Temperature/salt tolerance and oil recovery of xanthan gum solution enhanced by surface-modified nanosilicas

Amide-and alkyl-modified nanosilicas(AANPs)were synthesized and introduced into Xanthan gum(XG)solution,aiming to improve the temperature/salt tolerance and oil recovery.The rheological behaviors of XG/AANP hybrid dispersions were systematically studied at different concentrations,temperatures and inorganic salts.At high temperature(75C)and high salinity(10,000 mg,L1 NaCl),AANPs increase the apparent viscosity and dynamic modulus of the XG solution,and XG/AANP hybrid dispersion exhibits elastic-dominant properties.The most effective concentrations of XG and AANP interacting with each other are 1750 mg·L^(-1) and 0.74 wt%,respectively.The temperature tolerance of XG solution is not satisfactory,and high temperature further weakens the salt tolerance of XG.However,the AANPs significantly enhance the viscoelasticity the XG solution through hydrogen bonds and hydrophobic effect.Under reservoir conditions,XG/AANP hybrid recovers approximately 18.5%more OOIP(original oil in place)than AANP and 11.3%more OOIP than XG.The enhanced oil recovery mechanism of the XG/AANP hybrid is mainly increasing the sweep coefficient,the contribution from the reduction of oil-water interfacial tension is less.

Eco-friendly aqueous foam stabilized by cellulose microfibers with great salt tolerance and high temperature resistance

A low-cost eco-friendly aqueous foam,especially the robust foam with great tolerance to high salinity and high temperature,is in great demand in the oil industry,e.g.,oil and gas well or geothermal well drilling.Herein,an ultra-stable aqueous foam was developed using the biodegradable cellulose microfiber(CMF)as a foam stabilizer.The foam stabilized by CMF shows excellent tolerance to the high concentration of NaCl(6.0 wt%)and CaCl_(2)(0.25 wt%)and the related drainage half-life times(T_(0.5))reach 1750 and 2340 s respectively.By contrast,the foams without CMF are completely drained(T_(0.5)=0 s)when NaCl concentration is greater than 6.0 wt%or CaCl_(2) concentration is greater than 0.20 wt%.Notably,T0.5 of the foams stabilized by CMF at these saline concentrations still can maintain above 1000 s even after aging at 120℃ for 16 h,exhibiting an outstanding foam-stabilizing performance at high temperature.Experimental results suggest that the salt and high-temperature tolerance of CMF in foam stabilization is attributed to the electrically uncharged surfaces,the formation of a gel-like structure and the excellent thermal stability.This work not only provides a promising candidate of aqueous foam stabilizer to deal with high temperature and high salinity but also presents a natural-based solution for an environmentally friendly drilling industry in the future.

Advances in deciphering salt tolerance mechanism in maize

Maize(Zea mays L.)is a global cereal crop whose demand is projected to double by 2050.Along with worsening of farmland salinization,salt stress has become a major environmental threat to the sustainability of maize production worldwide.Accordingly,there is an urgent need to decipher salt-tolerant mechanisms and facilitate the breeding of salt-tolerant maize.As salt tolerance is a complex trait regulated by multiple genes,and maize germplasm varies widely in salt tolerance,efforts have been devoted to the identification and application of quantitative-trait loci(QTL)for salt tolerance.QTL associated with ion regulation,osmotic tolerance,and other aspects of salt tolerance have been discovered using genomewide association studies(GWAS),linkage mapping,and omics-based approaches.This review highlights recent advances in the molecular-level understanding of salt stress response in maize,in particular in(a)the discovery of salt-tolerance QTL,(b)the mechanisms of salt tolerance,(c)the development of salttolerant maize cultivars,and(d)current challenges and future prospects.

SlTPP4 participates in ABA-mediated salt tolerance by enhancing root architecture in tomato

Salinity tolerance is an important physiological index for crop breeding.Roots are typically the first plant tissue to withstand salt stress.In this study,we found that the tomato(Solanum lycopersicum)trehalose-6-phosphate phosphatase(SlTPP4)gene is induced by abscisic acid(ABA)and salt,and is mainly expressed in roots.Overexpression of SlTPP4 in tomato enhanced tolerance to salt stress,resulting in better growth performance.Under saline conditions,SlTPP4 overexpression plants demonstrated enhanced sucrose metabolism,as well as increased expression of genes related to salt tolerance.At the same time,expression of genes related to ABA biosynthesis and signal transduction was enhanced or altered,respectively.In-depth exploration demonstrated that SlTPP4 enhances Casparian band development in roots to restrict the intake of Na^(+).Our study thus clarifies the mechanism of SlTPP4-mediated salt tolerance,which will be of great importance for the breeding of salt-tolerant tomato crops.

