The influence of salinity on cell ultrastructures and photosynthetic apparatus of barley genotypes differing in salt stress tolerance

A hydroponic experiment was conducted to elucidate the difference in growth and cell ultrastructure between Tibetan wild and cultivated barley genotypes under moderate (150 mM NaCl) and high (300 mM NaCl) salt stress. The growth of three barley genotypes was reduced significantly under salt stress, but the wild barley XZ16 (tolerant) was less affected relative to cultivated barley Yerong (moderate tolerant) and Gairdner (sensitive). Meanwhile, XZ16 had lower Na⁺ and higher K⁺ concentrations in leaves than other two genotypes. In terms of photosynthetic and chlorophyll fluorescence parameters, salt stress reduced maximal photochemical efficiency (F _v/F _m), net photosynthetic rate (Pn), stomatal conductance (Gs), and intracellular CO₂ concentration (Ci). XZ16 showed relatively smaller reduction in comparison with the two cultivated barley genotypes. The observation of transmission electron microscopy found that fundamental cell ultrastructure changes happened in both leaves and roots of all barley genotypes under salt NaCl stress, with chloroplasts being most changed. Moreover, obvious difference could be detected among the three genotypes in the damage of cell ultrastructure under salt stress, with XZ16 and Gairdner being least and most affected, respectively. It may be concluded that high salt tolerance in XZ16 is attributed to less Na⁺ accumulation and K⁺ reduction in leaves, more slight damage in cell ultrastructure, which in turn caused less influence on chloroplast function and photosynthesis.     

Effect of salt stress on photosynthesis and physiological parameters of three contrasting barley genotypes

In order to understand the physiological traits important in conferring salt tolerance in three barley genotypes, this study was performed under field conditions with three water salinity levels (2, 10, and 18 dS m–1). High salinity decreased net photosynthetic rate, transpiration rate, and stomatal conductance, K⁺ concentration, K⁺:Na⁺ ratio, and grain yield, but increased electrolyte leakage and Na⁺ content. Under 10 and 18 dS m–1 salinity, Khatam (salt-tolerant) had the maximum stomatal conductance, K⁺, K⁺:Na⁺ ratio, and the grain yield, and a minimum Na⁺ content and electrolyte leakage, whereas Morocco (salt-sensitive) had the lowest net photosynthetic rate, stomatal conductance, K⁺ content, K⁺:Na⁺ ratio, and grain yield, and the highest Na⁺ content and electrolyte leakage. This study showed that tolerant genotypes of barley may avoid Na⁺ accumulation in aboveground parts, facilitating a higher photosynthetic rate and higher grain yield.     

Infection of Barley by <Emphasis Type="Italic">Ramularia</Emphasis> <Emphasis Type="Italic">collo-cygni</Emphasis>: Scanning Electron Microscopic Investigations

Ramularia collo-cygni causes leaf spots on barley (Hordeum vulgare), a disease of growing economical importance. Scanning electron microscopy was used to study the life cycle of the fungus on barley during the vegetation period and in winter. The infectious stage started with conidium germination on the surface and the penetration into the leaf via the stomatal pore where the hyphae grew within the cells that became necrotic. The conidiophores emerged through the stomatal pore. On older leaves, however, they frequently emerged apart from it and the results suggested a pushing apart of adjacent cell walls of the epidermal cells. An assessment of the amount of conidium formation of one heavily infested barley plant resulted in 4.05 × 10⁶ conidia per plant. For the first time, conidiophores, conidium production and germination of conidia were also observed in winter on barley and on maize leaves.     

