Salinity Effects on Guard Cell Proteome in Chenopodium quinoa.

Epidermal fragments enriched in guard cells (GCs) were isolated from the halophyte quinoa (Chenopodium quinoa Wild.) species, and the response at the proteome level was studied after salinity treatment of 300 mM NaCl for 3 weeks. In total, 2147 proteins were identified, of which 36% were differentially expressed in response to salinity stress in GCs. Up and downregulated proteins included signaling molecules, enzyme modulators, transcription factors and oxidoreductases. The most abundant proteins induced by salt treatment were desiccation-responsive protein 29B (50-fold), osmotin-like protein OSML13 (13-fold), polycystin-1, lipoxygenase, alpha-toxin, and triacylglycerol lipase (PLAT) domain-containing protein 3-like (eight-fold), and dehydrin early responsive to dehydration (ERD14) (eight-fold). Ten proteins related to the gene ontology term "response to ABA" were upregulated in quinoa GC; this included aspartic protease, phospholipase D and plastid-lipid-associated protein. Additionally, seven proteins in the sucrose-starch pathway were upregulated in the GC in response to salinity stress, and accumulation of tryptophan synthase and L-methionine synthase (enzymes involved in the amino acid biosynthesis) was observed. Exogenous application of sucrose and tryptophan, L-methionine resulted in reduction in stomatal aperture and conductance, which could be advantageous for plants under salt stress. Eight aspartic proteinase proteins were highly upregulated in GCs of quinoa, and exogenous application of pepstatin A (an inhibitor of aspartic proteinase) was accompanied by higher oxidative stress and extremely low stomatal aperture and conductance, suggesting a possible role of aspartic proteinase in mitigating oxidative stress induced by saline conditions.

Effect of salinity on physiological, biochemical and photostabilizing attributes of two genotypes of quinoa (Chenopodium quinoa Willd.) exposed to arsenic stress.

Soil salinity and arsenic (As) contamination are serious environmental problems. To investigate the effects of salinity on As uptake and physiological and biochemical attributes of quinoa (Chenopodiumquinoa Willd.), a hydroponic experiment was performed. One-month old healthy plants of two quinoa genotypes; Vikinga and A7 were transplanted in plastic tubs containing half strength Hoagland's nutrient solution. Plants were exposed to different levels of As (0, 150 and 300 µM), salinity (0, 150 and 300 mM) and their combinations (150 µM As + 300 mM NaCl; 300 µM As + 300 mM NaCl) for five weeks. Results revealed that combined application of salinity and As caused more pronounced reduction in growth, chlorophyll contents and caused more oxidative damage in both quinoa genotypes. Under combined application of salinity and As, Na+ concentration was increased whereas As content was decreased in plant tissues. Quinoa genotype A7 was more tolerant than Vikinga against salinity, As and their combination perhaps because of less uptake of toxic ions and higher activities of antioxidant enzymes (SOD, CAT, POD). Bioconcentration factor (BCF), translocation factor (TF) and tolerance index (TI) indicated that genotype A7 can be successfully employed for phytostabilization of As contaminated saline soils.

Acid treated biochar enhances cadmium tolerance by restricting its uptake and improving physio-chemical attributes in quinoa (Chenopodium quinoa Willd.).

Heavy metals contamination of soil especially with cadmium (Cd) is a serious environmental concern in the current industrial era. Biochar serves as an excellent ameliorating agent depending upon its properties and application rates. In the pot scale study, effect of acid treated (AWSB) and untreated wheat straw biochar (WSB) was studied on physiology, grain yield, Cd accumulation, and tolerance of quinoa with possible health risks. Different levels of Cd (0, 25, 50 and 75 mg kg-1), AWSB and WSB (1% and 2% (w/w)) were applied in soil. Accumulation of Cd in control plant tissues led to oxidative stress which was shown in terms of increased lipid peroxidation. While biochar application relieved the oxidative damage as confirmed by the low production of H2O2 and TBARS contents. Application of AWSB improved plant growth, pigment contents and gas exchange attributes by limiting the accumulation of Cd in root, shoot and grain of quinoa. Results revealed a significant improvement in the activity of superoxide (SOD), catalase (CAT), ascorbate peroxidase (APX) and peroxidase (POD) with biochar at elevated levels of Cd in soil. Target Hazard Quotient (THQ) remained < 1 in the quinoa grains with WSB and AWSB under Cd stress. These results revealed that AWSB most effectively alleviated Cd toxicity in quinoa thereby decreasing Cd accumulation and regulation of Cd induced oxidative stress triggered by the antioxidant enzymatic system.

