Cheng-Lie Zhang

Effects of Silicon on Growth of Wheat Under Drought

Abstract Plants of wheat growing in pots with silicon (Si) applied before sowing had greater plant height, leaf area, and dry materials compared to those without Si applied in well watering conditions. Drought stress was applied by withholding watering for 12 days from 26‐day old seedlings. In the stress conditions, plants growing in Si‐applied soil could maintain higher relative water content (RWC), water potential and leaf area compared to those without Si applied. Moreover, the Si applied plant dry materials were not significantly changed by drought while those of plants growing in pots without Si applied were significantly decreased, and this was mainly due to growth inhibition of the shoots. Drought stressed wheat growing in pots with Si applied had a significantly greater leaf weight ratio (LWR) and lower specific leaf area (SLA) compared to those of stressed plants in the absence of applied Si. This demonstrates that the leaves of stressed plants growing in pots with Si applied were thicker compared to those without Si applied. This may have a beneficial effect by reducing the transpirational loss of water and maintain high RWC and water potential. Therefore, application of Si may be one of the available pathways to improve growth of this crop and increase its production in arid or semi arid areas.

Evidence for the involvement of nitric oxide and reactive oxygen species in osmotic stress tolerance of wheat seedlings: Inverse correlation between leaf abscisic acid accumulation and leaf water loss

Nitric oxide (NO) and reactive oxygen species (ROS) play important roles in both abscisic acid (ABA) signaling and stress-induced ABA accumulation. However, little is known about their physiological roles in the whole plant. In this study, the effects of NO and ROS on leaf water control and the roles of ABA were determined using wheat (Triticum aestivum L.) seedlings. As compared with the control, osmotic stress reduced leaf water loss (LWL) while it increased leaf ABA content. The effects of osmotic stress on LWL and ABA contents were partially reversed by NO scavengers or NO synthase (NOS) inhibitors. Furthermore, sodium nitroprusside (SNP) at concentrations between 0.01 and 10 mM all reduced LWL efficiently and induced ABA accumulation in a dose-dependent manner. When ABA synthesis was inhibited by fluridone or actidione, the effects of SNP on LWL were partially reversed. These results suggest that NO is involved in leaf water maintenance of wheat seedlings under osmotic stress, and one of the possible mechanisms is by stimulating ABA synthesis. The ROS scavengers used in our experiments had no effects on either LWL or ABA accumulation induced by osmotic stress. However, all ROS induced LWL reduction and ABA accumulation significantly. Hydrogen peroxide had the same effects as SNP on LWL and induced ABA accumulation in a dose-dependent manner but had a maximal effect at 1 mM. Fluridone reversed the effects of H2O2 on both LWL reduction and ABA accumulation, while actidione had no effect. These results suggest that ROS are also involved in leaf water maintenance of wheat seedlings by stimulating ABA biosynthesis, but with a different mechanism to that of NO. The ABA-independent mechanism in NO/ROS regulation of leaf water balance is discussed, in relation to our results.

Gradual Drought Under Field Conditions Influences the Glutathione Metabolism, Redox Balance and Energy Supply in Spring Wheat

Glutathione (GSH) metabolism, redox balance and energy supply in spring wheat (Triticum aestivum L.) during gradual drought stress under field conditions were investigated. Although levels of total and reduced GSH were decreased, the ratio of GSH/GSSG (glutathione disulfide) was markedly increased by drought. Levels of GSH biosynthetic precursors, cysteine (Cys) and γ-glutamylcysteine (γ-GC), and the activities of their biosynthetic enzymes, γ-glutamylcysteine synthetase (γ-GCS) and glutathione synthetase (GSHS) were also significantly increased in stressed plants. Glutathione reductase (GR) activity, which is responsible for the conversion of GSSG to GSH, was also increased under this field stress. However, two other important enzymes in GSH metabolism, glutathione peroxidase (GP) and glutathione S-transferase (GST), showed decreased activity in the droughted plants. These results suggest that the higher ratio of GSH/GSSG, the rate of GSH biosynthesis and the capacity of its redox cycling rather than GSH accumulation might be essential for drought resistance of plants. Activities of the two key Calvin-cycle enzymes possessing exposed sulfhydryl groups, NADP+-dependent glyceraldehydes-3-phosphate dehydrogenase (G3PD) and fructose-1,6-bisphosphatase (FBPase) were not affected by drought stress, whereas, activity of the key enzyme in the pentose-phosphate pathway (PPP), 6-phosphogluconate dehydrogenase (6-PGD), increased in the droughted plants. The ratios of NADPH/NADP+, NADH/NAD+ and ATP/ADP increased in the droughted plants, indicating that an up-regulation of the reduced redox state and the energy supply in the plant cells might be an important physiological strategy for plants responding to drought stress. A simple correlation between the high ratio of GSH/GSSG, the rate of GSH biosynthesis and the redox cycle and the high reduction states of redox status in the plant cells was also observed under field drought.

