Silicon-mediated amelioration of zinc toxicity in rice (Oryza sativa L.) seedlings

Background and aims

Silicon (Si) was suggested to enhance plant resistance to toxic elements, and its beneficial role was mainly based on external and internal plant mechanisms. This work aimed at investigating the internal effect of Si on zinc (Zn) detoxification to rice (Oryza sativa L., cv. Tian You 116) seedlings.

Methods

In a hydroponic experiment, we examined the uptake, xylem loading and localization of Zn in rice seedlings under the condition of 200?μM Zn contamination with the additional silicate supply at three levels ( 0, 0.5 and 1.8?mM).

Results

The silicate addition significantly increased the seedling biomass, and decreased Zn concentration in both root and shoot of seedlings and in xylem sap flow. Zinpyr-1 fluorescence test and Energy-dispersive X-ray spectroscopy analysis showed the concentration of biologically active Zn²⁺ decreased, and Zn and Si co-localized in the cell wall of metabolically less active tissues, especially in sclerenchyma of root. The fractionation analysis further supported silicate supply increased about 10% the cell wall bound fraction of Zn.

Conclusions

This study suggests the Si-assisted Zn tolerance of rice is mainly due to the reduction of uptake and translocation of excess Zn, and a stronger binding of Zn in the cell wall of less bioactive tissues might also contribute to some degree.     

Soil microalgae modulate grain arsenic accumulation by reducing dimethylarsinic acid and enhancing nutrient uptake in rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.)

Background and aims

Microalgae are ubiquitous in paddy soils. However, their roles in arsenic (As) accumulation and transport in rice plants remains unknown.

Methods

Two green algae and five cyanobacteria were used in pot experiments under continuously flooded conditions to ascertain whether a microalgal inoculation could influence rice growth and rice grain As accumulation in plants grown in As-contaminated soils.

Results

The microalgal inoculation greatly enhanced nutrient uptake and rice growth. The presence of representative microalga Anabaena azotica did not significantly differ the grain inorganic As concentrations but remarkably decreased the rice root and grain DMA concentrations. The translocation of As from roots to grains was also markedly decreased by rice inoculated with A. azotica. This subsequently led to a decrease in the total As concentration in rice grains.

Conclusions

The results of the study indicate that the microalgal inoculation had a strong influence on soil pH, soil As speciation, and soil nutrient bioavailability, which significantly affected the rice growth, nutrient uptake, and As accumulation and translocation in rice plants. The results suggest that algae inoculation can be an effective strategy for improving nutrient uptake and reducing As translocation from roots to grains by rice grown in As-contaminated paddy soils.

     

Differential toxicity and accumulation of inorganic and methylated arsenic in rice

Background and aims

Efficient accumulation of arsenic (As) in rice (Oryza sativa L.) poses a potential health risk to rice consumers. The aim of this study was to investigate the mechanisms of uptake, transport and distribution of inorganic arsenic (As_i) and dimethylarsinic acid (DMA) in rice plants.

Methods

Rice was exposed to As_i (As(V)) and DMA in hydroponics. High-performance liquid chromatography inductively coupled plasma mass spectrometry (HPLC-ICP-MS) and synchrotron X-ray fluorescence (SXRF) microprobe were used to determine As concentration and the in situ As distribution.

Results

DMA induced abnormal florets before flowering and caused a sharp decline in the seed setting rate after flowering compared to As_i. Rice grains accumulated 2-fold higher DMA than As_i. The distribution of As_i concentration (root?>?leaf?>?husk?>?caryopsis) in As(V) treatments was different from that of the DMA concentration (caryopsis?>?husk?>?root?≥?leaf) in DMA treatments. SXRF showed that As_i mainly accumulated in the vascular trace of caryopsis with limited distribution to the endosperm, whereas DMA was observed in both tissues.

Conclusions

DMA tended to accumulate in caryopsis and induced higher toxicity to the reproductive tissues resulting in markedly reduced grain yield, whereas As_i mainly remained in the vegetative tissues and had no significant effect on yield. DMA is more toxic than As_i to the reproductive tissues when both of them are at similar concentrations in nutrient solution.     

Effects of silicon on aluminum toxicity in upland rice plants

Aims

This study aimed to determine the capacity of Si to mitigate Al toxicity in upland rice plants (Oryza sativa L.) by evaluating plant growth and the Si and Al uptake kinetics.