The combined effect of Cr(III) and NaCl determines changes in metal uptake, nutrient content, and gene expression in quinoa (Chenopodium quinoa Willd.).

Many areas of the world are affected simultaneously by salinity and heavy metal pollution. Halophytes are considered as useful candidates in remediation of such soils due to their ability to withstand both osmotic stress and ion toxicity deriving from high salt concentrations. Quinoa (Chenopodium quinoa Willd) is a halophyte with a high resistance to abiotic stresses (drought, salinity, frost), but its capacity to cope with heavy metals has not yet been fully investigated. In this pot experiment, we investigated phytoextraction capacity, effects on nutrient levels (P and Fe), and changes in gene expression in response to application of Cr(III) in quinoa plants grown on saline or non-saline soil. Plants were exposed for three weeks to 500 mg kg-1 soil of Cr(NO3)3·9H2O either in the presence or absence of 150 mM NaCl. Results show that plants were able tolerate this soil concentration of Cr(III); the metal was mainly accumulated in roots where it reached the highest concentration (ca. 2.6 mg g-1 DW) in the presence of NaCl. On saline soil, foliar Na concentration was significantly reduced by Cr(III). Phosphorus translocation to leaves was reduced in the presence of Cr(III), while Fe accumulation was enhanced by treatment with NaCl alone. A real-time RT-qPCR analysis was conducted on genes encoding for sulfate, iron, and phosphate transporters, a phytochelatin, a metallothionein, glutathione synthetase, a dehydrin, Hsp70, and enzymes responsible for the biosynthesis of proline (P5CS), glycine betaine (BADH), tocopherols (TAT), and phenolic compounds (PAL). Cr(III), and especially Cr(III)+NaCl, affected transcript levels of most of the investigated genes, indicating that tolerance to Cr is associated with changes in phosphorus and sulfur allocation, and activation of stress-protective molecules. Moderately saline conditions, in most cases, enhanced this response, suggesting that the halophytism of quinoa could contribute to prime the plants to respond to chromium stress.

Soil drenching of paclobutrazol: An efficient way to improve quinoa performance under salinity.

Salinity extent and severity is rising because of poor management practices on agricultural lands, possibility lies to grow salt-tolerant crops with better management techniques. Therefore, a highly nutritive salt-tolerant crop quinoa with immense potential to contribute for future food security was selected for this investigation. Soil drenching of paclobutrazol (PBZ; 20 mg l-1 ) was used to understand the ionic relations, gaseous exchange characteristics, oxidative defense system and yield under saline conditions (400 mM NaCl) including normal (0 mM NaCl) and no PBZ (0 mg l-1 ) as controls. The results revealed that salinity stress reduced the growth and yield of quinoa through perturbing ionic homeostasis with the consequences of overproduction of reactive oxygen species (ROS), oxidative damages and reduced photosynthesis. PBZ improved the quinoa performance through regulation of ionic homeostasis by decreasing Na+ , Cl- , while improving K+ , Mg2+ and Ca2+ concentration. It also enhanced the antioxidative system including ascorbic acid, phenylalanine ammonia-lyase, polyphenol oxidase and glutathione peroxidase, which scavenged the ROS (H2 O2 and O2 •- ) and lowered the oxidative damages (malondialdehyde level) under salinity in roots and more specifically in leaf tissues. The photosynthetic rate and stomatal conductance consequently improved (16 and 21%, respectively) in salt-stressed quinoa PBZ-treated compared to the non-treated ones and contributed to the improvement of panicle length (33%), 100-grain weight (8%) and grain yield (38%). Therefore, PBZ can be opted as a shotgun approach to improve quinoa performance and other crops under high saline conditions.