Puccinellia tenuiflora exhibits stronger selectivity for K+ over Na+ than wheat

Abstract The characteristics of selectivity for K+ over Na+ by the roots of the halophyte Puccinellia tenuiflora were investigated in comparison with the glycophyte wheat (Triticum aestivum). Under various NaCl concentrations, the concentrations of K+ in the shoots of P. tenuiflora were 16–24% lower than those of wheat, whereas the concentrations of K+ in Puccinellia roots were 2.8–4.0 times higher than those of wheat. In 200 mM NaCl, the concentrations of Na+ in shoots of P. tenuiflora and wheat were similar under high K+ levels, but the concentrations of Na+ in wheat were 1.6 times higher than those in Puccinellia under low K+ levels. The concentrations of K+ in roots of P. tenuiflora were 1.5–2.0 times higher than those of wheat under low K+ levels. Formulas are given for calculating net selective absorption (SA) capacity and selective transport (ST) capacity by roots for K+ over Na+. We interpret SA as the net capacity of selectively absorbing K+ over Na+ by epidermal and cortical cells of whole plant roots into the root symplast. ST could reflect the net capacity of selection for K+ over Na+ transport from whole root stelar symplast to the xylem vessels. The lower ST value of P. tenuiflora might be the reason for accumulation of K+ in its roots. The SA values of P. tenuiflora and wheat were approximately equivalent in the low-affinity K+ uptake range. The SA values of the former were about two times higher than that of the latter in the high-affinity K+ uptake range, showing the root high-affinity K+ uptake system of the halophyte P. tenuiflora has a stronger capacity for K+ uptake.

Photosynthetic Electron Transport, Photophosphorylation, and Antioxidants in Two Ecotypes of Reed (Phragmites Communis Trin.) from Different Habitats

We compared chloroplast photochemical properties and activities of some chloroplast-localised enzymes in two ecotypes of Phragmites communis, swamp reed (SR, C3-like) and dune reed (DR, C4-like) plants growing in the desert region of north-west China. Electron transport rates of whole electron transport chain and photosystem (PS) 2 were remarkably lower in DR chloroplasts. However, the electron transport rate for PS1 in DR chloroplasts was more than 90 % of the activity similar in the SR chloroplasts. Activities of Mg2+-ATPase and cyclic and non-cyclic photophosphorylations were higher in DR chloroplasts than in the SR ones. The activities of chloroplast superoxide dismutase (SOD) and ascorbate peroxidase (APX), both localised at or near the PS1 complex and serving to scavenge active oxygen around PS1, and the content of ascorbic acid, a special substrate of APX in chloroplast, were all higher in DR chloroplasts. Hence reed, a hydrophytic plant, when subjected to intense selection pressure in dune habitat, elevates its cyclic electron flow around PS1. In consequence, it provides extra ATP required by C4 photosynthesis. Combined high activities of active oxygen scavenging components in DR chloroplasts might improve protection of photosynthetic apparatus, especially PS1, from the damage of reactive oxygen species. This offers new explanation of photosynthetic performance of plant adaptation to long-term natural drought habitat, which is different from those, subjected to the short-term stress treatment or even to the artificial field drought.

Up-regulation of glutathione metabolism and changes in redox status involved in adaptation of reed (Phragmites communis) ecotypes to drought-prone and saline habitats

The glutathione (GSH) metabolic characteristics and redox balance in three ecotypes of reed (Phragmites communis), swamp reed (SR), dune reed (DR), and heavy salt meadow reed (HSMR), from different habitats in desert regions of northwest China were investigated. The DR possessed the highest rate of GSH biosynthesis and metabolism with the lowest levels of total and reduced GSH and its biosynthetic precursors, gamma-glutamylcysteine (gamma-EC) and cysteine (Cys), of the three reed ecotypes. This suggests that a higher rate of GSH biosynthesis and metabolism, but not GSH accumulation, might be involved in the adaptation of this terrestrial reed ecotype to its dry habitat. The HSMR shared this profile although it exhibited the highest reduced thiol levels of the three ecotypes. Two key enzymes in the Calvin-cycle possessing exposed sulfhydryl groups, NADP(+)-dependent glyceraldehydes-3-phosphate dehydrogenase (G3PD) and fructose-1,6-bisphosphatase (FBPase), and other two key enzymes in the pentose-phosphate pathway (PPP), glucose-6-phosphate dehydrogenase (G6PDH) and 6-phosphogluconate dehydrogenase (6-PGD), had very similar activities in the three reed ecotypes. Compared to the SR, the DR and HSMR had higher ratios of NADPH/NADP+ and NADH/NAD+, indicating that a more reduced redox status in the plant cells might be involved in the survival and adaptation of the two terrestrial reed ecotypes to long-term drought and salinity, respectively. These results suggest that changes of GSH metabolism and redox balance were important components of the adaptation of reed, a hydrophilic plant, to more extreme dune and saline habitats. The coordinated up-regulations of the rate of GSH biosynthesis and metabolism and reduction state of redox status of plant cells, conferred on the plant high resistance or tolerance to long-term drought and salinity.