A cluster of mutagenesis revealed an osmotic regulatory role of the OsPIP1 genes in enhancing rice salt tolerance

Aquaporins play important regulatory roles in improving plant abiotic stress tolerance.To better understand whether the Os PIP1 genes collectively dominate the osmotic regulation in rice under salt stress,a cluster editing of the Os PIP1;1,Os PIP1;2 and Os PIP1;3 genes in rice was performed by CRISPR/Cas9 system.Sequencing showed that two mutants with Cas9-free,line 14 and line 18 were successfully edited.Briefly,line 14 deleted a single C base in both the Os PIP1;1 and Os PIP1;3 genes,and inserted a single T base in the Os PIP1;2 gene,respectively.While line 18 demonstrated an insertion of a single A base in the Os PIP1;1gene and a single T base in both the Os PIP1;2 and Os PIP1;3 genes,respectively.Multiplex editing of the Os PIP1 genes significantly inhibited photosynthetic rate and accumulation of compatible metabolites,but increased MDA contents and osmotic potentials in the mutants,thus delaying rice growth under salt stress.Functional loss of the Os PIP1 genes obviously suppressed the expressions of the Os PIP1,Os SOS1,Os CIPK24 and Os CBL4 genes,and increased the influxes of Na+and effluxes of K^(+)/H^(+)in the roots,thus accumulating more Na+in rice mutants under salt stress.This study suggests that the Os PIP1 genes are essential modulators collectively contributing to the enhancement of rice salt stress tolerance,and multiplex editing of the Os PIP1 genes provides insight into the osmotic regulation of the PIP genes.

The Improvement of Soybean Salt Tolerance by Overexpressed GmPAO1

Polyamines play an important regulatory role during plant growth and development and adversity stress,and polyamine oxidase(PAO)is involved in polyamine catabolism.In this study,an up-regulated polyamine oxidase gene GmPAO1 was obtained by transcriptome sequencing analysis and screening at soybean seedling stages.Also,its expression pattern and function were analyzed.The identification results of transgenic GmPAO1 soybean positive lines showed that the relative expression level of GmPAO1 in the overexpressed lines was increased under salt stress.With increasing stress concentration,the seed germination rate decreased.However,the seed germination rate of the overexpressed lines was significantly higher than that of the control lines,and the phenotypic character of the root systems was also better than that of the control lines.The measurement of superoxide dismutase(SOD)and peroxidase(POD)activities and malondialdehyde and hydrogen peroxide contents revealed that the overexpressed soybean lines significantly increased the SOD and POD activities,significantly reducing the malondialdehyde content.Although the hydrogen peroxide content in the transformed plants gradually increased,the hydrogen peroxide content in the overexpression lines was still lower than that in the gene editing lines.Based on this,it was preliminarily judged that GmPAO1 can improve soybean salt tolerance.

Thellungiella halophila ST5 improves salt tolerance in cotton

Background: Salinity is a major abiotic stress to global agriculture which hampers crop growth and development, and eventually reduces yield. Transgenic technology is an e ective and e cient approach to improve crop salt tolerance but depending on the availability of e ective genes. We previously isolated Salt Tolerance5(ThST5) from the halophyte Thellungiella halophila, an ortholog of Arabidopsis SPT4-2 which encodes a transcription elongation factor. However, SPT4-2-confered salt tolerance has not been evaluated in crops yet. Here we report the evaluation of Th ST5-conferred salt tolerance in cotton(Gossypium hirsutum L.).Results: The ThST5 overexpression transgenic cotton plants displayed enhanced tolerance to salt stress during seed germination and seedling stage compared with wild type. Particularly, the transgenic plants showed improved salinity tolerance as well as yield under saline field conditions. Comparative transcriptomic analysis showed that ThST5 improved salt tolerance of transgenic cotton mainly by maintaining ion homeostasis. In addition, ThST5 also orchestrated the expression of genes encoding antioxidants and salt-responsive transcription factors.Conclusion: Our results demonstrate that ThST5 is a promising candidate to improve salt tolerance in cotton.

