transmission electron microscopic observations showed that under 100 μM cadmium treatment, the structure of chloroplast was impaired and the cell wall of vascular bundle cells in leaves and xylem and phloem cells in roots turned thicker compared to control. In summary, morphophysiological characteristic analysis demonstrated sweet sorghum can absorb cadmium and the growth is not negatively affected by mild level cadmium stress; thus, it is a promising material for the phytoremediation of cadmium-contaminated soils considering its economic benefit. This study also points out potential strategies to improve the phytoremediation capacity of sweet sorghum through genetic modification of transporters and cell wall components.

Highlight Unlike in glycophytes, NaCl facilitates the nitrate uptake in the euhalophyte S. europaea, while calcium signalling plays important roles in this process., Improving crop nitrogen (N) use efficiency under salinity is essential for the development of sustainable agriculture in marginal lands. Salicornia europaea is a succulent euhalophyte that can survive under high salinity and N-deficient habitat conditions, implying that a special N assimilation mechanism may exist in this plant. In this study, phenotypic and physiological changes of S. europaea were investigated under different nitrate and NaCl levels. The results showed that NaCl had a synergetic effect with nitrate on the growth of S. europaea. In addition, the shoot nitrate concentration and nitrate uptake rate of S. europaea were increased by NaCl treatment under both low N and high N conditions, suggesting that nitrate uptake in S. europaea was NaCl facilitated. Comparative proteomic analysis of root plasma membrane (PM) proteins revealed 81 proteins, whose abundance changed significantly in response to NaCl and nitrate. These proteins are involved in metabolism, cell signalling, transport, protein folding, membrane trafficking, and cell structure. Among them, eight proteins were calcium signalling components, and the accumulation of seven of the above-mentioned proteins was significantly elevated by NaCl treatment. Furthermore, cytosolic Ca2+ concentration ([Ca2+]cyt) was significantly elevated in S. europaea under NaCl treatment. The application of the Ca2+ channel blocker LaCl3 not only caused a decrease in nitrate uptake rate, but also attenuated the promoting effects of NaCl on nitrate uptake rates. Based on these results, a possible regulatory network of NaCl-facilitated nitrate uptake in S. europaea focusing on the involvement of Ca2+ signalling was proposed.

Phosphorus (P) is an essential mineral nutrient for plant growth and development. Low availability of inorganic phosphate (orthophosphate; Pi) in soil seriously restricts the crop production, while excessive fertilization has caused environmental pollution. Pi acquisition and homeostasis depend on transport processes controlled Pi transporters, which are grouped into five families so far: PHT1, PHT2, PHT3, PHT4, and PHT5. This review summarizes the current understanding on plant PHT families, including phylogenetic analysis, function, and regulation. The potential application of Pi transporters and the related regulatory factors for developing genetically modified crops with high phosphorus use efficiency (PUE) are also discussed in this review. At last, we provide some potential strategies for developing high PUE crops under salt or drought stress conditions, which can be valuable for improving crop yields challenged by global scarcity of water resources and increasing soil salinization.

A betA gene encoding choline dehydrogenase from Escherichia coli was transformed into cotton (Gossypium hirsutum L.) via Agrobacterium-mediated transformation. Transgenic cotton plants exhibited improved tolerance to chilling due to accumulation of glycinebetaine (GB). The results of our experiment showed that GB contents of leaves of transgenic lines 1, 3, 4, and 5, both before and after chilling stress, were significantly higher than those of wild-type (WT) plants. At 15°C, transgenic lines 1, 3, 4, and 5 exhibited higher germination capacity as determined by the germination speed and final germination percentage and, displayed less inhibition in seedling shoot growth rate than WT plants. Under chilling stress, transgenic lines 4 and 5 maintained higher relative water content, upper carbon dioxide (CO2) fixation capacity and PSII electron transfer rate, better osmotic adjustment (OA), a lower percentage of ion leakage, and less lipid membrane peroxidation when compared with WT plants. Chilling resistance of the transgenic lines was demonstrated to be positively correlated with GB content under chilling stress. The high levels of GB in transgenic cotton plants might not only protect the integrity of cell membrane from chilling damage, but also be involved in OA which alleviated chilling induced water stress. Moreover, under chilling-stressed conditions, transgenic cotton plants enhanced stomatal conductance, PSII electron transport rate, and further leaf photosynthesis through accumulating high levels of GB.

Salicornia europaea is a succulent euhalophyte that belongs to the Chenopodiaceae family. It is found that moderate concentration of NaCl can dramatically stimulate the growth of S. europaea plants. To elucidate the mech- anism underlying the phenomenon, morphological and physiological changes of S. europaea in response to dif- ferent ions, including cations (Na ? ,K ? ,L i ? ,C s ? ) and anions (Cl - ,N O 3 - ,C H 3COO - ) were investigated, and the effects of Na ? ,C l - and K ? on the growth of S. europaea were also studied. Na ? was more effective than K ? and Cl - in stimulating shoot succulence, cell expansion, and stomatal opening. Plants treated with Na ? (including NaCl, Na ? , NaNO3) showed better plant growth, increased pho- tosynthesis and less cell membrane damage than those untreated and treated with 200 mM of Cl - and K ? (including KCl and KNO3). Both SEM-X-Ray microanal- ysis and flame emission results revealed that well devel- oped S. europaea plants had a higher content of sodium but lower potassium and chlorine. It is concluded that sodium plays a more important role in the growth and development of S. europaea than potassium and chloride.

