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Since regulatory roles of calcium (Ca) and melatonin (MT) in physiological responses of plants to salinity stress are lacking, various Dracocephalum kotschyi genotypes (Bojnord, Urmia, Fereydunshahr, and Semirom) were pretreated with exogenous Ca (5 mM), MT (100 µM), and Ca + MT in the presence of salt (75 mM NaCl). In addition measuring the concentration of phenolic compounds by high-performance liquid chromatography (HPLC), histochemical evaluations of essential oils and phenolic compounds in glandular trichomes of leaf samples was performed by light microscope. Salt stress reduced shoot fresh (SFW) and dry weight (SDW), leaf area (LA), relative water content (RWC), and maximum efficiency of photosystem II (Fv /Fm ), but enhanced total phenolic content (TPC) and total flavonoids content (TFC), phenolic compounds concentrations, 2,2-diphenylpicrylhydrazyl (DPPH) radical scavenging capacity, electrolyte leakage (EL), proline, and hydrogen peroxide (H2 O2 ) concentrations, and Na+ /K+ and essential oils and TPC of the glandular trichomes of leaves in all D. kotschyi genotypes. Foliar spraying of Ca, MT, and particularly Ca + MT on D. kotschyi seedlings improved SFW, SDW, RWC, TPC, TFC, proline, and phenolic compounds concentrations, Fv /Fm , and DPPH radical scavenging capacity, but reduced H2 O2 , EL, and Na+ /K+ in the leaves and essential oils and TPC in the glandular trichomes of all genotypes under both nonstress and salt stress conditions. These findings indicate that the crosstalk between MT and Ca synergistically improves salt tolerance, TPC and TFC, phenolic compounds concentration, and essential oils accumulation in glandular trichomes of different D. kotschyi genotypes.

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In order to address the lacuna of data on the response of physiological and biochemical attributes and chemical compounds of glandular trichomes of Iranian borage (Echium amoenum Fisch. & C.A.Mey.) to saline water (0, 25, 50, 75, and 100 mM NaCl) an experiment was conducted on 13 genotypes. Genotypic differences and salt-induced modifications in chlorophyll concentration and fluorescence, plant growth, relative water content, proline concentration, antioxidant defense, and chemical compounds of glandular trichomes upon exposure to salt stress were observed. Chlorophyll and carotenoids concentrations and catalase (EC 1.11.1.6) and ascorbate peroxidase (EC 1.11.1.11) activities were either enhanced or remained unchanged in the presence of moderate salt concentrations (i.e. 25 and 50 mM NaCl) in a majority of the genotypes. Though, 75 and 100 mM NaCl were modestly and severely detrimental, respectively, to the majority of the genotypes. The 75 and 100 mM NaCl led to substantial increases and decreases in the Na+ and K+, respectively, resulting in notable increase in the Na+/K+. Increases in proline, total phenolic compounds, and alkaloids concentrations, essential oils, alkaloids, and phenolic compounds of the glandular trichomes were concomitant to decreases in the relative water content, leaf area, maximum quantum efficiency of photosystem II, shoot and root dry masses. This study revealed, for the first time, that Iranian borage tolerates 25 and 50 mM NaCl and antioxidative enzymes as well as secondary metabolites such as alkaloids and phenolic compounds accumulated mainly in the trichomes play key role in this regard.

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This study investigated: (1) the effects of various concentrations of melatonin (MT) and Ca2+; and (2) the impact of crosstalk between these signal molecules on photosynthesis and salt tolerance of Dracocephalum kotschyi Boiss. Results indicated that 5mM CaCl2, as well as 100µM MT were the best concentrations for increasing shoot dry weight, leaf area, SPAD index, maximum quantum efficiency of photosystem II (Fv/Fm), and decreasing malondialdehyde content under salinity stress. The impact of MT on growth and photosynthesis was closely linked to its effect on enhancing antioxidant enzyme activities in leaves. Application of p-chlorophenylalanine, as an inhibitor of MT biosynthesis, negated the impacts of MT on the aforementioned attributes. Salinity and MT boosted cytosolic Ca2+ concentration. Exogenous MT, as well as Ca2+, enhanced tolerance index, membrane stability, leaf area, the content of chlorophyll (Chl) a, Chl b, and carotenoids (Car), Fv/Fm, and stomatal conductance under salinity stress. These impacts of MT were eliminated by applying a calmodulin antagonist, a Ca2+ chelator and a Ca2+ channel blocker. These novel findings indicate that the MT-induced effects on photosynthetic parameters and salt-evoked oxidative stress were mediated through calcium/calmodulin (Ca2+/CaM) signalling.

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In this study, the role of calcium/calmodulin (Ca2+/CaM) and melatonin (Mel) as two signal molecules in inducing systemic salt tolerance of Dracocephalum kotschyi Boiss. was investigated. Salinity stress (100 mM NaCl) reduced plant growth and induced ionic, osmotic, and oxidative damages in D. kotschyi leaves. Detection of cytosolic free Ca2+ ([Ca2+]cyt) by the Fura-2 method and the measurement of endogenous Mel by GC-MS demonstrated that salinity induced Ca2+ burst and increased endogenous Mel content in D. kotschyi leaves. Root pretreatment with 5 mM Ca2+ or 100 µM Mel recovered plant growth, reduced leaf electrolytic leakage, H2O2, and MDA contents and improved membrane integrity not only at the application site (roots), but also at the untreated distal parts (leaves) under salt stress. Rhizospheric treatment with Mel and Ca2+ triggered systemic tolerance in D. kotschyi, as judged from improving RWC, increasing proline content, modulating Na+, K+, and Ca2+ homeostasis, and enhancing the activities of SOD, CAT, APX, and POD in the leaves of salt-stressed plants. Mel augmented [Ca2+]cyt, but the rhizospheric application of Ca2+ antagonists impaired the latter responses. Furthermore, root pretreatment with Ca2+ increased Mel content, but the application of p-chlorophenylalanine (as an inhibitor of Mel biosynthesis) decreased the above attributes in the leaves of Ca2+-treated plants, leading to an arrest in the Ca2+-induced systemic salt tolerance. These novel results suggest that interaction of Ca2+/CaM and Mel is involved in overcoming salt-induced ionic, osmotic, and oxidative damages and Ca2+ and Mel may act as long-distance signals for inducing systemic salt tolerance in D. kotschyi.