Plant salt tolerance and Na^+ sensing and transport

Salinity is a global challenge to agricultural production. Understanding Na^+ sensing and transport in plants under salt stress will be of benefit for breeding robustly salt-tolerant crop species. In this review, first, possible salt stress sensor candidates and the root meristem zone as a tissue harboring salt stress-sensing components are proposed. Then,the importance of Na^+ exclusion and vacuolar Na^+ sequestration in plant overall salt tolerance is highlighted. Other Na^+ regulation processes, including xylem Na^+ loading and unloading, phloem Na^+ recirculation, and Na^+ secretion, are discussed and summarized.Along with a summary of Na^+ transporters and channels, the molecular regulation of Na^+ transporters and channels in response to salt stress is discussed. Finally, some largely neglected issues in plant salt stress tolerance, including Na^+ concentration in cytosol and the role of Na^+ as a nutrient, are reviewed and discussed.

Salt tolerance in rice:Physiological responses and molecular mechanisms

Crop yield loss due to soil salinization is an increasing threat to agriculture worldwide.Salt stress drastically affects the growth,development,and grain productivity of rice(Oryza sativa L.),and the improvement of rice tolerance to salt stress is a desirable approach for meeting increasing food demand.The main contributors to salt toxicity at a global scale are Na^(+)and Cl^(-)ions,which affect up to 50%of irrigated soils.Plant responses to salt stress occur at the organismic,cellular,and molecular levels and are pleiotropic,involving(1)maintenance of ionic homeostasis,(2)osmotic adjustment,(3)ROS scavenging,and(4)nutritional balance.In this review,we discuss recent research progress on these four aspects of plant physiological response,with particular attention to hormonal and gene expression regulation and salt tolerance signaling pathways in rice.The information summarized here will be useful for accelerating the breeding of salt-tolerant rice.

ZmWRKY104 positively regulates salt tolerance by modulating ZmSOD4 expression in maize

Salinity impairs plant growth, limiting agricultural development. It is desirable to identify genes responding to salt stress and their mechanism of action. We identified a function of the Zea mays WRKY transcription factor, Zm WRKY104, in salt stress response. Zm WRKY104 was localized in the nucleus and showed transcriptional activation activity. Phenotypic and physiological analysis showed that overexpression of Zm WRKY104 in maize increased the tolerance of maize to salt stress and alleviated salt-induced increases in O;accumulation, malondialdehyde(MDA) content, and percent of electrolyte leakage. Further investigation showed that Zm WRKY104 increased SOD activity by regulating Zm SOD4 expression. Yeast onehybrid, electrophoretic mobility shift test, and chromatin immunoprecipitation–quantitative PCR assay showed that Zm WRKY104 bound directly to the promoter of Zm SOD4 by recognizing the W-box motif in vivo and in vitro. Phenotypic, physiological, and biochemical analysis showed that Zm SOD4 increased salt tolerance by alleviating salt-induced increases in O;accumulation, MDA content, and percent of electrolyte leakage under salt stress. Taken together, our results indicate that Zm WRKY104 positively regulates Zm SOD4 expression to modulate salt-induced O;accumulation, MDA content, and percent of electrolyte leakage, thus affecting salt stress response in maize.

Haplotype variations in QTL for salt tolerance in Chinese wheat accessions identified by marker-based and pedigree-based kinship analyses

Most modern wheat cultivars were selected on the basis of yield-related indices measured under optimal fertilizer and irrigation inputs.With climate change,land degradation and salinity caused by sea water encroachment,wheat is increasingly subjected to environmental stress.Moreover,expanding urbanization increasingly encroaches upon prime agricultural land in countries like China,and alternative cropping areas must be found.Some of these areas have moderate constraining factors,such as salinity.Therefore,it is important to investigate whether current genetic materials and breeding procedures are maintaining adequate variability to address future problems caused by abiotic stress.In this study,a panel of 307 wheat accessions,including local landraces,exotic cultivars used in Chinese breeding programs and Chinese cultivars released during different periods since1940,were subjected to a genome-wide association study to dissect the genetic basis of salinity tolerance.Both marker-based and pedigree-based kinship analyses revealed that favorable haplotypes were introduced in some exotic cultivars as well as a limited number of Chinese landraces from the 1940 s.However,improvements in salinity tolerance during modern breeding are not as obvious as that of yield.To broaden genetic diversity for increasing salt tolerance,there is a need to refocus attention on local landraces that have high degrees of salinity tolerance and carry rare favorable alleles that have not been exploited in breeding.

