Analogy of silicon and boron in plant nutrition.

Silicon (Si) and boron (B) are a class of elements called metalloids, which have properties like metals and non-metals. Si is classified as a quasi-essential element, while B is a micronutrient element for plants. Nowadays, numerous discoveries have shown the analogy of silicon and boron in plant nutrition. In this minireview, the molecular mechanisms for the transport of these two metalloids are compared. We also discussed the chemical forms of Si and B and their functional similarity in response to environmental stresses in plants. In conclusion, it can be proposed that cell wall-bound silicon rather than silica might partially replace boron for plant growth, development, and stress responses, and the underlying mechanism is the Si contribution to B in its structural function.

Regulation of thermodynamics and kinetics of silica nucleation during the silicification process in higher plants.

The formation mechanism of SiO2 aggregates is controversial because two contrasting hypotheses are often proposed to explain plant silicification. In this review, we summarize the physicochemical fundamentals of amorphous silica nucleation and discuss how plants regulate the process of silicification by influencing the thermodynamics and kinetics of silica nucleation. At silicification positions, plants overcome the thermodynamic barrier by establishing the supersaturation of the H4SiO4 solution and reducing the interfacial free energy. Among the thermodynamic-drivers, the establishment of supersaturation of H4SiO4 solution mainly depends on the expression of Si transporters for H4SiO4 supply, evapotranspiration for concentrating Si, and the other solutes in H4SiO4 solution for influencing the dissolution equilibrium of SiO2; while the interfacial free energy was reduced seemingly by the overexpression Na+/H+ antiporter SOS1 in high NaCl-stressed rice. Moreover, some kinetic-drivers, such as silicification-related proteins (Slp1 and PRP1) and new cell wall components, are actively expressed or synthesized by plants to interact with silicic acid, thereby reducing the kinetic barrier. According to classical nucleation theory, when the thermodynamic barrier is overcome, the super-saturated silicic acid solution (such as H4SiO4 in xylem sap) does not necessarily have to precipitate, just has the potential ability to precipitation. Thus, based on the mediators of SiO2 deposition at the thermodynamic-driven stage, it is difficult to evaluate whether the process of plant silicification is active or passive. We conclude that the characteristics of kinetic-drivers determine the mechanism of plant silicification.

Mutation of CESA1 phosphorylation site influences pectin synthesis and methylesterification with a role in seed development.

Cell wall biogenesis is required for the production of seeds of higher plants. However, little is known about regulatory mechanisms underlying cell wall biogenesis during seed formation. Here we show a role for the phosphorylation of Arabidopsis cellulose synthase 1 (AtCESA1) in modulating pectin synthesis and methylesterification in seed coat mucilage. A phosphor-null mutant of AtCESA1 on T166 (AtCESA1T166A) was constructed and introduced into a null mutant of AtCESA1 (Atcesa1-1). The resulting transgenic lines showed a slight but significant decrease in cellulose contents in mature seeds. Defects in cellulosic ray architecture along with reduced levels of non-adherent and adherent mucilage were observed on the seeds of the AtCESA1T166A mutant. Reduced mucilage pectin synthesis was also reflected by a decrease in the level of uronic acid. Meanwhile, an increase in the degree of pectin methylesterification was also observed in the seed coat mucilage of AtCESA1T166A mutant. Change in seed development was further reflected by a delayed germination and about 50% increase in the accumulation of proanthocyanidins, which is known to bind pectin and inhibit seed germination as revealed by previous studies. Taken together, the results suggest a role of AtCESA1 phosphorylation on T166 in modulating mucilage pectin synthesis and methylesterification as well as cellulose synthesis with a role in seed development.

Exogenous Application of Low-Concentration Sugar Enhances Brassinosteroid Signaling for Skotomorphogenesis by Promoting BIN2 Degradation.