Isolation and characterization of the betalain biosynthesis gene involved in hypocotyl pigmentation of the allotetraploid Chenopodium quinoa.

In quinoa seedlings, the pigment betalain accumulates in the hypocotyl. To isolate the genes involved in betalain biosynthesis in the hypocotyl, we performed ethyl methanesulfonate (EMS) mutagenesis on the CQ127 variety of quinoa seedlings. While putative amaranthin and celosianin II primarily accumulate in the hypocotyls, this process produced a green hypocotyl mutant (ghy). This MutMap+ method using the quinoa draft genome revealed that the causative gene of the mutant is CqCYP76AD1-1. Our results indicated that the expression of CqCYP76AD1-1 was light-dependent. In addition, the transient expression of CqCYP76AD1-1 in Nicotiana benthamiana leaves resulted in the accumulation of betanin but not isobetanin, and the presence of a polymorphism in CqCYP76A1-2 in the CQ127 variety was shown to have resulted in its loss of function. These findings suggested that CqCYP76AD1-1 is involved in betalain biosynthesis during the hypocotyl pigmentation process in quinoa. To our knowledge, CqCYP76AD1-1 is the first quinoa gene identified by EMS mutagenesis using a draft gene sequence.

Differential response of quinoa genotypes to drought and foliage-applied H2O2 in relation to oxidative damage, osmotic adjustment and antioxidant capacity.

Quinoa (Chenopodium quinoa Willd.), a highly nutritious grain crop, is resistant to abiotic stresses (drought, salinity, and cold) and offers an alternate crop to endure harsh environmental conditions under the face of climate change. Naturally, quinoa genome displays a wide degree of variabilities in drought tolerance strategies. Therefore, the present study was designed to investigate drought tolerance variations and stress tolerance enhancement in four quinoa genotypes (Pichaman, Colorado-407D, IESP and 2-Want) thorough foliage-applied H2O2 with the purpose of identifying suitable genotype for water limited environments. The plants were exposed to two watering regimes (75% and 30% pot WHC) and foliage-applied H2O2 treatments (15 mM). The drought stress significantly reduced plant growth, relative water contents, chlorophyll and carotenoids contents and increased ROS production (H2O2 and O2•-) resulting in higher oxidative damage in all quinoa genotypes. Besides, drought stress significantly enhanced the antioxidants (SOD, PPO, and PAL) activity, total soluble sugars, proline, AsA contents and increased the total accumulation of measured inorganic ions in all quinoa genotypes. The PCA analysis indicated that parameters related to osmotic adjustment and antioxidant capacity were more pronounced in 2-Want and IESP genotypes, while parameters depicting oxidative damage were higher in Colorado-407D and more specifically in Pichaman. However, foliage-applied H2O2 effectively improved the osmolytes accumulation, antioxidants activity and K+/Na+ ratio which increased water relations, reduced lipid peroxidation and ultimately resulted in higher plant growth. Overall, 2-Want and IESP genotypes were found relatively more drought resistant, while exogenous application of H2O2 can be opted for more improvement in osmotic adjustment and antioxidant system, which may further enhance drought tolerance, even in sensitive genotypes of quinoa, such as Pichaman.

Epidermal bladder cells confer salinity stress tolerance in the halophyte quinoa and Atriplex species.