Methods

Plants were grown for 40 days, after which the Si and Al uptake kinetics (Cmin, Km and Imax) were analyzed. Then, the shoots and roots were separated, and the dry matter, root morphology and Si and Al concentration and accumulation in the plant were evaluated.

Results

Aluminum decreased plant growth and the Si uptake capacity by decreasing the root growth and Si transport system efficiency in the upland rice roots (> Km and > Cmin). Silicon mitigated Al toxicity in the upland rice plants by decreasing Al transport to the plant shoots, although it did not reduce the Al uptake rate (Imax). Si treatment increased the growth of upland rice plant shoots grown in the presence of Al without influencing the root growth. The alleviation of Al toxicity by Si is more evident in the susceptible upland rice cultivar Maravilha.

Conclusions

Silicon mitigated Al toxicity in the upland rice plants by decreasing Al transport to the plant shoots but did not reduce the Al uptake rate by roots.

     

Grain Unloading of Arsenic Species in Rice

Rice (Oryza sativa) is the staple food for over half the world''s population yet may represent a significant dietary source of inorganic arsenic (As), a nonthreshold, class 1 human carcinogen. Rice grain As is dominated by the inorganic species, and the organic species dimethylarsinic acid (DMA). To investigate how As species are unloaded into grain rice, panicles were excised during grain filling and hydroponically pulsed with arsenite, arsenate, glutathione-complexed As, or DMA. Total As concentrations in flag leaf, grain, and husk, were quantified by inductively coupled plasma mass spectroscopy and As speciation in the fresh grain was determined by x-ray absorption near-edge spectroscopy. The roles of phloem and xylem transport were investigated by applying a ± stem-girdling treatment to a second set of panicles, limiting phloem transport to the grain in panicles pulsed with arsenite or DMA. The results demonstrate that DMA is translocated to the rice grain with over an order magnitude greater efficiency than inorganic species and is more mobile than arsenite in both the phloem and the xylem. Phloem transport accounted for 90% of arsenite, and 55% of DMA, transport to the grain. Synchrotron x-ray fluorescence mapping and fluorescence microtomography revealed marked differences in the pattern of As unloading into the grain between DMA and arsenite-challenged grain. Arsenite was retained in the ovular vascular trace and DMA dispersed throughout the external grain parts and into the endosperm. This study also demonstrates that DMA speciation is altered in planta, potentially through complexation with thiols.Paddy rice (Oryza sativa) is particularly effective, compared to other cereals, at accumulating arsenic (As) in shoot and grain (Williams et al., 2007b). Rice is the staple food for over half the world''s population (Fageria, 2007) and rice represents a significant dietary source of inorganic As, a class 1, nonthreshold carcinogen, particularly in Southeast Asia (Meharg et al., 2009). Inorganic As levels in rice grain are problematic even where soil As is at background levels, derived from geogenic sources (Lu et al., 2009; Meharg et al., 2009). However, widespread pollution of paddy soils with As, leading to further elevation of grain As, has occurred in some regions due to base and precious mining (Liao et al., 2005; Zhu et al., 2008), irrigation of paddies with As-elevated groundwaters (e.g. Meharg and Rahman, 2003; Williams et al., 2006), and the use of arsenical pesticides (Williams et al., 2007a). Unlike other cereal grains, paddy rice cultivation is dependent of soils being anaerobic, and it is this anoxia that gives rise to elevated As concentrations in the plant. Anaerobic soil conditions lead to the mobilization of As as arsenite, where under aerobic systems arsenate dominates (Xu et al., 2008). Arsenite is efficiently assimilated by rice roots through silicic acid transport pathway (Ma et al., 2008).Knowledge of As metabolism and partitioning within plants, particularly rice, is still developing rapidly (Zhao et al., 2009). Several studies have now shown that As in rice vegetative tissue and grain is predominantly speciated as inorganic As and the methylated species dimethylarsinic acid (DMA), with variable, though low, levels of monomethyl arsonic acid (MMA; Abedin et al., 2002a; Williams et al., 2005, 2006; Norton et al., 2009). Arsenate is an analog of phosphate and competes with phosphate for rice root uptake (Abedin et al., 2002a) while arsenite is taken up by rice roots via silicic acid transporters (Ma et al., 2008). Abedin et al. (2002b) demonstrated that the methylated species DMA and MMA are also taken up by rice plants although at a much slower rate than inorganic As, with the protonated neutral forms also transported through silicic acid pathway (Li et al., 2009). Arsenate is reduced to arsenite within the rice root (Xu et al., 2008; Zhao et al., 2009), which then enters the xylem via a silicic acid/arsenite effluxer (Ma et al., 2008; Zhao et al., 2009). Arsenite may be detoxified through complexation with thiol-rich peptides including phytochelatins (PCs) and glutathione followed by sequestration into vacuoles (Bleeker et al., 2006; Raab et al., 2007b; Zhao et al., 2009). Raab et al. (2007a) found that while methylated As species are taken up by rice roots much less efficiently than inorganic species, they appear to be translocated within the plant more efficiently. The comparative contributions of xylem and phloem transport, in translocation of As to the grain, are unknown.The main species within rice grain, along with DMA, are inorganic As, particularly arsenite, which may be complexed with thiols (Williams et al., 2005; Lombi et al., 2009). Nutrients are unloaded into the grain from the ovular vascular trace (OVT) into the nucellar tissue and from there are uploaded, via the apoplast into the filial tissue (the aleurone and the endosperm; Krishnan and Dayanandan, 2003). Lombi et al. (2009) recently suggested that this may represent a physiological barrier that As species cross with differential efficiency. However, the transport and unloading of As to/into the grain, which are key processes in terms of human exposure to this contaminant, are far from being fully understood.This study investigated the differential efficiency with which important As species are translocated and unloaded into the rice grain and the comparative contributions of phloem and xylem transport. Rice panicles were excised below the flag leaf node during grain development, 10 DPA, and treated to a hydroponically administered 48-h pulse of arsenite, arsenate, arsenite glutathione, or DMA. Total As concentrations in flag leaf, grain, and husk samples for each treatment were quantified by inductively coupled plasma mass spectroscopy (ICP-MS), and As speciation in the fresh grain was determined by x-ray absorption near-edge spectroscopy (XANES) analysis. To evaluate the contributions of phloem versus xylem transport, a stem-girdling treatment was applied, using steam to destroy phloem cells in a second set of panicles prior to a pulse of either DMA or arsenite. The spatial unloading of As species into the developing grain was examined by synchrotron x-ray fluorescence (XRF) mapping, and fluorescence microtomography for the DMA and arsenite treatments.     

