· Root responses to lack of iron (Fe) have mainly been studied in nutrient solution experiments devoid of silicon (Si). Here we investigated how Si ameliorates Fe deficiency in cucumber (Cucumis sativus) with focus on the storage and utilization of Fe in the root apoplast. · A combined approach was performed including analyses of apoplastic Fe, reduction-based Fe acquisition and Fe-mobilizing compounds in roots along with the expression of related genes. · Si-treated plants accumulated higher concentrations of root apoplastic Fe, which rapidly decreased when Fe was withheld from the nutrient solution. Under Fe-deficient conditions, Si also increased the accumulation of Fe-mobilizing compounds in roots. Si supply stimulated root activity of Fe acquisition at the early stage of Fe deficiency stress through regulation of gene expression levels of proteins involved in Fe acquisition. However, when the period of Fe deprivation was extended, these reactions further decreased as a consequence of Si-induced enhancement of the Fe status of the plants. · This work provides new evidence for the beneficial role of Si in plant nutrition and clearly indicates that Si-mediated alleviation of Fe deficiency includes an increase of the apoplastic Fe pool in roots and an enhancement of Fe acquisition.

Silicon (Si) is not classified as an essential element for plants, but numerous studies have demonstrated its beneficial effects in a variety of species and environmental conditions, including low nutrient availability. Application of Si shows the potential to increase nutrient availability in the rhizosphere and root uptake through complex mechanisms, which still remain unclear. Silicon-mediated transcriptional regulation of element transporters for both root acquisition and tissue homeostasis has recently been suggested as an important strategy, varying in detail depending on plant species and nutritional status. Here, we summarize evidence of Si-mediated acquisition, uptake and translocation of nutrients: nitrogen (N), phosphorus (P), potassium (K), calcium (Ca), magnesium (Mg), sulfur (S), iron (Fe), zinc (Zn), manganese (Mn), copper (Cu), boron (B), chlorine (Cl), and nickel (Ni) under both deficiency and excess conditions. In addition, we discuss interactions of Si-with beneficial elements: aluminum (Al), sodium (Na), and selenium (Se). This review also highlights further research needed to improve understanding of Si-mediated acquisition and utilization of nutrients and vice versa nutrient status-mediated Si acquisition and transport, both processes which are of high importance for agronomic practice (e.g., reduced use of fertilizers and pesticides).

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The results suggest that Si enhances remobilization of Fe from older to younger leaves by a more efficient NA-mediated Fe transport via the phloem. In addition, from this and previous work, a model is proposed of how Si acts to improve Fe homeostasis under Fe deficiency in cucumber.