Epidermal bladder cells (EBCs) have been postulated to assist halophytes in coping with saline environments. However, little direct supporting evidence is available. Here, Chenopodium quinoa plants were grown under saline conditions for 5 weeks. One day prior to salinity treatment, EBCs from all leaves and petioles were gently removed by using a soft cosmetic brush and physiological, ionic and metabolic changes in brushed and non-brushed leaves were compared. Gentle removal of EBC neither initiated wound metabolism nor affected the physiology and biochemistry of control-grown plants but did have a pronounced effect on salt-grown plants, resulting in a salt-sensitive phenotype. Of 91 detected metabolites, more than half were significantly affected by salinity. Removal of EBC dramatically modified these metabolic changes, with the biggest differences reported for gamma-aminobutyric acid (GABA), proline, sucrose and inositol, affecting ion transport across cellular membranes (as shown in electrophysiological experiments). This work provides the first direct evidence for a role of EBC in salt tolerance in halophytes and attributes this to (1) a key role of EBC as a salt dump for external sequestration of sodium; (2) improved K+ retention in leaf mesophyll and (3) EBC as a storage space for several metabolites known to modulate plant ionic relations.

Oxidative stress and antioxidant responses to increasing concentrations of trivalent chromium in the Andean crop species Chenopodium quinoa Willd.

Quinoa (Chenopodium quinoa Willd), an ancient Andean seed crop, exhibits exceptional nutritional properties and resistance to abiotic stress. The species' tolerance to heavy metals has, however, not yet been investigated nor its ability to take up and translocate chromium (Cr). This study aimed to investigate the metabolic adjustments occurring upon exposure of quinoa to several concentrations (0.01-5mM) of CrCl3. Young hydroponically grown plants were used to evaluate Cr uptake, growth, oxidative stress, and other biochemical parameters three and/or seven days after treatment. Leaves accumulated the lowest amounts of Cr, while roots and stems accumulated the most at low and at high metal concentrations, respectively. Fresh weight and photosynthetic pigments were reduced only by the higher Cr(III) doses. Substantially increased lipid peroxidation, hydrogen peroxide, and proline levels were observed only with 5mM Cr(III). Except for a significant decrease at day 7 with 5mM Cr(III), total polyphenols and flavonoids maintained control levels in Cr(III)-treated plants, whereas antioxidant activity increased in a dose-dependent manner. Maximum polyamine accumulation was observed in 1mM CrCl3-treated plants. Even though α- and γ-tocopherols also showed enhanced levels only with the 1mM concentration, tyrosine aminotransferase (TAT, EC 2.6.1.5) activity increased under Cr(III) treatment in a dose- and time-dependent manner. Taken together, results suggest that polyamines, tocopherols, and TAT activity could contribute to tolerance to 1mM Cr(III), but not to the highest concentration that, instead, generated oxidative stress.

Silver nanoparticles enhance the mitigation of osmotic stress in Chenopodium quinoa microshoots grown under in vitro osmo-stressing conditions.

Osmotic stress is one of the main destructive abiotic factors that hinder plant growth and development. In this research, the role of silver nanoparticles (Ag NPs) in mitigating the negative impact of osmotic stress on in vitro grown Chenopodium quinoa (Quinoa 6 Line; Q6) was investigated to determine whether Ag NPs were able to reduce the negative effects on the in vitro grown cultures of the Q6 line. The explants were subcultured onto a special osmostressing media containing sucrose, sorbitol, or mannitol at different levels (0.1, 0.2, 0.3, and 0.4 mol/L) to mimic the osmotic stressing environment for four weeks. Then, stress physiological responses of in vitro grown Q6 under the induced osmotic stress were investigated to determine the highest stress level that the microshoots could tolerate. Next, Ag NPs; 25, 50, and 75 mg/L were added to the medium that contained the highest stress level of the induced osmotic stress to determine if their addition improved the physiological performance of the Q6 microshoots under the most severe osmotic agent levels. The results revealed that 0.4 mol/L sucrose, 0.3 mol/L sorbitol, and 0.3 mol/L mannitol were the highest stress levels that the microshoots could tolerate. The addition of 75 mg/L Ag NPs to the previous highest stress levels resulted in a significant increase in the following: stem length (SL), leaves number (LN), fresh weight (FW), dry weight (DW), total chlorophyll, protein, calcium (Ca), and phosphorus (P) contents, while it caused a reduction in proline, sodium (Na) ions, and potassium (K) ions. These results indicate that the negative consequences of osmotic stress on Q6 quinoa microshoots could be mitigated by adding specific concentrations of Ag NPs to the culture medium.

