Silicon (Si) being considered as a non-essential element for plant growth and development finds its role in providing several benefits to the plant, especially under stress conditions. Thus, Si can be regarded as “multi-talented” quasi-essential element. It is the most abundant element present in the earth’s crust after oxygen predominantly as a silicon dioxide (SiO2), a form plants cannot utilize. Plants take up Si into their root from the soil in the plant-available forms (PAF) such as silicic acid or mono silicic acid [Si(OH)4 or H4SiO4]. Nevertheless, besides being abundantly available, the PAF of Si in the soil is mostly a limiting factor. To improve Si-uptake and derived benefits therein in plants, understanding the molecular basis of Si-uptake and transport within the tissues has great importance. Numerous Si-transporters (influx and efflux) have been identified in both monocot and dicot plants. A difference in the root anatomy of both monocot and dicot plants leads to a difference in the Si-uptake mechanism. In the present review, Si-transporters identified in different species, their evolution and the Si-uptake mechanism have been addressed. Further, the role of Si in biotic and abiotic stress tolerance has been discussed. The information provided here will help to plan the research in a better way to develop more sustainable cropping system by harnessing Si-derived benefits.

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Role of silicon in plant stress tolerance: opportunities to achieve a sustainable cropping system.

Silicon (Si) is the most copious element of existence in the lithosphere but still it has not been added into the essential element list. The imperative role of Si in triggering growth and development of plants has been identified. It is of paramount importance in regulating overall physiological and metabolic characteristics of the plants. Being considered as a non-essential element, it has been known to occur at about 30%, majority of its presence is there in rocks as mineral salts. It has been regarded as multitalented or quasi-element on earth's crust that can be efficiently taken up by plants and translocated further towards aerial parts via transpiration phenomenon. It has also been known to mitigate different biotic and abiotic stressed conditions from plants as the need of the hour owing to its eco-friendly nature. However, the mechanisms associated with their stress attenuation are associated with Reactive Oxygen Species (ROS) scavenging, activation of antioxidative defense responses and phytohormonal signaling. Also, biotic stress factors can be ameliorated through accumulation of Si within epidermal tissues or pathogenesis-related host defense mechanisms. To explore further, omics-mediated studies have been further discussed to shed light on the stress mitigating processes. Further, to improve our understanding for Si-mediated benefits in plants we need to explore the molecular mechanisms of Si uptake, transport and gene expression studies revealing their mitigate properties. In the present review, we have evolved the Si-based studies in plants associated with its transport, uptake and accumulation. Apart from this, we have also discussed about their role in ameliorating stresses from plants by activating their defenses. Moreover, their roles in plant hormonal crosstalk have also been elucidated. Above all, we have also revealed the role of Si-Nanoparticles (SiNPs) in improving stress potential of plants along with stimulation of plant productivities via omics-based approaches.

Silicon and Plants: Current Knowledge and Future Prospects

Salinity affects around 20% of all arable land while an even larger area suffers from recurrent drought. Together these stresses suppress global crop production by as much as 50% and their impacts are predicted to be exacerbated by climate change. Infrastructure and management practices can mitigate these detrimental impacts, but are costly. Crop breeding for improved tolerance has had some success but is progressing slowly and is not keeping pace with climate change. In contrast, Silicon (Si) is known to improve plant tolerance to a range of stresses and could provide a sustainable, rapid and cost-effective mitigation method. The exact mechanisms are still under debate but it appears Si can relieve salt stress via accumulation in the root apoplast where it reduces "bypass flow of ions to the shoot. Si-dependent drought relief has been linked to lowered root hydraulic conductance and reduction of water loss through transpiration. However, many alternative mechanisms may play a role such as altered gene expression and increased accumulation of compatible solutes. Oxidative damage that occurs under stress conditions can be reduced by Si through increased antioxidative enzymes while Si-improved photosynthesis has also been reported. Si fertilizer can be produced relatively cheaply and to assess its economic viability to improve crop stress tolerance we present a cost-benefit analysis. It suggests that Si fertilization may be beneficial in many agronomic settings but may be beyond the means of smallholder farmers in developing countries. Si application may also have disadvantages, such as increased soil pH, less efficient conversion of crops into biofuel and reduced digestibility of animal fodder. These issues may hamper uptake of Si fertilization as a routine agronomic practice. Here, we critically evaluate recent literature, quantifying the most significant physiological changes associated with Si in plants under drought and salinity stress. Analyses show that metrics associated with photosynthesis, water balance and oxidative stress all improve when Si is present during plant exposure to salinity and drought. We further conclude that most of these changes can be explained by apoplastic roles of Si while there is as yet little evidence to support biochemical roles of this element.