Distinct responses to ozone of abaxial and adaxial stomata in three Euramerican poplar genotypes

Ozone induces stomatal sluggishness, which impacts photosynthesis and transpiration. Stomatal responses to variation of environmental parameters are slowed and reduced by ozone and may be linked to difference of ozone sensitivity. Here we determine the ozone effects on stomatal conductance of each leaf surface. Potential causes of this sluggish movement, such as ultrastructural or ionic fluxes modification, were studied independently on both leaf surfaces of three Euramerican poplar genotypes differing in ozone sensitivity and in stomatal behaviour. The element contents in guard cells were linked to the gene expression of ion channels and transporters involved in stomatal movements, directly in microdissected stomata. In response to ozone, we found a decrease in the stomatal conductance of the leaf adaxial surface correlated with high calcium content in guard cells compared with a slight decrease on the abaxial surface. No ultrastructural modifications of stomata were shown except an increase in the number of mitochondria. The expression of vacuolar H⁺/Ca²⁺‐antiports (CAX1 and CAX3 homologs), β‐carbonic anhydrases (βCA1 and βCA4) and proton H⁺‐ATPase (AHA11) genes was strongly decreased under ozone treatment. The sensitive genotype characterized by constitutive slow stomatal response was also characterized by constitutive low expression of genes encoding vacuolar H⁺/Ca²⁺‐antiports.     

Does root-sourced ABA have a role in mediating growth and stomatal responses to soil compaction in tomato (Lycopersicon esculentum)?

Isogenic wild-type (Ailsa Craig) and abscisic acid (ABA)-deficient mutant (flacca) genotypes of tomato were used to examine the role of root-sourced ABA in mediating growth and stomatal responses to compaction. Plants were grown in uniform soil columns providing low to moderate bulk densities (1.1–1.5 g cm^?3), or in a split-pot system, which allowed the roots to divide between soils of the same or differing bulk density (1.1/1.5 g cm^?3). Root and shoot growth and leaf expansion were reduced when plants were grown in compacted soil (1.5 g cm^?3) but leaf water status was not altered. However, stomatal conductance was affected, suggesting that non-hydraulic signal(s) transported in the transpiration stream were responsible for the observed effects. Xylem sap and foliar ABA concentrations increased with bulk density for 10 and 15 days after emergence (DAE), respectively, but were thereafter poorly correlated with the observed growth responses. Growth was reduced to a similar extent in both genotypes in compacted soil (1.5 g cm^?3), suggesting that ABA is not centrally involved in mediating growth in this severely limiting ‘critical’ compaction stress treatment. Growth performance in the 1.1/1.5 g cm^?3 split-pot treatment of Ailsa Craig was intermediate between the uniform 1.1 and 1.5 g cm^?3 treatments, whereas stomatal conductance was comparable to the compacted 1.5 g cm^?3 treatment. In contrast, shoot dry weight and leaf area in the split-pot treatment of flacca were similar to the 1.5 g cm^?3 treatment, but stomatal conductance was comparable to uncompacted control plants. These results suggest a role for root-sourced ABA in regulating growth and stomatal conductance during ‘sub-critical’ compaction stress, when genotypic differences in response are apparent. The observed genotypic differences are comparable to those previously reported for barley, but occurred at a much lower bulk density, reflecting the greater sensitivity of tomato to compaction. By alleviating the severe growth reductions induced when the entire root system encounters compacted soil, the split-pot approach has important applications for studies of the role of root-sourced signals in compaction-sensitive species such as tomato.     

Cell kinetics,development of stomata and some effects of colchicine in barley

Summary The developmental sequence of the formation of stomatal complexes in the leaf epidermis of barley was studied. Cell-kinetic parameters were obtained from two genotypes — Early Bonus and eceriferum-g, a mutant with an abnormal stomatal pattern. The distribution of mitotic frequencies as a function of position in the stomatal rows was analyzed at each stage of development leading to mature stomata. Regression curves obtained for each stage showed that the distributions were stage-specific. Thus, the mitotic frequencies presented similar values throughout the portion of the file where the first asymmetrical divisions take place, had a parabolic distribution for the stage of subsidiary formation, and showed a linear shape, with a negative slope, for the stage of guard-mother-cell divisions. The mutant genotype differed from the normal by having a faster rate of ordinary guard-mother-cell divisions as a function of position in the row. The higher level of subsidiary cell formation in the mutant was interpreted as a consequence of a displacement of the used markers, suggesting a precocious initiation of subsidiary-cell formation in eceriferum-g. Time estimations of the length of the cell cycles were obtained by cell-population studies after a pulse with colchicine. Eceriferum-g appeared to have slower cell cycles. The leaves treated with colchicine showed a shift in the elongation axis of the cells. Autoradiography after treatment with ³H-thymidine showed ineorporation of the label in the portions of the row proximal to all three peaks of divisions indicating that all mitoses were preceded by the usual period of DNA synthesis. Labelling of lateral cells at a mature stage suggested DNA synthesis leading to endoploidy.     