Multivariate characterization of salicylic acid and potassium induced physio-biochemical and phytoremediation responses in quinoa exposed to lead and cadmium contamination.

The levels of soils pollutants such as lead (Pb) and cadmium (Cd) have significantly increased recently resulting in ecological disturbances and threatening crop production. Various amendments have been employed to enhance the tolerance of crops to withstand Cd and Pb stresses. However, the role of combined application of potassium (K) and of salicylic acid (SA) for Cd and Pb stress mitigation and phytoremediation by quinoa (Chenopodium quinoa Willd) has not been comprehended well. In the present study, the effect of 10 mM K and 0.1 mM SA was tested on the quinoa plants subjected to 250 µM Pb and/or 100 µM Cd. The Pb and Cd treatments were applied separately or together. Phytotoxicity induced by Pb and Cd resulted in drastic decrease (>60%) in chlorophyll contents, stomatal conductance, and plant biomass. The collective treatment of Pb and Cd induced an increase in the concentration of hydrogen peroxide (13-fold) and lipid peroxidation (16-fold) that resulted in a 61% reduction in membrane stability. The application of 10 mM K and/or 0.1 mM SA was remarkable in mitigating the adverse effect of Pb and Cd. The reduction in plant biomass was 17% when 10 mM K and 0.1 mM SA were applied together under the combined treatment of both the metals. The simultaneous application of K and SA effectively mitigated oxidative stress by enhancing the activities of superoxide dismutase, peroxidase, ascorbate peroxidase, and catalase by 12, 10, 7 and 10-folds respectively. The positive effect of K and SA on these attributes resulted in a remarkable reduction in metal accumulation and translocation and lipid peroxidation. The stressed plants supplemented with K and SA exhibited a significant improvement in the membrane stability index, chlorophyll content, and stomatal conductance. This study concluded that the combined application of K and SA could be a good approach for reducing Pb and Cd phytotoxicity in quinoa and enhancing their phytostabilization potential in the contaminated soils.

Effect of exogenous γ-aminobutyric acid on physiological property, antioxidant activity, and cadmium uptake of quinoa seedlings under cadmium stress.

Increasing cadmium (Cd) pollution has negative effects on quinoa growth and production. Gamma-aminobutyric acid (GABA) confers plants with stress resistance to heavy metals; however, the mechanism remains unclear. We explored the effects of exogenous GABA on the physiological characteristics, antioxidant capacity, and Cd accumulation of quinoa seedlings under Cd stress using hydroponic experiments. Partial least-squares regression was used to identify key physical and chemical indices of seedlings affecting Cd accumulation. Compared with those of the CK group, exposure to 10 and 25 µmol·L-1 Cd significantly reduced the photosynthetic pigment contents, photosynthesis, and biomass accumulation of quinoa seedlings; resulted in shorter and thicker roots; decreased the length of the lateral roots; decreased the activities of superoxide dismutase (SOD) and peroxide (POD); and increased H2O2 and malondialdehyde (MDA) contents. Exogenous GABA reduced the Cd content in the stem/leaves and roots of quinoa seedlings under Cd stress by 13.22-21.63% and 7.92-28.32%, decreased Cd accumulation by 5.37-6.71% and 1.91-4.09%, decreased the H2O2 content by 38.21-47.46% and 45.81-55.73%, and decreased the MDA content by 37.65-48.12% and 29.87-32.51%, respectively. GABA addition increased the SOD and POD activities in the roots by 2.78-5.61% and 13.81-18.33%, respectively, under Cd stress. Thus, exogenous GABA can reduce the content and accumulation of Cd in quinoa seedlings by improving the photosynthetic characteristics and antioxidant enzyme activity and reducing the degree of lipid peroxidation in the cell membrane to alleviate the toxic effect of Cd stress on seedling growth.

Differential activity of plasma and vacuolar membrane transporters contributes to genotypic differences in salinity tolerance in a Halophyte Species, Chenopodium quinoa.