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Is Silicon a Panacea for Alleviating Drought and Salt Stress in Crops

Silicon (Si), the second most predominant element in the earth crust consists of numerous benefits to plant. Beneficial effect of Si has been apparently visible under both abiotic and biotic stress conditions in plants. Supplementation of Si improved physiology and yield on several important agricultural and horticultural crops. Salinity is one of the major abiotic stresses that affect growth and yield. The presence of high concentration of salt in growing medium causes oxidative, osmotic, and ionic stresses to plants. In extreme conditions salinity affects soil, ground water, and limits agricultural production. Si ameliorates salt stress in several plants. The Si mediated stress mitigation involves various regulatory mechanisms such as photosynthesis, detoxification of harmful reactive oxygen species using antioxidant and non-antioxidants, and proper nutrient management. In the present review, Si mediated alleviation of salinity stress in plants through the regulation of photosynthesis, root developmental changes, redox homeostasis equilibrium, and regulation of nutrients have been dealt in detail.

Mechanisms of Silicon-Mediated Amelioration of Salt Stress in Plants

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.

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

Low temperature is one of the major abiotic stresses which severely affects the productivity and the geographical distribution of rice (Oryza sativa). Silicon is considered a broad spectrum alleviator to combat stress in rice plant. Rice root absorbs silicon by a silicon transporter, Low silicon gene 1 (Lsi1). To gain a better understanding of cold stress responses triggered by overexpression of Lsi1 in rice (Oryza sativa L.), we carried out physiological and molecular studies between Lsi1-overexpression Dular (Lsi1-D) and its wild type (WD). Two leaf stage rice seedlings of above mentioned both lines were treated at 15 °C/12 °C (day/night) for 7 days. WD seedling leaves were turned comparatively yellow as compared to Lsi1-D seedlings. Microscopic studies showed significantly more deposition of silicon bodies in epidermal cells of Lsi1-D leaf seedlings in comparison with WD leaves. Lsi1-D leaves comparatively, depicted more SOD, POD and CAT activity, chlorophyll a, b contents in consistency with more silicon concentration. Protein extraction was carried out from whole seedling of both lines and further analyzed by tandem mass tag quantitative proteomics approach with double replicates. Among 393 reproducible proteins, 63 were up-regulated and 39 proteins were down-regulated. The total cold responsive differential proteins were involved in several processes, i.e. photosynthesis, signal transduction, redox homeostasis, hormone metabolism, carbohydrate metabolism, cell wall organization, N-assimilation, protein processing and secondary metabolism. We confirmed up-regulation of key proteins involved in cold-responsive pathway at mRNA level through qPCR such as chlorophyll a–b binding protein 1, peroxidase 2, signaling G-proteins RIC1, aquaporin PIP1.2, 1, 4-alpha-glucan branching enzyme, germin-like protein subfamily 2 member 4 and germin-like protein subfamily 8 member 2. In conclusion, our study provides new insights into cold stress responses in rice seedlings triggered by Lsi1-overexpression defense pathway.

Haiping Zheng

Papers

Quantitative proteomics study on Lsi1 in regulation of rice (Oryza sativa L.) cold resistance