The osmolyte-producing endophyte Streptomyces albidoflavus OsiLf-2 induces drought and salt tolerance in rice via a multi-level mechanism

Drought and salinity are major environmental stresses that impair crop growth and productivity worldwide. Improving drought and salt tolerance of crops with microbial mutualists is an effective and environmentally sound strategy to meet the demands of the ever-growing world population. In the present study, we found that the Streptomyces albidoflavus Osi Lf-2, a moderately salt-tolerant endophytic actinomycete, produced abundant osmolytes, including proline, polysaccharides, and ectoine. Inoculation with Osi Lf-2 increased the osmotic-adjustment ability of the rice host by increasing the proline content(by250.3% and 49.4%) and soluble sugar(by 20.9% and 49.4%) in rice under drought and salt conditions, relative to the uninoculated control. Osi Lf-2 increased stress responses in the rice host at the physiological and biochemical levels(photosynthesis efficiency, osmolytes and antioxidant content), and the gene level(osmolytes synthesis, stress-responsive and ion-transport related genes), raising rice yields under both greenhouse and saline–alkaline soil conditions. The use of endophytic actinomycetes offers a promising biotechnological approach to developing stress-tolerant plants.

The NADPH oxidase OsRbohA increases salt tolerance by modulating K^(+)homeostasis in rice

Rice(Oryza sativa L.)is a staple cereal for more than two thirds of the world's population.Soil salinity severely limits rice growth,development,and grain yield.It is desirable to elucidate the mechanism of rice's salt-stress response.As the major source of H_(2)O_(2),NADPH oxidase(Rboh)is believed to be involved in salt-stress tolerance.However,the function and mechanism of rice Rboh in salt stress response remain unclear.In this study,we found that the expression of OsRbohA was up-regulated by NaCl treatment in the shoots and roots of rice seedlings.Knockout of OsRbohA reduced the tolerance of rice to salt stress.Knockout of OsRbohA blocked NaCl-induced increases of NADPH activity and H_(2)O_(2) content in roots.OsRboh A knockout inhibited root growth and disrupted K^(+)homeostasis by reducing the expression of K^(+) transporters and channel-associated genes(OsGORK,OsAKT1,OsHAK1,and OsHAK5)in roots under NaCl treatment.Under NaCl treatment,OsRbohA knockout also reduced subcellular K^(+) contents of the plasma membrane and soluble fraction.Overexpression of OsRbohA increased the expression of K^(+) transporters and channel-associated genes and reduced the loss of K^(+) ions in roots.These results indicate that OsRboh A-mediated H_(2)O_(2) accumulation modulates K^(+) homeostasis,thereby increasing salt tolerance in rice.

Calcineurin B-like protein 5(SiCBL5)in Setaria italica enhances salt tolerance by regulating Na^(+)homeostasis

Salinity,a major abiotic stress,reduces plant growth and severely limits agricultural productivity.Plants regulate salt uptake via calcineurin B-like proteins(CBLs).Although extensive studies of the functions of CBLs in response to salt stress have been conducted in Arabidopsis,their functions in Setaria italica are still poorly understood.The foxtail millet genome encodes seven CBLs,of which only SiCBL4 was shown to be involved in salt response.Overexpression of SiCBL5 in Arabidopsis thaliana sos3-1 mutant rescued its salt hypersensitivity phenotype,but that of other SiCBLs(SiCBL1,SiCBL2,SiCBL3,SiCBL6,and SiCBL7)did not rescue the salt hypersensitivity of the Atsos3-1 mutant.SiCBL5 harbors an N-myristoylation motif and is located in the plasma membrane.Overexpression of SiCBL5 in foxtail millet increased its salt tolerance,but its knockdown increased salt hypersensitivity.Yeast two-hybrid and firefly luciferase complementation imaging assays showed that SiCBL5 physically interacted with SiCIPK24 in vitro and in vivo.Cooverexpression of SiCBL5,SiCIPK24,and SiSOS1 in yeast conferred a high-salt-tolerance phenotype.Compared to wild-type plants under salt stress conditions,SiCBL5 overexpressors showed lower accumulations of Na^(+) and stronger Na^(+) efflux,whereas RNAi-SiCBL5 plants showed higher accumulations of Na^(+) and weaker Na^(+) efflux.These results indicate that SiCBL5 confers salt tolerance in foxtail millet by modulating Na^(+) homeostasis.

