Haijun Gong

Beneficial effects of silicon on salt and drought tolerance in plants

Soil salinity and drought are major abiotic factors that limit crop growth and productivity worldwide. Indeed, soil salinity and drought disrupt the cellular ionic and osmotic balance. Although silicon (Si) is generally considered nonessential for plant growth and developments, Si uptake by plants can alleviate both biotic and abiotic stresses. Silicon application could therefore improve crop production under adverse climate and soil conditions. Several reports have reviewed the benefits of silicon application on crop growth, but the mechanisms of silicon action have not been systematically discussed. Here, we review recent advances on silicon uptake, transport, and accumulation in plants and how silicon alleviates salinity toxicity and drought stress. The major points are the following: (1) both passive and active silicon uptake may coexist in plants; (2) although silicon transporters have been identified in some plants, more silicon transporters remain to be identified, and the process of silicon transport needs further clarification; (3) the mechanisms for silicon-mediated tolerance of salinity and drought have been extensively investigated at both physiological and biochemical levels. The physiological aspects include increasing water uptake by roots, maintaining nutrient balance, decreasing water loss from leaves, and promoting photosynthetic rate. At the biochemical level, silicon may improve antioxidant defense abilities by increasing the activities of antioxidant enzymes and the contents of non enzymatic antioxidants; silicon may also contribute to osmotic adjustment and increase photosynthetic enzymatic activities; and (4) silicon can regulate the levels of endogenous plant hormones under stress conditions, whereas silicon involvement in signaling and regulation of gene expression related to increasing stress tolerance remains to be explored.

Effects of silicon on defense of wheat against oxidative stress under drought at different developmental stages

The effects of silicon application before sowing on the drought-induced oxidative stress and antioxidant defense in wheat (Triticum aestivum L.) were investigated. Drought stress was applied by withholding watering till sampling at booting or filling stage. Application of Si increased the water potential of drought-stressed plants at filling stage, whereas it did not at booting stage. The superoxide dismutase (SOD) activity was inhibited and peroxidase (POD) activity was enhanced by drought at booting stage, and no differences were observed due to the Si treatment. At filling stage, however, application of Si increased the SOD activity and decreased the POD activity of drought-stressed plants. The catalase (CAT) activity was slightly increased by drought only in the absence of Si and at booting stage. The activity of glutathione reductase (GR) was not greatly influenced. Application of Si did not change the contents of H2O2, total soluble protein and protein carbonyl of drought-stressed plants at booting stage, whereas at filling stage, it decreased the content of H2O2 and protein carbonyl and increased the content of total soluble protein. The content of thiobarbituric acid reactive substances (TBARS) and the activities of acid phospholipase (AP) and lipoxygenase (LOX) in drought-stressed plants were also decreased by application of Si at both stages.

Silicon decreases chloride transport in rice (Oryza sativa L.) in saline conditions

Silicon can alleviate salt damage to plants, although the mechanism(s) still remains to be elucidated. In this paper, we report the effect of silicon on chloride transport in rice (Oryza sativa L.) seedlings in saline conditions. In the absence of salinity, silicon enhanced the growth of shoots, but not roots in three cultivars (cv. GR4, IR36, and CSR10). Salinity reduced the growth of both shoots and roots in all three genotypes. In saline conditions, addition of silicon to the culture solution again improved the growth of shoots, but not of roots. Under these saline conditions, the concentrations of chloride in the shoot were markedly decreased by adding silicon and the ratio of K(+)/Cl(-) was significantly increased, while the concentration of chloride in the roots was unchanged. The decrease in chloride concentration in the shoot was correlated with the decrease in transpirational bypass flow in rice, as shown by the transport of the apoplastic tracer trisodium-8-hydroxy-1,3,6-pyrenetrisulphonic acid (PTS). Addition of silicon increased the net photosynthetic rate, stomata conductance, and transpiration of salt-stressed plants in cv. IR36, indicating that the reduction of chloride (and sodium) uptake by silicon was not through a reduction in transpiration rate. Silicon addition also increased the instantaneous water use efficiency of salt-stressed plants, while it did not change the relative growth rate of shoots. The results suggest that silicon addition decreased transpirational bypass flow in the roots, and therefore decreased the transport of chloride to the shoot.

