Uncommon heavy metals, metalloids and their plant toxicity: a review

Silicon is beneficial to plant growth and helps plants to overcome abiotic and biotic stresses by preventing lodging (falling over) and increasing resistance to pests and diseases, as well as other stresses. Silicon is essential for high and sustainable production of rice, but the molecular mechanism responsible for the uptake of silicon is unknown. Here we describe the Low silicon rice 1 (Lsi1) gene, which controls silicon accumulation in rice, a typical silicon-accumulating plant. This gene belongs to the aquaporin family and is constitutively expressed in the roots. Lsi1 is localized on the plasma membrane of the distal side of both exodermis and endodermis cells, where casparian strips are located. Suppression of Lsi1 expression resulted in reduced silicon uptake. Furthermore, expression of Lsi1 in Xenopus oocytes showed transport activity for silicon only. The identification of a silicon transporter provides both an insight into the silicon uptake system in plants, and a new strategy for producing crops with high resistance to multiple stresses by genetic modification of the root's silicon uptake capacity.

Silicon is present in plants in amounts equivalent to those of such macronutrient elements as calcium, magnesium, and phosphorus, and in grasses often at higher levels than any other inorganic constituent. Yet except for certain algae, including prominently the diatoms, and the Equisetaceae (horsetails or scouring rushes), it is not considered an essential element for plants. As a result it is routinely omitted from formulations of culture solutions and considered a nonentity in much of plant physiological research. But silicon-deprived plants grown in conventional nutrient solutions to which silicon has not been added are in many ways experimental artifacts. They are often structurally weaker than silicon-replete plants, abnormal in growth, development, viability, and reproduction, more susceptible to such abiotic stresses as metal toxicities, and easier prey to disease organisms and to herbivores ranging from phytophagous insects to mammals. Many of these same conditions afflict plants in silicon-poor soils-and there are such. Taken together, the evidence is overwhelming that silicon should be included among the elements having a major bearing on plant life.

Silicon (Si) in plants provides structural support and improves tolerance to diseases, drought and metal toxicity. Shoot Si concentrations are generally considered to be greater in monocotyledonous than in non-monocot plant species. The phylogenetic variation in the shoot Si concentration of plants reported in the primary literature has been quantified. Studies were identified which reported Si concentrations in leaf or non-woody shoot tissues from at least two plant species growing in the same environment. Each study contained at least one species in common with another study. Meta-analysis of the data revealed that, in general, ferns, gymnosperms and angiosperms accumulated less Si in their shoots than non-vascular plant species and horsetails. Within angiosperms and ferns, differences in shoot Si concentration between species grouped by their higher-level phylogenetic position were identified. Within the angiosperms, species from the commelinoid monocot orders Poales and Arecales accumulated substantially more Si in their shoots than species from other monocot clades. A high shoot Si concentration is not a general feature of monocot species. Information on the phylogenetic variation in shoot Si concentration may provide useful palaeoecological and archaeological information, and inform studies of the biogeochemical cycling of Si and those of the molecular genetics of Si uptake and transport in plants.