Soil silicon amendments increase resistance of sugarcane to stalk borer Eldana saccharina Walker (Lepidoptera: Pyralidae) under field conditions

Background and aims

Soil amendment with silicon (Si) can significantly increase resistance of susceptible sugarcane cultivars grown in pots to stalk borer Eldana saccharina (Lepidoptera: Pyralidae). This study tested the hypothesis that a single application of silicate can increase resistance to E. saccharina and increase yield in field-grown sugarcane.

Methods

Two Si materials (Calmasil® and Slagment® at 4 and 8 t/ha) were applied at planting to a field trial extending over three successive crops and incorporating three sugarcane cultivars varying in borer susceptibility.

Results

Both materials, especially Slagment, significantly increased soil, leaf and stalk Si content, but leaf Si levels seldom exceeded 0.5 %. Silicon treatment significantly reduced percent stalks bored in all three crops and stalk length bored in the second ratoon crop, but did not affect borer numbers per 100 stalks (E/100) or increase cane or sucrose yield. Borer damage and E/100 were significantly and consistently reduced in the resistant cultivar.

Conclusions

We argue that if leaf Si% in field sugarcane can be elevated to or exceed 0.8 %, using materials that release Si slowly, substantial reductions in stalk damage and sucrose loss could be achieved in susceptible cultivars in low-Si soils.     

Silicon ameliorates manganese toxicity by regulating manganese transport and antioxidant reactions in rice (Oryza sativa L.)

Background and aims

This study aimed to investigate the roles of silicon (Si) in ameliorating manganese (Mn) toxicity in two rice (Oryza sativa L.) cultivars: i.e. cv. Xinxiangyou 640 (XXY), a Mn-sensitive cultivar and cv. Zhuliangyou 99 (ZLY), a Mn-tolerant cultivar.

Methods

Plants were cultured in nutrient solution containing normal Mn (6.7 μM) or high Mn (2.0 mM), both with or without Si supply at 1.5 mM Si.