The regulatory role of silicon on water relations, photosynthetic gas exchange, and carboxylation activities of wheat leaves in field drought conditions

The effects of silicon on water relations, photosynthetic gas exchange, and carboxylation activities of wheat (Triticum aestivum L.) leaves were investigated in field drought conditions. Silicon application improved the leaf relative water content and water potential under drought. The leaf net photosynthetic rate and stomatal conductance were significantly decreased between 7:30 and 17:30 under drought, whereas silicon application increased the leaf net photosynthetic rate between 7:30 and 15:30 with an exception at 9:30. Silicon application also increased the leaf stomatal conductance at 13:30 and 17:30 under drought. The leaf transpiration rate was decreased by drought but it was increased by silicon from 13:30 to 17:30. The intercellular CO2 concentration was increased at 7:30 under drought, while it was decreased most of the time from midday to the afternoon. The leaf stomatal limitation was increased under drought from 11:30 to 17:30, whereas it was intermediate in silicon treated plants. The instantaneous water use efficiency was significantly increased by silicon application at 7:30 under drought. Silicon application slightly decreased the activity of ribulose-1, 5-bisphosphate carboxylase, but it increased the activity of phosphoenolpyruvate carboxylase and the concentration of inorganic phosphorus under drought. These results suggest that silicon could improve the photosynthetic ability of wheat in field drought conditions, and both stomatal and non-stomatal factors were involved in the regulation. In the early morning (at 7:30), the non-stomatal factor was the main contributor; 9:30 was a turning point, after which the stomatal factor was the main contributor.

Contrast in chloride exclusion between two grapevine genotypes and its variation in their hybrid progeny

Potted grapevines of 140 Ruggeri (Vitis berlandieri × Vitis rupestris), a good Cl− excluder, and K 51-40 (Vitis champinii × Vitis riparia ‘Gloire’), a poor Cl− excluder, and of a family obtained by crossing the two genotypes, were used to examine the inheritance of Cl− exclusion. Rooted leaves were then used to further investigate the mechanism for Cl− exclusion in 140 Ruggeri. In both a potting mix trial (plants watered with 50 mM Cl−) and a solution culture trial (plants grown in 25 mM Cl−), the variation in Cl− accumulation was continuous, indicating multiple rather than single gene control for Cl− exclusion between hybrids within the family. Upper limits of 42% and 35% of the phenotypic variation in Cl− concentration could be attributed to heritable sources in the potting mix and solution culture trials, respectively. Chloride transport in roots of rooted leaves of both genotypes appeared to be via the symplastic pathway, since addition of 8-hydroxy-1,3,6-pyrenetrisulphonic acid (PTS), an apoplastic tracer, revealed no obvious PTS fluorescence in the laminae of either genotype, despite significant accumulation of Cl− in laminae of K 51-40 during the PTS uptake period. There was no significant difference in either unidirectional 36Cl− flux (10 min) or 36Cl− uptake (3 h) into roots of rooted leaves exposed to 5, 10, or 25 mM Cl−. However, the percentage of 36Cl− transported to the lamina (3 h) was significantly lower in 140 Ruggeri than in K 51-40, supporting reduced Cl− loading into xylem and implicating the root stele in the Cl− exclusion mechanism.

Silicon Enhances Water Stress Tolerance by Improving Root Hydraulic Conductance in Solanum lycopersicum L.

Silicon (Si) can improve drought tolerance in plants, but the mechanism is still not fully understood. Previous research has been concentrating on Si’s role in leaf water maintenance in Si accumulators, while little information is available on its role in water uptake and in less Si-accumulating plants. Here, we investigated the effects of Si on root water uptake and its role in decreasing oxidative damage in relation to root hydraulic conductance in tomato (Solanum lycopersicum ‘Zhongza No.9’) under water stress. Tomato seedlings were subjected to water stress induced by 10% (w/v) polyethylene glycol-6000 in the absence or presence of 2.5 mM added silicate. The results showed that Si addition ameliorated the inhibition in tomato growth and photosynthesis, and improved water status under water stress. The root hydraulic conductance of tomato plants was decreased under water stress, and it was significantly increased by added Si. There was no significant contribution of osmotic adjustment in Si-enhanced root water uptake under water stress. The transcriptions of plasma membrane aquaporin genes were not obviously changed by Si under water stress. Water stress increased the production of reactive oxygen species and induced oxidative damage, while added Si reversed these. In addition, Si addition increased the activities of superoxide dismutase and catalase and the levels of ascorbic acid and glutathione in the roots under stress. It is concluded that Si enhances the water stress tolerance via enhancing root hydraulic conductance and water uptake in tomato plants. Si-mediated decrease in membrane oxidative damage may have contributed to the enhanced root hydraulic conductance.