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Melatonin (Mel) and calcium (Ca2+) have a regulatory role in the induction of specialized metabolites production and defensive responses against stresses. Therefore, in this study, the effects of Mel and Ca2+ and the possible relationship between them in the increase of the production of phenolic compounds in Dracocephalum kotschyi Boiss. under both control and salinity stress conditions were investigated. The results showed that 75 mM NaCl reduced shoot dry biomass but elevated H2O2 content, electrolyte leakage (EL) level, total phenolic and flavonoid contents (TPC and TFC), and DPPH scavenging capacity. Salinity stress also upregulated gene expression of phenylalanine ammonia-lyase (PAL) and rosmarinic acid synthase (RAS), as well as the activities of PAL and tyrosine ammonia-lyase (TAL) enzymes. Pre-treatment of the plants with CaCl2 and Mel affected these attributes in a dose-dependent manner. Application of 5 mM Ca2+ and 100 µM Mel improved shoot dry biomass and reduced the level of EL and H2O2 content but enhanced TPC and TFC, DPPH scavenging capacity, PAL and TAL activities, PAL and RAS transcripts, and content of rosmarinic acid (RA), luteolin flavone (LF) and apigenin flavone (AF) under salinity stress. Pre-treatment of D. kotschyi with lanthanum chloride (LaCl3) as a plasma membrane channel blocker, ethylene glycol tetra-acetic acid (EGTA) as a Ca2+ chelator and trifluoperazine (TFP) as a calmodulin (CaM) antagonist, impaired Mel effects on the above attributes under salinity stress. In contrast, pre-treatment with p-chlorophenylalanine (p-CPA), as an inhibitor of Mel biosynthesis, did not impair the impacts of Ca2+ on the production of phenolic compounds in salt-exposed plants. These results suggested that the effect of Mel on the induction of phenolic compounds production requires the influx of extracellular Ca2+ into the cells and is dependent on Ca2+/CaM signaling.

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To examine the physiological role of hexose transporters in determining the sink strength of individual fruits, the regulation of hexose transporters gene expression was studied when the sink/source ratio was artificially altered under the greenhouse condition; this was done in two cultivars of tomato, i.e. Grandella and Isabella. The sink/source ratio treatments included: saving one fruit per truss (1F), two fruits per truss (2F), three fruits per truss (3F) and no fruit pruning (control). The results showed that fruit thinning could increase starch, sucrose, and hexose contents in the fruits; it could also modulate the activity of the key enzymes and the expression of tomato hexose transporter genes (LeHTs). Based on the relative transcript levels, all examined LeHTs were unregulated at the end of cell division and the cell expansion stage of fruit development, but the strongest expression level observed at the onset of ripening was related to LeHT1 and LeHT2. Given the concomitancy of cell wall invertase (EC 3.2.1.26) activity and the LeHTs relative expression cell wall, invertase activity seemed to be involved in the expression level of LeHTs. The increased trends of the LeHTs expression with the decrease of the sink/source ratio confirmed the role of hexose transporters in determining the sink strength of the tomato fruits.

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Cadmium (Cd) can stress plants by affecting various physiological functions. Cd stress-response mechanisms were investigated in two genotypes of domesticated safflower (Carthamus tinctorius) and a population of wild safflower (Carthamus oxycantous) to explore potential differences in tolerance mechanisms of these species. A hydroponic experiment was conducted with 6-day-old safflower plants. Genotypes AC-Sterling (tolerant) and Saffire (semi-tolerant) from C. tinctorius, and Arak (sensitive) a population from C. oxycantouswere subjected to three concentrations of Cd (i.e., 0, 1, and 20 µM CdCl2). Genotypic differences were detected in Cdtolerance index, Cd concentration in shoots and roots, Cd translocation to shoots, Cd bound to cell walls, superoxide dismutase (SOD) activity, lipid peroxidation, and phytochelatins accumulation in safflower plants upon exposure to CdCl2. Results indicate that genotypic differences were more obvious in the presence of low (i.e., 1 µM) rather than high (i.e., 20 µM) CdCl2 concentrations. Comparing genotypes, root and shoot Cd accumulation was highest in the semi-tolerant genotype. Cadmium translocation to shoots was increased with increasing tolerance. The percentage of Cd bound to root cell walls was higher in the tolerant genotype, but only with low CdCl2 addition. Furthermore, in the tolerant genotype, SOD activity was lowest in both roots and shoots with low CdCl2 addition but highest with high CdCl2 addition, while the opposite was found for phytochelatins. Lipid peroxidation was decreased with Cd tolerance at both CdCl2 concentrations. We conclude that safflower relies mainly on binding Cd to the cell walls and the formation of phytochelatins in root and shoot tissues, in order to handle the Cd stress, evidenced by lessening Cd-induced lipid peroxidation.

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