An efficient regeneration system of barley cultivars from leaf base segments

A simple and reliable regeneration system from leaf bases of barley (Hordeum vulgare L.) has been developed. The in vitro regeneration frequencies of seven commercial barley genotypes were compared using segments from the first leaves of 5-d-old seedlings. The regeneration frequency ranged from 31.56 to 72.22 % among the barley genotypes. Murashige and Skoog medium supplemented with 6-benzyladenine (0.5 mg dm⁻³) and kinetin (0.5 mg dm⁻³) was optimum for the regeneration. Longitudinal cut of the segments or the removal of coleoptiles further increased plantlet regeneration frequency.     

Effects of Infection Density on the Size of Barley and Wheat Leaf Rust Colonies before and on the Size of Uredia after the Start of Sporulation

Three barley genotypes were exposed to four different inoculum densities of barley leaf rust, Puccinia hordei. Colony area well before first sporulation and uredia size well after the start of sporulation were measured. In a second series two barley and two wheat cultivars were exposed to four different inoculum densities of barley and wheat leaf rust (P. recondita f. sp. tritici), respectively. Before the sporulation is initiated the colony size was independent of the uredia density. This was valid for densities ranging from approximately 7 to over 200 uredia per cm² leaf area. After the sporulation had started the uredia size was strongly dependent on the uredia density. The size of the uredia was approximately halved when the uredia density increased from about 10 to about 150 per cm². The urediospore production per uredium decreased much stronger with increased uredia density.     

Development of a Differential Set for the Race analysis of Puccinia hordei Otth.

Investigations for the race analysis of P. hordei (location Cologne, W.-Germany) showed, that most of the resistant genotypes in the International differential set with Pa₁ to Pa₈ were ineffective for that purpose. Out of 80 selected barley genotypes with different degree of resistance against the field-collection RG 1975 hosts with a strongly differentiating effect were chosen. This was done using three parameters: mean resistance x?w, its standard deviation x?w and the range z. They were calculated from evaluating the reaction types of 80 barley genotypes infected with 116 single-pustule lines of barley brown rust. Eight hosts with strongly differentiating reactions towards the 116 rust lines were composed into a differential set, which could characterize the heterogeneity of the Cologne P. hordei-population essentially better and more precisely than the International differential set.     

Leaf morphological plasticity and stomatal conductance in three Populus alba L. genotypes subjected to salt stress

The effect of salt stress on leaf morphology and functionality was studied in three Populus alba genotypes differing in tolerance to salinity: 6K3 (sensitive), 2AS11 (moderately tolerant), and 14P11 (tolerant). Plants were subjected to an intense and progressive salt stress from 50 to 250 mM NaCl by 50 mM steps at 10-day intervals. The micromorphological results highlighted phenotypic variation among the three genotypes already in control plants, with the genotype 14P11 having significantly smaller epidermal cells and higher stomatal density. Salt-treated plants modulated differently the expansion of stomata compared with epidermal cells. Regression analysis showed significant correlations between decrease of stomatal area and stomatal conductance (g_s) in genotypes 14P11 and 6K3. So, the common reduction of stomatal area could be an early mechanism to save water in this species. However, only genotype 14P11 showed further significant decrease of this trait under the highest salinity level, combined with a significant reduction in leaf length. In addition, this genotype showed the lowest leaf abscission rate at the end of salt stress period. The genotype 6K3 was severely affected by leaf necrosis and showed the highest leaf abscission rate in salt stress conditions. In the moderately tolerant genotype 2AS11, an intermediate plastic behaviour in both leaf morphology and physiology was observed during the experiment. The phenotypic variation among the three genotypes in terms of micromorphology and stomatal conductance is discussed in relation to plant functionality in salt stress conditions. Overall results suggest that leaf morphological habit contributes to salt tolerance in P. alba.     