In plants, seedling growth is subtly controlled by multiple environmental factors and endogenous phytohormones. The cross-talk between sugars and brassinosteroid (BR) signaling is known to regulate plant growth; however, the molecular mechanisms that coordinate hormone-dependent growth responses with exogenous sucrose in plants are incompletely understood. Skotomorphogenesis is a plant growth stage with rapid elongation of the hypocotyls. In the present study, we found that low-concentration sugars could improve skotomorphogenesis in a manner dependent on BR biosynthesis and TOR activation. However, accumulation of BZR1 in bzr1-1D mutant plants partially rescued the defects of skotomorphogenesis induced by the TOR inhibitor AZD, and these etiolated seedlings displayed a normal phenotype like that of wild-type seedlings in response to both sucrose and non-sucrose treatments, thereby indicating that accumulated BZR1 sustained, at least partially, the sucrose-promoted growth of etiolated seedlings (skotomorphogenesis). Moreover, genetic evidence based on a phenotypic analysis of bin2-3bil1bil2 triple-mutant and gain-of-function bin2-1 mutant plant indicated that BIN2 inactivation was conducive to skotomorphogenesis in the dark. Subsequent biochemical and molecular analyses enabled us to confirm that sucrose reduced BIN2 levels via the TOR-S6K2 pathway in etiolated seedlings. Combined with a determination of the cellulose content, our results indicated that sucrose-induced BIN2 degradation led to the accumulation of BZR1 and the enhancement of cellulose synthesis, thereby promoting skotomorphogenesis, and that BIN2 is the converging node that integrates sugar and BR signaling.

Plant silicon-cell wall complexes: Identification, model of covalent bond formation and biofunction.

Silicon (Si) is the second most abundant element on earth crust, consisting primarily of silicate minerals. Si is found in the tissues of almost all terrestrial plants and is mostly deposited in specialized cells or cell walls as amorphous silica. Numerous discoveries have shown that in addition to non-covalent interactions through amorphous silica, Si can form covalent bonds with plant cell wall components such as hemicelluloses, pectin and lignin. The covalent bonds may be formed via Si-O-C linkages between monosilicic acid (H4SiO4) and cis-diols of cell wall polysaccharides or lignin. The covalently bound organosilicon, independent of amorphous inorganic silica, may play a crucial role in plant cell wall structure and remodeling and thus plant growth and its resistance against biotic and abiotic stresses. This review discusses the existing research on the discovery of plant silicon-cell wall complexes and proposes a model of their covalent bond formation and biofunction.

Cell wall-bound silicon optimizes ammonium uptake and metabolism in rice cells.

Background and Aims: Turgor-driven plant cell growth depends on cell wall structure and mechanics. Strengthening of cell walls on the basis of an association and interaction with silicon (Si) could lead to improved nutrient uptake and optimized growth and metabolism in rice (Oryza sativa). However, the structural basis and physiological mechanisms of nutrient uptake and metabolism optimization under Si assistance remain obscure. Methods: Single-cell level biophysical measurements, including in situ non-invasive micro-testing (NMT) of NH4+ ion fluxes, atomic force microscopy (AFM) of cell walls, and electrolyte leakage and membrane potential, as well as whole-cell proteomics using isobaric tags for relative and absolute quantification (iTRAQ), were performed. Key Results: The altered cell wall structure increases the uptake rate of the main nutrient NH4+ in Si-accumulating cells, whereas the rate is only half in Si-deprived counterparts. Conclusions: Rigid cell walls enhanced by a wall-bound form of Si as the structural basis stabilize cell membranes. This, in turn, optimizes nutrient uptake of the cells in the same growth phase without any requirement for up-regulation of transmembrane ammonium transporters. Optimization of cellular nutrient acquisition strategies can substantially improve performance in terms of growth, metabolism and stress resistance.

iTRAQ-based proteomic analysis reveals the mechanisms of silicon-mediated cadmium tolerance in rice (Oryza sativa) cells.

Silicon (Si) can alleviate cadmium (Cd) stress in rice (Oryza sativa) plants, however, the understanding of the molecular mechanisms at the single-cell level remains limited. To address these questions, we investigated suspension cells of rice cultured in the dark environment in the absence and presence of Si with either short- (12 h) or long-term (5 d) Cd treatments using a combination of isobaric tags for relative and absolute quantitation (iTRAQ), fluorescent staining, and inductively coupled plasma mass spectroscopy (ICP-MS). We identified 100 proteins differentially regulated by Si under the short- or long-term Cd stress. 70% of these proteins were down-regulated, suggesting that Si may improve protein use efficiency by maintaining cells in the normal physiological status. Furthermore, we showed two different mechanisms for Si-mediated Cd tolerance. Under the short-term Cd stress, the Si-modified cell walls inhibited the uptake of Cd ions into cells and consequently reduced the expressions of glycosidase, cell surface non-specific lipid-transfer proteins (nsLTPs), and several stress-related proteins. Under the long-term Cd stress, the amount of Cd in the cytoplasm in Si-accumulating (+Si) cells was decreased by compartmentation of Cd into vacuoles, thus leading to a lower expression of glutathione S-transferases (GST). These results provide protein-level insights into the Si-mediated Cd detoxification in rice single cells.