An H(+)-PPase gene, TsVP from Thellungiella halophila, was transferred into two cotton (Gossypium hirsutum) varieties (Lumianyan19 and Lumianyan 21) and southern and northern blotting analysis showed the foreign gene was integrated into the cotton genome and expressed. The measurement of isolated vacuolar membrane vesicles demonstrated that the transgenic plants had higher V-H(+)-PPase activity compared with wild-type plants (WT). Overexpressing TsVP in cotton improved shoot and root growth, and transgenic plants were much more resistant to osmotic/drought stress than the WT. Under drought stress conditions, transgenic plants had higher chlorophyll content, improved photosynthesis, higher relative water content of leaves and less cell membrane damage than WT. We ascribe these properties to improved root development and the lower solute potential resulting from higher solute content such as soluble sugars and free amino acids in the transgenic plants. In this study, the average seed cotton yields of transgenic plants from Lumianyan 19 and Lumianyan 21 were significantly increased compared with those of WT after exposing to drought stress for 21 days at flowering stage. The average seed cotton yields were 51 and 40% higher than in their WT counterparts, respectively. This study benefits efforts to improve cotton yields in arid and semiarid regions.

Increasing soil salinity threatens crop productivity worldwide. High soil salinity is usually accompanied by the low availability of many mineral nutrients. Here, we investigated the potential role that the H(+)- PPase could play in optimizing P use efficiency under salinity in plants. Transgenic Arabidopsis plants overexpressing either SeVP1 or SeVP2 from Salicornia europaea outperformed the wild-types under low phosphate (Pi) as well as low Pi plus salt conditions. Our results suggested that H(+)-PPase could increase external Pi acquisition through promoting root development and upregulating phosphate transporters, thus to protect plants from Pi limiting stress. This study provides a potential strategy for improving crop yields challenged by the co-occurrence of abiotic stresses.

Exploiting the negative biochemical interference between plants and algal species has been suggested as a method to control harmful algal blooms. In this work, we investigated the inhibitory effect of the salt marsh halophyte Salicornia europaea against the marine alga Skeletonema costatum. S. europaea suppressed the growth of S. costatum in a nutrient-sufficient co-culture system, indicating that the inhibition of algal growth was because of the phytotoxic effect of S. europaea, rather than nutrient competition. We tested aqueous and organic extracts from S. europaea roots against S. costatum. The organic extracts inhibited growth and affected the cell size and chlorophyll a content of S. costatum in a dose-dependent manner. Among the three tested organic extracts, the methanol extract had the greatest effects on S. costatum, followed by butanol extract, and then the chloroform extract. Two flavonoids, rutin and quercetin-3-β-D-glucoside, were identified in the methanol extract by high performance liquid chromatography. The concentration of rutin was much higher than that of quercetin-3-β-D-glucoside. In an algal bioassay, rutin inhibited the growth of S. costatum and the inhibitory effect increased with increasing rutin concentration and with decreasing initial algal density. Therefore, we concluded that S. europaea negatively affects the growth of S. costatum, and that rutin, a metabolite of S. europaea, may play a role in this inhibitory effect.

Euhalophyte Salicornia europaea L., one of the most salt-tolerant plant species in the world, can tolerate more than 1000 mM NaCl. To study the salt tolerance mechanism of this plant, the effects of different NaCl concentrations on plant growth, as well as Na(+) accumulation and distribution at organ, tissue, and subcellular levels, were investigated. Optimal growth and an improved photosynthetic rate were observed with the plant treated with 200-400 mM NaCl. The Na(+) content in the shoots was considerably higher than that in the roots of S. europaea. The Na(+) in S. europaea cells may act as an effective osmotic adjuster to maintain cell turgor, promoting photosynthetic competence and plant growth. The results from the SEM-X-ray and TEM-X-ray microanalyses demonstrate that Na(+) was compartmentalized predominantly into the cell vacuoles of shoot endodermis tissues. Accordingly, the transcript amounts of SeNHX1, SeVHA-A, and SeVP1 increased significantly with increased NaCl concentration, suggesting their important roles in Na(+) sequestration into the vacuoles. Therefore, a multiple sodium compartmentalization mechanism is proposed to enhance further the salt tolerance of S. europaea.

Inhibition of lycopene cyclization decreased the salt tolerance of the euhalophyte Salicornia europaea L. We isolated a β-lycopene cyclase gene SeLCY from S. europaea and transformed it into Arabidopsis with stable expression. Transgenic Arabidopsis on post-germination exhibited enhanced tolerance to oxidative and salt stress. After 8 and 21 d recovery from 200 mM NaCl treatment, transgenic lines had a higher survival ratio than wild-type (WT) plants. Three-week-old transgenic plants treated with 200 mM NaCl showed better growth than the WT with higher photosystem activity and less H(2)O(2) accumulation. Determination of endogenous pigments of Arabidopsis treated with 200 mM NaCl for 0, 2 or 4 d demonstrated that the transgenic plants retained higher contents of carotenoids than the WT. Furthermore, to compare the difference between SeLCY and AtLCY from Arabidopsis, we used viral vector mediating ectopic expression of SeLCY and AtLCY in Nicotiana benthamiana. Although LCY genes transformation increased the salt tolerance in tobacco, there is no significant difference between SeLCY- and AtLCY-transformed plants. These findings indicate that SeLCY transgenic Arabidopsis improved salt tolerance by increasing synthesis of carotenoids, which impairs reactive oxygen species and protects the photosynthesis system under salt stress, and as a single gene, SeLCY functionally showed no advantage for salt tolerance improvement compared with AtLCY.