Physiological analysis reveals relatively higher salt tolerance in roots of Ilex integra than in those of Ilex purpurea

Determining the responses of candidate plants to salt stress is a prerequisite for selecting and breeding suitable plants with high salt tolerance to grow in coastal mudfl at areas with high salinity.Here,2-year cutting seedlings of Ilex purpurea Hassk.(local species)and I.integra Thunb.(introduced species)were grown in pots in a glasshouse and irrigated with a Hoagland-NaCl solution at 0,24,and 48 h.Root samples were collected at 0,1,6,24,and 72 h,and concentration of Na^(+)ion;content of proline,soluble carbohydrate,malondialdehyde(MDA),H_(2)O_(2) and ascorbate;and activity of three key antioxidative enzymes were measured.Roots of I.integra accumulated relatively less Na^(+)and had less membrane lipid peroxidation and H_(2)O_(2) during salt stress,thus indicating a relatively higher salt tolerance than roots of I.purpurea.Values for ascorbate content and antioxidant enzymatic activity suggest that the antioxidant ascorbate and antioxidative catalase may play substantial roles for scavenging reactive oxygen species in I.integra roots during salt treatment.Thus,I.integra is apparently more suitable for growing in local highly saline coastal mudfl ats.

Study on Relationship Between Differential Proteins of Bacillus cereus LBR-4 and Its Salt Tolerance Mechanism

In order to explore the salt tolerance mechanism of Bacillus cereus LBR-4 with salinity of 14%NaCl,differential proteomic analysis of the whole protein of LBR-4 strain expressed under 14%NaCl high salinity condition and normalculture condition(1%NaCl)was studied by two-dimensional electrophoresis and mass spectrometry.The isoelectric point of most detected proteins was between pH 4-7 and the molecular weight distribution was 10-70 ku.Compared with the normal culture condition,the expression level of 118 protein spots in the whole protein expression map changed significantly(accounting for 25.2%of the total protein spots).The expression level of 78 protein spots increased significantly,including 22 new protein spots that appeared under high salt stress.The expression levels of 40 protein spots decreased significantly,including 18 protein spots that disappeared under high salt stress.By mass spectrometry,six distinct differentially expressed protein spotswere dihydroxy acid dehydratase,cell division protein FtsZ,iron sulfur cluster synthesis protein SufD,unknown carboxylase YngE,hypothetical acetaldehyde dehydrogenase DhaS and phenylalanine acid tRNA ligase alpha subunit.It was speculated that under high salt stress,the cells had protective measures and the secretion of intracellular compatible solutes increased.The iron and sulfur clusters involved in various physiological reactions also activated the stressful suf synthesis pathway,and therate of cell division and reproduction was also slowed down and ensured the normal progress of physiological reactions inthe cells.

Evaluation of Thellungiella halophila ST7 for improving salt tolerance in cotton

Background: Gossypium hirsutum(upland cotton) is one of the principal fiber crops in the world. Cotton yield is highly affected by abiotic stresses, among which salt stress is considered as a major problem around the globe. Transgenic approach is efficient to improve cotton salt tolerance but depending on the availability of salt tolerance genes.Results: In this study we evaluated salt tolerance candidate gene ST7 from Thellungiella halophila, encoding a homolog of Arabidopsis aluminum-induced protein, in cotton. Our results showed that ThST7 overexpression in cotton improved germination under NaCl stress as well as seedling growth. Our field trials also showed that ThST7 transgenic cotton lines produced higher yield under salt stress conditions. The improved salt tolerance of the transgenic cotton lines was partially contributed by enhanced antioxidation as shown by diaminobenzidine(DAB) and nitrotetrazolium blue chloride(NBT) staining. Moreover, transcriptomic analysis of ThST7 overexpression lines showed a significant upregulation of the genes involved in ion homeostasis and antioxidation, consistent with the salt tolerance phenotype of the transgenic cotton.Conclusions: Our results demonstrate that ThST7 has the ability to improve salt tolerance in cotton. The ThST7 transgenic cotton may be used in cotton breeding for salt tolerance cultivars.

