Microbial communities play a pivotal role in the functioning of plants by influencing their physiology and development. While many members of the rhizosphere microbiome are beneficial to plant growth, also plant pathogenic microorganisms colonize the rhizosphere striving to break through the protective microbial shield and to overcome the innate plant defense mechanisms in order to cause disease. A third group of microorganisms that can be found in the rhizosphere are the true and opportunistic human pathogenic bacteria, which can be carried on or in plant tissue and may cause disease when introduced into debilitated humans. Although the importance of the rhizosphere microbiome for plant growth has been widely recognized, for the vast majority of rhizosphere microorganisms no knowledge exists. To enhance plant growth and health, it is essential to know which microorganism is present in the rhizosphere microbiome and what they are doing. Here, we review the main functions of rhizosphere microorganisms and how they impact on health and disease. We discuss the mechanisms involved in the multitrophic interactions and chemical dialogues that occur in the rhizosphere. Finally, we highlight several strategies to redirect or reshape the rhizosphere microbiome in favor of microorganisms that are beneficial to plant growth and health.

The rhizosphere microbiome: significance of plant beneficial, plant pathogenic, and human pathogenic microorganisms

Preface.- Contributors.- List of Abbreviations.- Section 1: Basic Principles: Introduction.-Sources of Heavy Metals and Metalloids in Soils.- Chemistry of Heavy Metals and Metalloids in Soils.- Methods for the Determination of Heavy Metals and Metalloids in Soils.- Effects of Heavy Metals and Metalloids on Soil Organisms.- Soil-Plant Relationships of Heavy Metals and Metalloids.- Heavy Metals and Metalloids as Micronutrients for Plants and Animals.-Critical Loads of Heavy Metals for Soils.- Section 2: Key Heavy Metals And Metalloids: Arsenic.- Cadmium.- Chromium and Nickel.- Cobalt and Manganese.- Copper.-Lead.- Mercury.- Selenium.- Zinc.- Section 3: Other Heavy Metals And Metalloids Of Potential Environmental Significance: Antimony.- Barium.- Gold.- Molybdenum.- Silver.- Thallium.- Tin.- Tungsten.- Uranium.- Vanadium.- Glossary of Specialized Terms.- Index.

Heavy metals in soils : trace metals and metalloids in soils and their bioavailability

Heavy metal hyperaccumulating plants: how and why do they do it? And what makes them so interesting?

Remediation of heavy metal(loid)s contaminated soils - To mobilize or to immobilize?

https://www.cell.com/trends/plant-science/pdf/S1360-1385(06)00163-4.pdf

Silicon uptake and accumulation in higher plants

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.

A silicon transporter in rice

Arsenic poisoning affects millions of people worldwide. Human arsenic intake from rice consumption can be substantial because rice is particularly efficient in assimilating arsenic from paddy soils, although the mechanism has not been elucidated. Here we report that two different types of transporters mediate transport of arsenite, the predominant form of arsenic in paddy soil, from the external medium to the xylem. Transporters belonging to the NIP subfamily of aquaporins in rice are permeable to arsenite but not to arsenate. Mutation in OsNIP2;1 (Lsi1, a silicon influx transporter) significantly decreases arsenite uptake. Furthermore, in the rice mutants defective in the silicon efflux transporter Lsi2, arsenite transport to the xylem and accumulation in shoots and grain decreased greatly. Mutation in Lsi2 had a much greater impact on arsenic accumulation in shoots and grain in field-grown rice than Lsi1. Arsenite transport in rice roots therefore shares the same highly efficient pathway as silicon, which explains why rice is efficient in arsenic accumulation. Our results provide insight into the uptake mechanism of arsenite in rice and strategies for reducing arsenic accumulation in grain for enhanced food safety.

https://www.pnas.org/content/pnas/105/29/9931.full.pdf

Transporters of arsenite in rice and their role in arsenic accumulation in rice grain