Solute levels and osmoregulatory enzyme activities in reed plants adapted to drought and saline habitats

The pattern of solute accumulation and the activities of key enzymes involved in metabolism of proline and betaine were investigated in three ecotypes of reed from different habitats: swamp reed (SR), dune reed (DR), and heavy salt meadow reed (HSR). The two terrestrial reed ecotypes, DR and HSR, exhibited a higher capacity for osmotic adjustment; they accumulated higher contents of K+ and Ca2+ in the leaves in comparison with SR. DR also had the highest soluble sugar content in its leaves. HSR has higher levels of Na+ in its root environment and this was reflected by considerable accumulation of Na+ in the HSR rhizome. However, the different zones of its leaf lamina (upper, middle and lower) did not exhibit increased levels of Na+, suggesting that HSR has the ability to accumulate Na+ in the rhizome to protect the shoots from excessive Na+ toxicity. DR and HSR had higher levels of proline and betaine in the leaves than did SR. This difference was consistent with the activities of the various biosynthetic enzymes: betaine aldehyde dehydrogenase (BADH), pyrroline-5-carboxylate reductase (P5CR) and ornithine-δ-aminotransferase (OAT) were enhanced in DR and HSR as compared to SR, whereas proline oxidase (PO) activities were inhibited. These findings suggest that changes in the activities of enzymes involved in osmotregulation might play important roles in the adaptation of reed, a hydrophilic plant, to more extreme dune and saline habitats. The relative contributions of the various proline synthetic pathways are also discussed.

Drought stress stimulates p-nitrophenyl phosphate hydrolysis rate of the plasma membrane H+-ATPase from wheat leaves

The influence of drought stress on the ATP and p-nitrophenyl phosphate (PNPP) hydrolysis activity by plasma membrane H+-ATPase was investigated using purified plasma membrane vesicles from wheat leaves by two-phase partitioning. Drought stress increased the ATPase activity, and the optimal pH was shifted from 6.5 to about 7.0. Drought stress also stimulated the PNPP hydrolysis rate. The Km for PNPP hydrolysis was moved from 4.49 ± 0.33 mM to 3.64 ± 0.12 mM. In addition, the PNPP hydrolysis was more sensitive to vanadate under drought compared to the control. However, the inhibitory effect of hydroxylamine on the ATPase was not changed by the present drought stress. In addtion, drought stress also decreased the trypsin activation of PNPP hydrolysis by PM H+-ATPase. These results suggested that drought stress altered the catalytic mechanism of the plasma membrane H+-ATPase, and the stimulation of its activity by drought stress was mainly due to increase of the catalytic activity of its phosphatase domain. It is also suggested that drought stress might alter the structure or property of the C-terminal end of PM H+-ATPase, therefore increasing the catalytic activity of the phosphatase domain.

Composition and characteristic differences in photosynthetic membranes of two ecotypes of reed (Phragmites communis L.) from different habitats

As compared with the swamp reed (SR) ecotype of Phragmites communis growing in the desert region of northwest China, plants of the dune reed (DR) ecotype from the same region possessed lower chlorophyll (Chl) content in leaves, and less thylakoids and grana stacks in chloroplasts. Tube gel electrophoresis without stain showed that the contents of Chl-protein (Pro) components related to photosystem 2 (PS2) were markedly lower in the DR thylakoid membranes than in the SR thylakoid membranes, while the contents of Chl-Pro components associated with PS1 were almost the same in both types. SDS-PAGE analysis indicated that the content of polypeptides of the light-harvesting Chl a/b complex of PS2 (LHC2) was lower in the DR thylakoids. Besides, the conformation of LHC2 within the DR thylakoid membranes was also altered as indicated by circular dichroism spectra. Hence in the DR, reduced energy harvesting by declining the size of LHC2 might be responsible for the down-regulated PS2 activity. Chl fluorescence parameters. Fv/Fm and quantum efficiency of PS2 (ΦPS2), were lower in the DR leaves than in the SR ones. However, non-photochemical quenching coefficient (qN) was greater in DR than that in SR, implying other energy dissipation way exists in the DR photosynthetic membranes.

Antioxidant defense system in Phragmites communis Trin. ecotypes

The antioxidant defense system in three ecotypes of reed (Phragmites communis Trin.), swamp reed (SR), dune reed (DR), and heavy salt meadow reed (HSMR), from northwest China were investigated. The HSMR possessed the highest ratio of ascorbate (ASC)/dehydroascorbate (DHA) and activities of superoxide dismutase (SOD) and catalase among the three reed ecotypes, whereas, the DR exhibited the highest ratio of glutathione/glutathione disulfide and activities of ASC peroxidase (APX) and DHA reductase. Malondialdehyde and hydrogen peroxide contents were highest in HSMR, intermediate in SR, and lowest in DR. In addition, different isoenzymes of glutathion reductase, APX, SOD and DHA were also observed in three reed ecotypes.