Halophytes species can be used as a highly convenient model system to reveal key ionic and molecular mechanisms that confer salinity tolerance in plants. Earlier, we reported that quinoa (Chenopodium quinoa Willd.), a facultative C3 halophyte species, can efficiently control the activity of slow (SV) and fast (FV) tonoplast channels to match specific growth conditions by ensuring that most of accumulated Na+ is safely locked in the vacuole (Bonales-Alatorre et al. (2013) Plant Physiology). This work extends these finding by comparing the properties of tonoplast FV and SV channels in two quinoa genotypes contrasting in their salinity tolerance. The work is complemented by studies of the kinetics of net ion fluxes across the plasma membrane of quinoa leaf mesophyll tissue. Our results suggest that multiple mechanisms contribute towards genotypic differences in salinity tolerance in quinoa. These include: (i) a higher rate of Na+ exclusion from leaf mesophyll; (ii) maintenance of low cytosolic Na+ levels; (iii) better K+ retention in the leaf mesophyll; (iv) a high rate of H+ pumping, which increases the ability of mesophyll cells to restore their membrane potential; and (v) the ability to reduce the activity of SV and FV channels under saline conditions. These mechanisms appear to be highly orchestrated, thus enabling the remarkable overall salinity tolerance of quinoa species.

Biochar reduces copper toxicity in Chenopodium quinoa Willd. In a sandy soil.

Mining, smelting, land applications of sewage sludge, the use of fungicides containing copper (Cu), and other human activities have led to widespread soil enrichment and contamination with Cu and potentially toxic conditions. Biochar (BC) can adsorb several substances, ranging from herbicides to plant-inhibiting allelochemicals. However, the range of potential beneficial effects on early-stage plant growth with regard to heavy metal toxicity is largely unexplored. We investigated the ameliorating properties of a forestry-residue BC under Cu toxicity conditions on early plant growth. Young quinoa plants () were grown in the greenhouse in the presence of 0, 2, and 4% BC application (w/w) added to a sandy soil with 0, 50, or 200 µg g Cu supplied. The plants without BC showed severe stress symptoms and reduced growth shortly after Cu application of 50 µg g and died at 200 µg Cu g. Increasing BC concentrations in the growth medium significantly increased the plant performance without Cu toxicity or under Cu stress. At the 4% BC application rate, the plants with 200 µg g Cu almost reached the same biomass as in the control treatment. In the presence of BC, less Cu entered the plant tissues, which had reduced Cu concentrations in the order roots, shoots, leaves. The amelioration effect also was reflected in the plant-soil system CO gas exchange, which showed clear signs of improvement with BC presence. The most likely ameliorating mechanisms were adsorption of Cu to negatively charged BC surfaces and an improvement of the water supply. Overall, BC seems to be a beneficial amendment with the potential to ameliorate Cu toxicity in sandy soils. Further research with a broad spectrum of different soil types, BCs, and crop plants is required.

Antiphytoviral activity of sesquiterpene-rich essential oils from four croatian teucrium species.

The purpose of this study was to compare the essential oil profiles of four Croatian Teucrium species (Lamiaceae), as determined by GC and GC/MS, with their antiphytoviral efficiency. A phytochemical analysis showed that T. polium, T. flavum, T. montanum and T. chamaedrys are characterized by similar essential oil compositions. The investigated oils are characterized by a high proportion of the sesquiterpene hydrocarbons ß-caryophyllene (7.1-52.0%) and germacrene D (8.7-17.0%). Other important components were ß-pinene from T. montanum and α-pinene from T. flavum. The investigated essential oils were proved to reduce lesion number in the local host Chenopodium quinoa Willd. infected with Cucumber Mosaic Virus (CMV), with reductions of 41.4%, 22.9%, 44.3% and 25.7%, respectively.

Exogenous application of moringa leaf extract improves growth, biochemical attributes, and productivity of late-sown quinoa.