A high salt tolerance and low adsorption drag reducer based on non-covalent enhancement

To formulate fluids with flowback water,produced water directly to improve the utilization rate of recycling and reduce the adsorption damage of slick water to reservoirs,a high salt tolerance and low adsorption drag reducer was designed and prepared by introducing polar cation fragments to enhance the non-covalent interactions between the chains.The drag reducer was characterized by IR and NMR.Friction resistance and viscosity tests were conducted to evaluate its salt resistance property.Static adsorption and dynamic adsorption retention tests were carried out to evaluate the damage of this reducer to shale reservoirs.The introduction of cation units into the molecular structure can weak the shielding effect of metal cations to some extent,so the drag reducer can keep a stable molecular structure and good resistant reducing performance under high salinity.The enhancement of non-covalent interaction between chains decreased the free polarity sites,further reduced the possibility of hydrogen bonding between drag reducer molecules and shale.In high salinity condition,both the adsorption capacity of the drag reducer on the shale surface and the average damage rate to the core permeability are low.Compared with the conventional salt-tolerant system,the overall liquid cost was reduced by 17%and the production per well increased by 44%.The application of this slick water system has achieved remarkable results.

Mapping QTLs for Salt Tolerance at Seeding Stage of Landrace Rice (Oryza sativa L.)

[Objectives] The identification of salt tolerant genetic loci in rice can provide a research basis for the molecular mechanism of salt tolerance and gene resources for improving salt tolerant cultivars. [Methods] Recombinant inbred lines(RILs) derived from Zhaxima, an indica landrace variety from Yunnan Province and Nanjing 46, an elite japonica variety with superior grain quality from Jiangsu Province were used. The salt tolerance at seeding stage in the RIL population was investigated as the phenotypic value. [Results] Combined with the linkage map, a total of 4 QTLs were detected: qSST-1, qSST-3, qSST-5 and qSST-11, located in rice chromosomes 1, 3, 5 and 11, respectively. All positive alleles were from the parent Nanjing 46. Three QTLs among them were not included in chromosome intervals the same as cloned rice salt tolerance genes, and thus were described as new candidate gene loci associated with seeding-stage salt tolerance. [Conclusions] This study provides important information for further exploration and utilization of new salt tolerant QTLs in rice. It is of great significance for improving the utilization of saline land in China and ensuring the stable rice production.

Cytokinin signaling promotes salt tolerance by modulating shoot chloride exclusion in maize

Excessive accumulation of chloride(Cl^(-))in the aboveground tissues under saline conditions is harmful to crops.Increasing the exclusion of Cl^(-) from shoots promotes salt tolerance in various crops.However,the underlying molecular mechanisms remain largely unknown.In this study,we demonstrated that a type A response regulator(ZmRR1)modulates Cl^(-) exclusion from shoots and underlies natural variation of salt tolerance in maize.ZmRR1 negatively regulates cytokinin signaling and salt tolerance,likely by interacting with and inhibiting His phosphotransfer(HP)proteins that are key mediators of cytokinin signaling.A naturally occurring non-synonymous SNP variant enhances the interaction between ZmRR1 and ZmHP2,conferring maize plants with a salt-hypersensitive phenotype.We found that ZmRR1 undergoes degradation under saline conditions,leading to the release of ZmHP2 from ZmRR1 inhibition,and subsequently ZmHP2-mediated signaling improves salt tolerance primarily by promoting Cl^(-) exclusion from shoots.Furthermore,we showed that ZmMATE29 is transcriptionally upregulated by ZmHP2-mediated signaling under highly saline conditions and encodes a tonoplast-located Cl^(-) transporter that promotes Cl^(-) exclusion from shoots by compartmentalizing Cl^(-) into the vacuoles of root cortex cells.Collectively,our study provides an important mechanistic understanding of the cytokinin signaling-mediated promotion of Cl^(-) exclusion from shoots and salt tolerance and suggests that genetic modification to promote Cl^(-) exclusion from shoots is a promising route for developing salt-tolerant maize.