Evaluation of salinity tolerance and analysis of allelic function of <Emphasis Type="Italic">HvHKT1</Emphasis> and <Emphasis Type="Italic">HvHKT2</Emphasis> in Tibetan wild barley

Tibetan wild barley is rich in genetic diversity with potential allelic variation useful for salinity-tolerant improvement of the crop. The objectives of this study were to evaluate salinity tolerance and analysis of the allelic function of HvHKT1 and HvHKT2 in Tibetan wild barley. Salinity tolerance of 189 Tibetan wild barley accessions was evaluated in terms of reduced dry biomass under salinity stress. In addition, Na⁺ and K⁺ concentrations of 48 representative accessions differing in salinity tolerance were determined. Furthermore, the allelic and functional diversity of HvHKT1 and HvHKT2 was determined by association analysis as well as gene expression assay. There was a wide variation among wild barley genotypes in salt tolerance, with some accessions being higher in tolerance than cultivated barley CM 72, and salinity tolerance was significantly associated with K⁺/Na⁺ ratio. Association analysis revealed that HvHKT1 and HvHKT2 mainly control Na⁺ and K⁺ transporting under salinity stress, respectively, which was validated by further analysis of gene expression. The present results indicated that Tibetan wild barley offers elite alleles of HvHKT1 and HvHKT2 conferring salinity tolerance.     

Levels of short-chain fatty acids and of abscisic acid in water-stressed and non-stressed leaves and their effects on stomata in epidermal strips and excised leaves

Straight-chain saturated fatty acids (C6-C11) and abscisic acid (ABA) accumulate in the leaves of Phaseolus vulgaris L. and Hordeum vulgare L. under water stress. ABA and certain of the fatty acids, particularly decanoic and undecanoic acid, can inhibit stomatal opening and cause stomatal closure in epidermal strips of Commelina communis L. depending on the incubating medium used. 10^-4 M (±)-ABA inhibits opening in media containing either high or relatively low concentrations of KCl but causes closure only in the latter medium. The fatty acids (at 10^-4 M) prevent opening in both media while significant closure of open stomata was caused only by undecanoic acid in both media and, additionally, by decanoic acid in the low-KCl medium. 10^-4 M formic acid also caused stomatal closure and prevented opening to significant extents in the low-KCl medium (it was not tested in the high-KCl medium). The efficacy of undecanoic acid in causing 50% inhibition of opening is about three orders of magnitude lower than that of ABA. At a concentration of 10^-3 M, nonanoic, decanoic and particularly undecanoic acid and all-trans-farnesol cause increased cell leakage in Beta vulgaris L. root tissue. Undecanoic acid (10^-4 M) also causes some loss of guard cell integrity in C. communis within 1.5 h of treatment. ABA (10^-4 M) reduces transpiration rates in barley and C. communis leaves when applied via the transpiration stream but decanoic and undecanoic acids did not have this effect. Transpiration was not affected when ABA or the fatty acids were applied to the leaf surfaces.Abbreviations ABA abscisic acid - RWC relative water content - SCFA short-chain fatty acids Deceased May 1977     

Physiological characterizations of three barley genotypes in response to low potassium stress

Plants adopt several strategies for fighting against low potassium (LK) stress. Our previous study identified some Tibetan wild barley accessions which show the higher LK tolerance than cultivated barley. However, the physiological mechanisms underlying the wild barley are not well understood. In this study, growth performance, elements content, SPAD value, photosynthetic parameters, and ATPase activities were measured to investigate the effect of LK stress on the two wild barley genotypes (XZ153 and XZ141) and one barley cultivar (ZD9) differing in LK tolerance. The results revealed that LK stress inhibited barley growth and induced reduction in dry weight, with XZ153 being least inhibited. Moreover, XZ153 had less reduction in photosynthetic rate, SPAD value, and K concentrations in the younger leaves under LK stress compared to the other two genotypes. Although the activities of H⁺/K⁺-ATPase and Ca²⁺/Mg²⁺-ATPase were increased significantly in all three genotypes in response to LK, the highest H⁺/K⁺-ATPase activity was observed in XZ153. The current results indicate that higher LK tolerance of XZ153 is partly attributed to its high capacity of transferring K from the old leaves to younger ones.     

