Unveiling mechanisms of silicon-mediated resistance to chromium stress in rice using a newly-developed hierarchical system.

Silicon (Si) has been well-known to enhance plant resistance to heavy-metal stress. However, the mechanisms by which silicon mitigates heavy-metal stress in plants are not clear. In particular, information regarding the role of Si in mediating resistance to heavy-metal stress at a single cell level is still lacking. Here, we developed a hierarchical system comprising the plant, protoplast, and suspension cell subsystems to investigate the mechanisms by which silicon helps to alleviate the toxic effects of trivalent chromium [Cr(III)] in rice. Our results showed that in whole-plant subsystem silicon reduced shoot Cr(III) concentration, effectively alleviating Cr(III) stress in seedlings and causing changes in antioxidant enzyme activities similar to those observed at lower Cr(III) concentrations without silicon added. However, in protoplast subsystem lacking the cell wall, no silicon deposition occurred, leading to insignificant changes in cell survival or antioxidation processes under Cr(III) stress. Conversely, in suspension cell subsystem, silicon supplementation substantially improved cell survival and changes in antioxidant enzyme activities under Cr(III) stress. This is due to the fact that >95% of silicon was on the cell wall, reducing Cr(III) concentration in cells by 7.7%-10.4%. Collectively, the results suggested that the silicon deposited on the cell wall hindered Cr(III) bio-uptake, which consequently delayed Cr(III)-induced changes in antioxidant enzyme activities. This research emphasizes the significance of cell walls in Si-alleviated heavy-metal stress and deepens our understanding of silicon functioning in plants. Furthermore, the hierarchical system has great potential for application in studying the functioning of other elements in plant cell walls.

Silicon-phosphorus pathway mitigates heavy metal stress by buffering rhizosphere acidification.

Heavy metal pollution threatens food security, and rhizosphere acidification will increase the bioavailability of heavy metals. As a beneficial element in plants, silicon can relieve heavy metal stress. However, less attention has been paid to its effects on plant rhizosphere processes. Here, we show that for Japonica (Nipponbare and Oochikara) and Indica (Jinzao 47) rice cultivars, the degree of root acidification was significantly reduced after silicon uptake, and the total organic carbon, citric acid, and malic acid concentrations in rice root exudates were significantly reduced. We further confirmed the results by q-PCR that the expressions of proton pump and organic acid secretion genes were down-regulated by 35-61 % after silicon treatment. Intriguingly, phosphorus allocation, an intensively studied mechanism of rhizosphere acidification, was altered by silicon treatment. Specifically, among total phosphorus in rice seedlings, the soluble proportion increased from 52.0 % to 61.7 %, while cell wall phosphorus decreased from 48.0 % to 32.3 %. Additionally, silicon-mediated alleviation of rhizosphere acidification has positive effects on relieving heavy metal stress. Simulation revealed that low acidification of the nutrient solution resulted in a decrease in bioavailable heavy metal concentrations, thereby reducing rice uptake. We further confirmed that the impediment of rhizosphere acidification led to free-state Cr3+ in solutions decreasing by 43 % and contributed up to 63 % of silicon's mitigation of Cr(III) stress. Overall, we propose a novel mechanism in which silicon reduces heavy metal absorption by increasing plant soluble phosphorus concentration and buffering rhizosphere acidification. This paper provides a unique insight into the role of silicon in plants and, more importantly, a theoretical reference for the rational application of silicon fertilizer to improve phosphorus utilization efficiency, alleviate heavy metal stress, and balance soil pH.