Silicon-Mediated Regulation of Sodium Distribution in Barley Plants

Salinity is one of the largest problems in the world today. Silicon (Si)-mediated increase in plant tolerance to saline environment has been well documented, while the underlying mechanisms remain unclear. Monosilicic acid, polysilicic acid, and sodium (Na) were analyzed in the apoplast and symplast of roots, stems and leaves of salt-stressed barley plants in dynamics. Sodium moved predominantly via apoplastic pathway. The dynamics of Na in apoplast represented a parabolic curve. Soluble Si in nutrient solution increased the total Na in the roots but restricted the Na root-to-shoot transport via apoplastic pathway and reduced Na accumulation in stems and leaves. Plant exposure to high concentration of Na resulted in increased polysilicic acids in the root symplast and stem apoplast and symplast. These increases are attributable to Si redistribution within plant with its accumulation in stressed tissue. Probably, Si moves in the form of polysilicic acid. Under optimum or low stress growth conditions, Si mainly accumulated in the roots and leaves. Under higher stress, this Si can be redistributed to a mostly stress-affected place.

Analysis of Phenotypic Characteristics and Sucrose Metabolism in the Roots of Raphanus sativus L.

The taproot of radish (Raphanus sativus L.) is an important sink organ; it is morphologically diverse and contains large amounts of secondary metabolites. Sucrose metabolism is believed to be important in the development of sink organs. We measured the amounts of glucose, fructose, and sucrose in the roots of sixty three radish accessions and analyzed the association between the sugar content and the root phenotype. Fructose content correlated with the root color and length characteristics, glucose was the most abundant sugar in the roots, and the sucrose content was very low, compared to that of the hexoses in most of the accessions. Expression analysis of the genes involved in sucrose metabolism, transportation, starch synthesis, and cell wall synthesis was performed through RNA sequencing. The genes encoding sucrose synthases (SUSY) and the enzymes involved in the synthesis of cellulose were highly expressed, indicating that SUSY is involved in cell wall synthesis in radish roots. The positive correlation coefficient (R) between the sucrose content and the expression of cell wall invertase and sugar transporter proteins suggest that hexose accumulation could occur through the apoplastic pathway in radish roots. A positive R score was also obtained when comparing the expression of genes encoding SUSY and fructokinase (FK), suggesting that the fructose produced by SUSY is mostly phosphorylated by FK. In addition, we concluded that sucrose was the most metabolized sugar in radish roots.

Overexpression of OsCASP1 Improves Calcium Tolerance in Rice

The Casparian strip domain protein 1 (OsCASP1) is necessary for the formation of the Casparian strip (CS) in the rice endodermis. It also controls Ca2+ transport to the stele. Here, we demonstrated that OsCASP1 overexpression enhanced Ca tolerance in rice. Under normal conditions, OsCASP1-overexpressed lines showed similar concentrations of essential metals in the roots and shoots compared to the wild type, while under high Ca conditions, Ca in the roots, shoots, and xylem sap of the OsCASP1-overexpressed lines was significantly decreased. This did not apply to other essential metals. Ca-inhibited growth was significantly alleviated in the OsCASP1-overexpressed lines. Furthermore, OsCASP1 overexpression resulted in earlier formation of both the CS and functional apoplastic barrier in the endodermis but did not induce ectopic CS formation in non-endodermal cell layers and affect suberin accumulation in the endodermis. These results indicate that the overexpression of OsCASP1 promotes CS formation in endodermal cells and inhibits Ca2+ transport by the apoplastic pathway, restricting Ca accumulation in the roots and shoots under high Ca conditions. Taken together, the results suggest that OsCASP1 overexpression is an effective way to improve rice adaptation to high Ca environments.

Phloem unloading via the apoplastic pathway is essential for shoot distribution of root-synthesized cytokinins