An α-galactosidase with an essential function during leaf development

The putative α-galactosidase gene HvSF11 of barley, previously shown to be expressed during dark induced senescence, is expressed in the growing/elongating zone of primary foliage leaves of barley. The amino acid sequence deduced from the full length HvSF11 cDNA contains a hydrophobic signal sequence at the N-terminus. Phylogenetic relationship of the HvSF11 encoded barley α-galactosidase to other α-galactosidases revealed high homology with the α-galactosidase encoded by the gene At5g08370 from Arabidopsis thaliana. We have isolated two independent heterozygous At5g08370 T-DNA insertion mutants from Arabidopsis thaliana, both of which have a higher number of rosette leaves with a curly surface leaf morphology and delayed flowering time in comparison to wildtype plants. Localization of the Arabidopsis α-galactosidase protein via GUS-tag revealed that the protein is associated with the cell wall. This result was confirmed by immunological detection of the orthologous barley protein in a protein fraction derived from cell walls of barley leaves. It is concluded that the α-galactosidase proteins from barley and Arabidopsis might fulfill an important role in leaf development by functioning in cell wall loosening and cell wall expansion.

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Association genetics,geography and ecophysiology link stomatal patterning in Populus trichocarpa with carbon gain and disease resistance trade‐offs

Stomata are essential for diffusive entry of gases to support photosynthesis, but may also expose internal leaf tissues to pathogens. To uncover trade‐offs in range‐wide adaptation relating to stomata, we investigated the underlying genetics of stomatal traits and linked variability in these traits with geoclimate, ecophysiology, condensed foliar tannins and pathogen susceptibility in black cottonwood (Populus trichocarpa). Upper (adaxial) and lower (abaxial) leaf stomatal traits were measured from 454 accessions collected throughout much of the species range. We calculated broad‐sense heritability (H²) of stomatal traits and, using SNP data from a 34K Populus SNP array, performed a genome‐wide association studies (GWAS) to uncover genes underlying stomatal trait variation. H² values for stomatal traits were moderate (average H² = 0.33). GWAS identified genes associated primarily with adaxial stomata, including polarity genes (PHABULOSA), stomatal development genes (BRASSINOSTEROID‐INSENSITIVE 2) and disease/wound‐response genes (GLUTAMATE‐CYSTEINE LIGASE). Stomatal traits correlated with latitude, gas exchange, condensed tannins and leaf rust (Melampsora) infection. Latitudinal trends of greater adaxial stomata numbers and guard cell pore size corresponded with higher stomatal conductance (g_s) and photosynthesis (A_max), faster shoot elongation, lower foliar tannins and greater Melampsora susceptibility. This suggests an evolutionary trade‐off related to differing selection pressures across the species range. In northern environments, more adaxial stomata and larger pore sizes reflect selection for rapid carbon gain and growth. By contrast, southern genotypes have fewer adaxial stomata, smaller pore sizes and higher levels of condensed tannins, possibly linked to greater pressure from natural leaf pathogens, which are less significant in northern ecosystems.     

Slow photosynthetic induction and low photosynthesis in Paphiopedilum armeniacum are related to its lack of guard cell chloroplast and peculiar stomatal anatomy