Quinoa (Chenopodium quinoa Willd.) has gained significant popularity among agricultural scientists and farmers throughout the world due to its high nutritive value. It is cultivated under a range of soil and climatic conditions; however, late sowing adversely affects its productivity and yield due to shorter growth period. Inorganic and organic phyto-stimulants are promising for improving growth, development, and yield of field crops under stressful environments. Field experiments were conducted during crop cultivation seasons of 2016-17 and 2017-18, to explore the role of inorganic (hydrogen peroxide and ascorbic acid) and organic [moringa leaf extract (MLE) and sorghum water extract (sorgaab)] phyto-stimulants in improving growth and productivity of quinoa (cultivar UAF-Q7). Hydrogen peroxide at 100 µM, ascorbic acid at 500 µM, MLE at 3% and sorgaab at 3% were exogenously applied at anthesis stage of quinoa cultivated under normal (November 21st and 19th during 2016 and 2017) and late-sown (December 26th and 25th during 2016 and 2017) conditions. Application of inorganic and organic phyto-stimulants significantly improved biochemical, physiological, growth and yield attributes of quinoa under late sown conditions. The highest improvement in these traits was recorded for MLE. Application of MLE resulted in higher chlorophyll a and b contents, stomatal conductance, and sub-stomatal concentration of CO2 under normal and late-sowing. The highest improvement in soluble phenolics, anthocyanins, free amino acids and proline, and mineral elements in roots, shoot and grains were observed for MLE application. Growth attributes, including plant height, plant fresh weight and panicle length were significantly improved with MLE application as compared to the rest of the treatments. The highest 1000-grain weight and grain yield per plant were noted for MLE application under normal and late-sowing. These findings depict that MLE has extensive crop growth promoting potential through improving physiological and biochemical activities. Hence, MLE can be applied to improve growth and productivity of quinoa under normal and late-sown conditions.

Silicon mitigates nutritional stress in quinoa (Chenopodium quinoa Willd.).

Nutritional deficiency is common in several regions of quinoa cultivation. Silicon (Si) can attenuate the stress caused by nutritional deficiency, but studies on the effects of Si supply on quinoa plants are still scarce. Given this scenario, our objective was to evaluate the symptoms in terms of tissue, physiological and nutritional effects of quinoa plants submitted to nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), and magnesium (Mg) deficiencies under Si presence. The experiment consisted of a factorial scheme 6 × 2, using a complete solution (CS), -N, -P, -K, -Ca, -Mg combined with absence and presence of Si (1.5 mmol L-1). Symptomatic, physiological, nutritional and evaluation vegetative were performed in quinoa crop. The deficiencies of N, P, K, Ca and Mg in quinoa cultivation caused visual symptoms characteristic of the deficiency caused by respective nutrients, hence decreasing the plant dry mass. However, Si supply attenuated the deficiency effects by preserving the photosynthetic apparatus, increasing the chlorophyll production, increasing the membrane integrity, and decreasing the electrolyte leakage. Thus, the Si supply attenuated the visual effects provided by deficiency of all nutrients, but stood out for N and Ca, because it reflected in a higher dry mass production. This occurred because, the Si promoted higher synthesis and protection of chlorophylls, and lower electrolyte leakage under Ca restriction, as well as decreased electrolyte leakage under N restriction.

Synergistic consequences of salinity and potassium deficiency in quinoa: Linking with stomatal patterning, ionic relations and oxidative metabolism.