Induction of tetraploidy in Juncus effusus by colchicine

Tetraploidy was induced in vitro in mat rush (Juncus effusus L.) cultivar Nonglin-4 by exposure to colchicine (0, 50, 100 and 500 mg dm−3) for 6, 12 and 24 h. Flow cytometric analysis was used to confirm the ploidy level. Anatomical and ultrastructural analyses at cellular and subcellular levels in tetraploid and diploid control plants revealed differences between diploid and tetraploid plants. The leaf epidermis had larger stomata but lower stomatal density in tetraploid plants. In addition, mesophyll cells in tetraploid plants appeared more compact and showed less intercellular spaces along with increased size of vascular bundles. However, a significant reduction of chlorophyll content was observed in tetraploid plants that might be the result of structural modification in the lamellar membranes of chloroplasts.

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.

Isolation and functional characterization of CsLsi1, a silicon transporter gene in Cucumis sativus

Cucumber (Cucumis sativus) is a widely grown cucurbitaceous vegetable that exhibits a relatively high capacity for silicon (Si) accumulation, but the molecular mechanism for silicon uptake remains to be clarified. Here we isolated and characterized CsLsi1, a gene encoding a silicon transporter in cucumber (cv. Mch-4). CsLsi1 shares 55.70 and 90.63% homology with the Lsi1s of a monocot and dicot, rice (Oryza sativa) and pumpkin (Cucurbita moschata), respectively. CsLsi1 was predominantly expressed in the roots, and application of exogenous silicon suppressed its expression. Transient expression in cucumber protoplasts showed that CsLsi1 was localized in the plasma membrane. Heterologous expression in Xenopus laevis oocytes showed that CsLsi1 evidenced influx transport activity for silicon but not urea or glycerol. Expression of cucumber CsLsi1-mGFP under its own promoter showed that CsLsi1 was localized at the distal side of the endodermis and the cortical cells in the root tips as well as in the root hairs near the root tips. Heterologous expression of CsLsi1 in a rice mutant defective in silicon uptake and the over-expression of this gene in cucumber further confirmed the role of CsLsi1 in silicon uptake. Our results suggest that CsLsi1 is a silicon influx transporter in cucumber. The cellular localization of CsLsi1 in cucumber roots is different from that in other plants, implying the possible effect of transporter localization on silicon uptake capability.

Analysis of the salt exclusion phenotype in rooted leaves of grapevine (Vitis spp.): Salt exclusion in rooted leaves of grapevine

Background and Aims Rooted leaves were used to analyse the salt exclusion phenotype in grapevines. Genotypes included rootstocks 140 Ruggeri and K51‐40 and cultivar Cabernet Sauvignon – respectively, good, poor and intermediate chloride excluders. Methods and Results We investigated the effect of short‐term salt treatment on the chloride, sodium and potassium concentration of organs and whole rooted leaves and the time course of chloride accumulation in salt‐treated, whole rooted leaves. The effect of Control and salt plus low and high nitrate concentration on chloride and nitrate concentration in organs was assessed. Salt treatment increased chloride and sodium concentration in all organs but had no effect on potassium concentration. The chloride concentration of whole rooted leaves and the sodium concentration of lamina and petiole were similar between salt‐treated rooted leaves of 140 Ruggeri and K51‐40; 140 Ruggeri accumulated more chloride in roots and less in leaf (petiole and lamina) than K51‐40. Roots of salt‐treated 140 Ruggeri and K51‐40 responded to higher external nitrate by decreasing chloride concentration and increasing nitrate concentration. Conclusions Restricted transport of chloride to the leaf and greater storage in roots is a feature of the chloride exclusion phenotype in rooted leaves. The short‐term capacity of 140 Ruggeri for chloride exclusion from the leaf was reduced in the salt plus high nitrate treatment. This was linked to higher nitrate and reduced chloride accumulation in roots. Significance of the Study The chloride exclusion phenotype in rooted leaves was shown to involve greater partitioning to roots and less to the leaf, rather than differences in net chloride accumulation by whole rooted leaves.