Paphiopedilum and Cypripedium are close relatives in the subfamily Cypripedioideae. Cypripedium leaves contain guard cell chloroplasts, whereas Paphiopedilum do not. It is unclear whether the lack of guard cell chloroplasts affects photosynthetic induction, which is important for understory plants to utilize sunflecks. To understand the role of guard cell chloroplasts in photosynthetic induction of Paphiopedilum and Cypripedium, the stomatal anatomy and photosynthetic induction of Paphiopedilum armeniacum and Cypripedium flavum were investigated at different ratios of red to blue light. The highest stomatal opening and photosynthesis of intact leaves in P. armeniacum were induced by irradiance enriched with blue light. Its stomatal opening could be induced by red light 250 µmol m^?2 s^?1, but the magnitude of stomatal opening was lower than those at the other light qualities. However, the stomatal opening and photosynthesis of C. flavum were highly induced by mixed blue and red light rather than pure blue or red light. The two orchid species did not differ in stomatal density, but P. armeniacum had smaller stomatal size than C. flavum. The stomata of P. armeniacum were slightly sunken into the leaf epidermis, while C. flavum protruded above the leaf surface. The slower photosynthetic induction and lower photosynthetic rate of P. armeniacum than C. flavum were linked to the lack of guard cell chloroplasts and specific stomatal structure, which reflected an adaptation of Paphiopedilum to periodic water deficiency in limestone habitats. These results provide evidence for the morphological and physiological evolution of stomata relation for water conservation under natural selection.     

Genotypic variation in response of barley to boron deficiency

Responses of a range of barley (Hordeum vulgare L.) genotypes to boron (B) deficiency were studied in two experiments carried out in sand culture and in the field at Chiang Mai, Thailand. In experiment 1, two barley genotypes, Stirling (two-row) and BRB 2 (six-row) and one wheat (Triticum aestivum L.) genotype, SW 41, were evaluated in sand culture with three levels of applied B (0, 0.1 and 1.0 μM B) to the nutrient solution. It was found that B deficiency depressed flag leaf B concentration at booting, grain number and grain yield of all genotypes. In barley Stirling, B deficiency also depressed number of spikes plant^-1, spikelets spike^-1 and straw yield. However, no significant difference between genotypes in flag leaf B concentration was found under low B treatments. Flag leaf B concentration below 4 mg kg^-1 was associated with grain set reduction and could, therefore, be used as a general indicator for B status in barley. In experiment 2, nine barley and two wheat genotypes were evaluated in the field on a low B soil with three levels of B. Boron levels were varied by applying either 2 t of lime ha^-1 (BL), no B (B0) or 10 kg Borax ha^-1 (B+) to the soil prior to sowing. Genotypes differed in their B response for grain spike^-1, grain spikelet^-1 and grain set index (GSI). The GSI of the B efficient wheat, Fang 60, exceeded 90% in all B treatments. The B inefficient wheat SW 41 and most of the barley genotypes set grain normally (GSI >80%) only at the B+. In B0 GSI of the barley genotypes ranged from 23% to 84%, and in BL from 19% to 65%. Three of the barley with severely depressed GSI in B0 and BL also had a decreased number of spikelets spike^-1. In experiment 3, 21 advanced barley lines from the Barley Thailand Yield Nursery 1997/98 (BTYN 1997/98) were screened for B response in sand culture with no added B. Grain Set Index of the Fang 60 and SW 41 checks were 98 and 65%, respectively, and GSI of barley lines ranged between 5 and 90%. One advanced line was identified as B efficient and two as moderately B efficient. The remaining lines ranked between moderately inefficient to inefficient. These experiments have established that there is a range of responses to B in barley genotypes. This variation in the B response was observed in vegetative as well as reproductive growth. Boron efficiency should be considered in breeding and selection of barley in low B soils. This revised version was published online in June 2006 with corrections to the Cover Date.     

Direct and acclimatory responses of stomatal conductance to elevated carbon dioxide in four herbaceous crop species in the field