Quinoa emerged as an ideal food security crop due to its exceptional nutritive profile and stress enduring potential and also deemed as model plant to study the salt-tolerance mechanisms. However to fill the research gaps of this imperative crop, the present work aimed to study the effect of potassium (K) deficiency either separately or in combination with salinity. First, we investigated the stomatal and physiological based variations in quinoa growth under salinity and K, then series of analytical tools were used with model approach to interpret the stomatal aperture (SA) and photosynthesis (Pn) changes. Results revealed that quinoa efficiently deployed antioxidants to scavenge the excessive reactive oxygen species (ROS), had high uptake and retention of K+, Ca2+, Mg2+ with Cl⁻ as charge balancing ion, increased stomata density (SD) and declined the SA to maintain the Pn which resulted the improved growth under salinity. Whereas, K-deficiency caused the stunted growth more severally under salinity due to disruption in ionic homeostasis, excessive ROS production elicited the oxidative damages, SD and SA reduced and ultimately declined in Pn. Our best fitted regression model explored that dependent variables like Pn and SA changed according to theirs signified explanatory variables with quantification per unit based as stomatal conductance (Gs, 51), SD (0.05), ROS (-0.79) and K+ (0.08), Cl⁻ (0.34) and Na+ (- 0.52) respectively. Overall, moderate salinity promoted the quinoa growth, while K-deficiency particularly with salinity reduced the quinoa performance by affecting stomatal and non-stomatal factors.

Cross-talk between nitric oxide, hydrogen peroxide and calcium in salt-stressed Chenopodium quinoa Willd. At seed germination stage.

Seed germination is critical for successful crop production and this growth stage can be very sensitive to salt stress depending on the plant's tolerance mechanisms. The pretreatment of Chenopodium quinoa (quinoa) seeds with CaCl2, H2O2 and sodium nitroprusside (SNP) limited the adverse effect of salt stress on seed germination. The pre-treated seeds showed a significant increase in germination rate, relative germination rate and germination index while the mean germination time was significantly reduced under both optimal and stress conditions. In parallel with seed germination, the negative effect of salt stress on the activity of α-amylase and ß-amylase was reduced in pre-treated seeds. The amylase enzymes are responsible for starch hydrolysis, so the reduction of amylase activity by salt stress resulted in higher starch content in the seeds and lower concentrations of water-soluble sugars such as glucose. Pretreatment stimulated amylase activity resulting in starch breakdown and increased content of water-soluble sugars in the salt-stressed seeds. Protein and amino acid contents were significantly enhanced in salt-stressed seeds, which were highlighted in pre-treated seeds. The findings of this study demonstrate that pretreatment of quinoa seeds with CaCl2, H2O2 and SNP at 5, 5 and 0.2 mM, respectively, concentration to achieve rapid germination at high levels under optimal and salt-stress conditions.

Low-temperature effect on enzyme activities involved in sucrose-starch partitioning in salt-stressed and salt-acclimated cotyledons of quinoa (Chenopodium quinoa Willd.) seedlings.

The effect of low temperature on growth, sucrose-starch partitioning and related enzymes in salt-stressed and salt-acclimated cotyledons of quinoa (Chenopodium quinoa Willd.) was studied. The growth of cotyledons and growing axes in seedlings grown at 25/20 degrees C (light/dark) and shifted to 5/5 degrees C was lower than in those only growing at 25/20 degrees C (unstressed). However, there were no significant differences between low-temperature control and salt-treated seedlings. The higher activities of sucrose phosphate synthase (SPS, EC 2.4.1.14) and soluble acid invertase (acid INV, EC 3.2.1.25) were observed in salt-stressed cotyledons; however, the highest acid INV activity was observed in unstressed cotyledons. ADP-glucose pyrophosphorylase (ADP-GPPase, EC 2.7.7.27) was higher in unstressed cotyledons than in stressed ones. However, between 0 and 4days the highest value was observed in salt-stressed cotyledons. The lowest value of ADP-GPPase was observed in salt-acclimated cotyledons. Low temperature also affected sucrose synthase (SuSy, EC 2.4.1.13) activity in salt-treated cotyledons. Sucrose and glucose were higher in salt-stressed cotyledons, but fructose was essentially higher in low-temperature control. Starch was higher in low-temperature control; however, the highest content was observed at 0day in salt-acclimated cotyledons. Results demonstrated that low temperature induces different responses on sucrose-starch partitioning in salt-stressed and salt-acclimated cotyledons. Data also suggest that in salt-treated cotyledons source-sink relations (SSR) are changed in order to supply soluble sugars and proline for the osmotic adjustment. Relationships between starch formation and SuSy activity are also discussed.