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      The controversies of silicon's role in plant biology

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          Abstract

          Contents Summary 67 I. Introduction 68 II. Silicon transport in plants: to absorb or not to absorb 69 III. The role of silicon in plants: not just a matter of semantics 71 IV. Silicon and biotic stress: beyond mechanical barriers and defense priming 76 V. Silicon and abiotic stress: a proliferation of proposed mechanisms 78 VI. The apoplastic obstruction hypothesis: a working model 79 VII. Perspectives and conclusions 80 Acknowledgements 81 References 81 SUMMARY: Silicon (Si) is not classified as an essential plant nutrient, and yet numerous reports have shown its beneficial effects in a variety of species and environmental circumstances. This has created much confusion in the scientific community with respect to its biological roles. Here, we link molecular and phenotypic data to better classify Si transport, and critically summarize the current state of understanding of the roles of Si in higher plants. We argue that much of the empirical evidence, in particular that derived from recent functional genomics, is at odds with many of the mechanistic assertions surrounding Si's role. In essence, these data do not support reports that Si affects a wide range of molecular-genetic, biochemical and physiological processes. A major reinterpretation of Si's role is therefore needed, which is critical to guide future studies and inform agricultural practice. We propose a working model, which we term the 'apoplastic obstruction hypothesis', which attempts to unify the various observations on Si's beneficial influences on plant growth and yield. This model argues for a fundamental role of Si as an extracellular prophylactic agent against biotic and abiotic stresses (as opposed to an active cellular agent), with important cascading effects on plant form and function.

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          A silicon transporter in rice.

          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.
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            SILICON.

            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.
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              Increasing CO2 threatens human nutrition.

              Dietary deficiencies of zinc and iron are a substantial global public health problem. An estimated two billion people suffer these deficiencies, causing a loss of 63 million life-years annually. Most of these people depend on C3 grains and legumes as their primary dietary source of zinc and iron. Here we report that C3 grains and legumes have lower concentrations of zinc and iron when grown under field conditions at the elevated atmospheric CO2 concentration predicted for the middle of this century. C3 crops other than legumes also have lower concentrations of protein, whereas C4 crops seem to be less affected. Differences between cultivars of a single crop suggest that breeding for decreased sensitivity to atmospheric CO2 concentration could partly address these new challenges to global health.
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                Author and article information

                Journal
                New Phytologist
                New Phytol
                Wiley
                0028646X
                July 14 2018
                Affiliations
                [1 ]Département de Phytologie; Faculté des Sciences de l'Agriculture et de l'Alimentation (FSAA); Université Laval; Québec QC G1V 0A6 Canada
                [2 ]Agriculture and Agri-Food Canada; 101 Route 100 Morden MB R6M 1Y5 Canada
                [3 ]Biosecurity and Food Safety; NSW Department of Primary Industries; Elizabeth Macarthur Agricultural Institute; Menangle NSW 2568 Australia
                [4 ]Graham Centre for Agricultural Innovation; Wagga Wagga NSW 2650 Australia
                [5 ]Institute of Plant Science and Resources; Okayama University; Chuo 2-20-1 Kurashiki 710-0046 Japan
                [6 ]School of Agriculture and Food; Faculty of Veterinary and Agricultural Sciences; The University of Melbourne; Melbourne Vic. 3010 Australia
                Article
                10.1111/nph.15343
                30007071
                a0d89202-31f8-4ee2-8003-b9efb48f7d6a
                © 2018

                http://doi.wiley.com/10.1002/tdm_license_1.1

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