Root-synthesized cytokinins are transported to the shoot and regulate the growth, development, and stress responses of aerial tissues. Previous studies have demonstrated that Arabidopsis (Arabidopsis thaliana) ATP binding cassette (ABC) transporter G family member 14 (AtABCG14) participates in xylem loading of root-synthesized cytokinins. However, the mechanism by which these root-derived cytokinins are distributed in the shoot remains unclear. Here, we revealed that AtABCG14-mediated phloem unloading through the apoplastic pathway is required for the appropriate shoot distribution of root-synthesized cytokinins in Arabidopsis. Wild-type rootstocks grafted to atabcg14 scions successfully restored trans-zeatin xylem loading. However, only low levels of root-synthesized cytokinins and induced shoot signaling were rescued. Reciprocal grafting and tissue-specific genetic complementation demonstrated that AtABCG14 disruption in the shoot considerably increased the retention of root-synthesized cytokinins in the phloem and substantially impaired their distribution in the leaf apoplast. The translocation of root-synthesized cytokinins from the xylem to the phloem and the subsequent unloading from the phloem are required for the shoot distribution and long-distance shootward transport of root-synthesized cytokinins. This study revealed a mechanism by which the phloem regulates systemic signaling of xylem-mediated transport of root-synthesized cytokinins from the root to the shoot.

Phloem loading in rice leaves depends strongly on the apoplastic pathway

Phloem loading is the first step in sucrose transport from source leaves to sink organs. The phloem loading strategy in rice remains unclear. To determine the potential phloem loading mechanism in rice, yeast invertase (INV) was overexpressed specifically in the cell wall by 35S promoter to block sugar transmembrane loading in rice. The transgenic lines exhibited obvious phloem loading suppression characteristics accompanied by the accumulation of sucrose and starch, restricted vegetative growth and decreased grain yields. The decreased sucrose exudation rate with p-chloromercuribenzenesulfonic acid (PCMBS) treatment also indicated that rice actively transported sucrose into phloem. Moreover, the expression level of OsSUT1 was much higher than that of other plasma membrane localized OsSUTs in the source leaf. Cross sections of the GUS transgenic plant showed that the signals of OsSUT1 and OsSUT5 occurred in the phloem companion cells. The ossut1 and ossut4 mutants presented a decrease of grain yield, implying important roles of OsSUTs in phloem loading. Based on these results, we conclude that rice uses the apoplastic loading as a major phloem loading strategy.

CHANGE IN CONTRIBUTION OF AQUAPORINS TO THE HYDRAULIC CONDUCTIVITY OF PLANTS DURING THE FORMATION OF APOPLASTIC BARRIERS UNDER SALINITY

The study of plant adaptation mechanisms during the salt stress is required to provide an increase in plant productivity under such conditions. Along with a decrease in the availability of water for plants, the NaCL-induced inhibition of plant growth is associated with the toxic effect of sodium ions. The formation of apoplastic barriers due to the deposition of suberin and lignin restricts passive ion diffusion. However, the formation of such barriers reduces the capacity of the apoplastic pathway for water movement. In these conditions the role of transmembrane water transport is increased. This process is provided by aquaporin water channels. Thus the purpose of this work was to determine the contribution of aquaporins to hydraulic conductivity of peas plants under salinity-induced apoplastic barrier formation. An only slight decrease in plants transpiration caused by mercury chloride in the absence of salinization was in accordance with the ideas the apoplast is the dominant pathway when the Casparian bands is not formed yet. Salt stress in our experiments accelerated the development of the Casparian bands formation which could be visualized as an appearance of suberin strips in root endodermis which in turn was accompanied by a decrease in hydraulic conductivity. The decrease in hydraulic conductivity in 2 times during the mercury chloride treatment under salinity confirmed that contribution of aquaporins to the total hydraulic conductivity was increased under conditions when Casparian bands have had formed.

Effects of major vein blockage and aquaporin inhibition on leaf hydraulics and stomatal conductance

The density and architecture of leaf veins determine the network and efficiency of water transport within laminae and resultant leaf gas exchange and vary widely among plant species. Leaf hydraulic conductance ( K leaf ) can be regulated by vein architecture in conjunction with the water channel protein aquaporin. However, our understanding of how leaf veins and aquaporins affect leaf hydraulics and stomatal conductance ( g s ) remains poor. By inducing blockage of the major veins and inhibition of aquaporin activity using HgCl 2 , we examined the effects of major veins and aquaporins on K leaf and g s in species with different venation types. A vine species, with thick first-order veins and low vein density, displayed a rapidly declined g s with high leaf water potential in response to vein blockage and a greatly reduced K leaf and g s in response to aquaporin inhibition, suggesting that leaf aquaporins are involved in isohydric/anisohydric stomatal behaviour. Across species, the decline in K leaf and g s due to aquaporin inhibition increased linearly with decreasing major vein density, possibly indicating that a trade-off function between vein architecture (apoplastic pathway) and aquaporin activity (cell-to-cell pathway) affects leaf hydraulics.