Silicon is an important nutrient for the optimal growth and sustainable production of rice. Rice accumulates up to 10% silicon in the shoot, and this high accumulation is required to protect the plant from multiple abiotic and biotic stresses. A gene, Lsi1, that encodes a silicon influx transporter has been identified in rice. Here we describe a previously uncharacterized gene, low silicon rice 2 (Lsi2), which has no similarity to Lsi1. This gene is constitutively expressed in the roots. The protein encoded by this gene is localized, like Lsi1, on the plasma membrane of cells in both the exodermis and the endodermis, but in contrast to Lsi1, which is localized on the distal side, Lsi2 is localized on the proximal side of the same cells. Expression of Lsi2 in Xenopus oocytes did not result in influx transport activity for silicon, but preloading of the oocytes with silicon resulted in a release of silicon, indicating that Lsi2 is a silicon efflux transporter. The identification of this silicon transporter revealed a unique mechanism of nutrient transport in plants: having an influx transporter on one side and an efflux transporter on the other side of the cell to permit the effective transcellular transport of the nutrients.

An efflux transporter of silicon in rice

The accumulation of silicon (Si) in the shoots varies considerably among plant species, but the mechanism responsible for this variation is poorly understood. The uptake system of Si was investigated in terms of the radial transport from the external solution to the root cortical cells and the release of Si from the cortical cells to the xylem in rice, cucumber, and tomato, which differ greatly in shoot Si concentration. Symplasmic solutions of the root tips were extracted by centrifugation. The concentrations of Si in the root-cell symplast in all species were higher than that in the external solution, although the concentration in rice was 3- and 5-fold higher than that in cucumber and tomato, respectively. A kinetic study showed that the radial transport of Si was mediated by a transporter with a Km value of 0.15 mM in all species, but with different Vmax values in the order of rice>cucumber>tomato. In the presence of the metabolic inhibitor 2,4-dinitrophenol, and at low temperature, the Si concentration in the root-cell symplast decreased to a level similar to that of the apoplasmic solution. These results suggest that both transporter-mediated transport and passive diffusion of Si are involved in the radial transport of Si and that the transporter-mediated transport is an energy-dependent process. The Si concentration of xylem sap in rice was 20- and 100-fold higher than that in cucumber and tomato, respectively. In contrast to rice, the Si concentration in the xylem sap was lower than that in the external solution in cucumber and tomato. A kinetic study showed that xylem loading of Si was also mediated by a kind of transporter in rice, but by passive diffusion in cucumber and tomato. These results indicate that a higher density of transporter for radial transport and the presence of a transporter for xylem loading are responsible for the high Si accumulation in rice.

Uptake system of silicon in different plant species.

Soluble ionic aluminum (Al) inhibits root growth and reduces crop production on acid soils Al-resistant cultivars of barley (Hordeum vulgare L) detoxify Al by secreting citrate from the roots, but the responsible gene has not been identified yet Here, we identified a gene (HvAACT1) responsible for the Al-activated citrate secretion by fine mapping combined with microarray analysis, using an Al-resistant cultivar, Murasakimochi, and an Al-sensitive cultivar, Morex This gene belongs to the multidrug and toxic compound extrusion (MATE) family and was constitutively expressed mainly in the roots of the Al-resistant barley cultivar Heterologous expression of HvAACT1 in Xenopus oocytes showed efflux activity for (14)C-labeled citrate, but not for malate Two-electrode voltage clamp analysis also showed transport activity of citrate in the HvAACT1-expressing oocytes in the presence of Al Overexpression of this gene in tobacco enhanced citrate secretion and Al resistance compared with the wild-type plants Transiently expressed green fluorescent protein-tagged HvAACT1 was localized at the plasma membrane of the onion epidermal cells, and immunostaining showed that HvAACT1 was localized in the epidermal cells of the barley root tips A good correlation was found between the expression of HvAACT1 and citrate secretion in 10 barley cultivars differing in Al resistance Taken together, our results demonstrate that HvAACT1 is an Al-activated citrate transporter responsible for Al resistance in barley

Namiki Mitani

Papers

A silicon transporter in rice

Transporters of arsenite in rice and their role in arsenic accumulation in rice grain

An efflux transporter of silicon in rice

Uptake system of silicon in different plant species.

An Aluminum-Activated Citrate Transporter in Barley