In order to separate the net effect of growth at elevated [CO₂] on stomatal conductance (g_s) into direct and acclimatory responses, mid‐day values of g_s were measured for plants grown in field plots in open‐topped chambers at the current ambient [CO₂], which averaged 350 μmol mol^?1 in the daytime, and at ambient + 350 μmol mol^?1[CO₂] for winter wheat, winter barley, potato and sorghum. The acclimatory response was determined by comparing g_s measured at 700 μmol mol^?1[CO₂] for plants grown at the two [CO₂]. The direct effect of increasing [CO₂] from 350 to 700 μmol mol^?1 was determined for plants grown at the lower concentration. Photosynthetic rates were measured concurrently with g_s. For all species, growth at the higher [CO₂] significantly reduced g_s measured at 700 μmol mol^?1[CO₂]. The reduction in g_s caused by growth at the higher [CO₂] was larger for all species on days with low leaf to air water vapour pressure difference for a given temperature, which coincided with highest conductances and also the smallest direct effects of increased [CO₂] on conductance. For barley, there was no other evidence for stomatal acclimation, despite consistent down‐regulation of photosynthetic rate in plants grown at the higher [CO₂]. In wheat and potato, in addition to the vapour pressure difference interaction, the magnitude of stomatal acclimation varied directly in proportion to the magnitude of down‐regulation of photosynthetic rate through the season. In sorghum, g_s consistently exhibited acclimation, but there was no down‐regulation of photosynthetic rate. In none of the species except barley was the direct effect the larger component of the net reduction in g_s when averaged over measurement dates. The net effect of growth at elevated [CO₂] on mid‐day g_s resulted from unique combinations of direct and acclimatory responses in the various species.     

Leaf extension in the slender barley mutant: delineation of the zone of cell expansion and changes in translatable mRNA during leaf development

The slender mutant of barley (Hordeum vulgare L.) results from an alteration to a single nuclear gene. Plants homozygous for the mutant allele have long, attenuated leaves as a result of a greatly increased extension rate. Although the growth rate at any one position in the extension zone appears not to differ between slender and normal (wild-type) barley, in slender the length of the zone over which cells extend is approximately 50% greater than that in normal barley. Epidermal cells are both longer and narrower in slender, so the whole-plant phenotype is mirrored at the cell level. Translation in vitro of RNA extracted from successive sections of the young primary leaf, followed by one-dimensional SDS-PAGE separation, facilitated the alignment of equivalent developmental stages in the two genotypes, but failed to demonstrate major differences between the two genotypes. Two-dimensional separation of translation products from total leaf tissue revealed a few small differences between normal and slender. Growth of plants at 8°C compared with 20°C caused changes in some translation products, with one (unknown) product decreasing in abundance in cold-treated normal tissue but not in slender tissue.     

Salicylic Acid Application Mitigates Oxidative Damage and Improves the Growth Performance of Barley under Drought Stress

Drought is a severe environmental constraint, causing a significant reduction in crop productivity across the world. Salicylic acid (SA) is an important plant growth regulator that helps plants cope with the adverse effects induced by various abiotic stresses. The current study investigated the potential effects of SA on drought tolerance efficacy in two barley (Hordeum vulgare) genotypes, namely BARI barley 5 and BARI barley 7. Ten-day-old barley seedlings were exposed to drought stress by maintaining 7.5% soil moisture content in the absence or presence of 0.5, 1.0 and 1.5 mM SA. Drought exposure led to severe damage to both genotypes, as indicated by phenotypic aberrations and reduction of dry biomass. On the other hand, the application of SA to drought-stressed plants protected both barley genotypes from the adverse effects of drought, which was reflected in the improvement of phenotypes and biomass production. SA supplementation improved relative water content and proline levels in drought-stressed barley genotypes, indicating the osmotic adjustment functions of SA under water-deficit conditions. Drought stress induced the accumulation of reactive oxygen species (ROS), such as hydrogen peroxide (H₂O₂) and superoxide (O₂ ^•− ), and the lipid peroxidation product malondialdehyde (MDA) in the leaves of barley plants. Exogenous supply of SA reduced oxidative damage by restricting the accumulation of ROS through the stimulation of the activities of key antioxidant enzymes, including superoxide dismutase (SOD), peroxidase (POD), catalase (CAT), ascorbate peroxidase (APX) and glutathione peroxidase (GPX). Among the three-applied concentrations of SA, 0.5 mM SA exhibited better mitigating effects against drought stress considering the phenotypic performance and biochemical data. Furthermore, BARI barley 5 showed better performance under drought stress than BARI barley 7 in the presence of SA application. Collectively, our results suggest that SA played a crucial role in improving water status and antioxidant defense strategy to protect barley plants from the deleterious effects of water deficiency.