Results

Plant growth was severely inhibited by high Mn in cv. XXY, but was enhanced by Si supply. In cv. XXY, Si-enhanced tolerance resulted from a restriction of Mn transport, whereas in cv. ZLY Mn uptake was depressed. In cv. XXY, high Mn significantly increased superoxide dismutase (SOD), catalase and ascorbate peroxidase activities but decreased non-protein thiols and glutathione concentrations, leading to accumulation of H₂O₂ and malondialdehyde. The addition of Si significantly counteracted high Mn-elevated malondialdehyde and H₂O₂ concentrations and enhanced plant growth. In cv. ZLY, high Mn considerably raised SOD activities and glutathione concentrations, thus leading to relatively low oxidative damage.

Conclusions

Si-enhanced Mn tolerance was attributed mainly to restricted Mn transport in cv. XXY but to depressed Mn uptake in cv. ZLY. Silicon mainly influenced non-enzymatic antioxidants in these two rice cultivars under high Mn stress.     

Influence of Sulfur on Transcription of Genes Involved in Arsenic Accumulation in Rice Grains

Arsenic (As) contamination of rice grains affects millions of people worldwide. In this study, we found that sulfur application (20As+120S) decreased As concentration in rice grains by 44 % compared to grains without sulfur application (20As+0S). Importantly, sulfur application decreased arsenate [As(V)] and arsenite [As(III)] concentration in rice grains significantly, while there was no significant effect on dimethylarsenate (DMA) concentration. To elucidate the molecular basis of As accumulation in rice grains, we performed Illumina sequencing to acquire the differentially expressed genes induced by arsenate and sulfur treatments. By contrast with the control, the expression of 1,000 genes was found to be changed significantly, with 46 genes up-regulated and 954 genes down-regulated in grains grown in arsenate-contaminated soil (20As+0S). Between samples of control and arsenate together with sulfur treatment (20As+120S), 1,169 genes expressed significantly differently, with 16 genes up-regulated and 1,153 genes down-regulated. Sulfur application significantly changed the expression of genes involved in As metabolism in rice grains, significantly down-regulated phosphate transporter gene OsPT23 and aquaporin gene OsTIP4;2, while ABC transporter genes (OsABCG5, OsABCI7_2 and OsABC6) and phytochelatin synthase genes (OsPCS1, OsPCS3 and OsPCS13) were up-regulated. These results provide an insight into the molecular basis of how sulfur assimilation regulates As accumulation in rice grains.     

Silencing of omega-5 gliadins in transgenic wheat eliminates a major source of environmental variability and improves dough mixing properties of flour

Background

Silicon (Si) application has been known to enhance the tolerance of plants against abiotic stresses. However, the protective mechanism of Si under heavy metals contamination is poorly understood. The aim of this study was to assess the role of Si in counteracting toxicity due to cadmium (Cd) and copper (Cu) in rice plants (Oryza sativa).

Results

Si significantly improved the growth and biomass of rice plants and reduced the toxic effects of Cd/Cu after different stress periods. Si treatment ameliorated root function and structure compared with non-treated rice plants, which suffered severe root damage. In the presence of Si, the Cd/Cu concentration was significantly lower in rice plants, and there was also a reduction in lipid peroxidation and fatty acid desaturation in plant tissues. The reduced uptake of metals in the roots modulated the signaling of phytohormones involved in responses to stress and host defense, such as abscisic acid, jasmonic acid, and salicylic acid. Furthermore, the low concentration of metals significantly down regulated the mRNA expression of enzymes encoding heavy metal transporters (OsHMA2 and OsHMA3) in Si-metal-treated rice plants. Genes responsible for Si transport (OsLSi1 and OsLSi2), showed a significant up-regulation of mRNA expression with Si treatment in rice plants.

Conclusion

The present study supports the active role of Si in the regulation of stresses from heavy metal exposure through changes in root morphology.     

Effect of selenium fertilization on the accumulation of cadmium and lead in rice plants

Background and Aims

The accumulation of cadmium and lead in rice (Oryza sativa L.) grains is a potential threat to human health. In this study, the effect of selenium fertilization on the uptake and translocation of cadmium and lead in rice plants was investigated.

Methods

Rice plants were cultivated using cadmium and lead contaminated soils with selenium addition at three concentrations (0, 0.5 and 1 mg kg^?1). At maturity, plants were harvested, and element concentrations in rice tissues were analyzed by using ICP-MS.

Results

Selenium application significantly increased selenium accumulation in rice grain, and markedly decreased cadmium and lead concentrations in rice tissues. In brown rice grains, selenium application reduced cadmium concentrations by 44.4 %, but had no significant effect on lead accumulation. Selenium application significantly decreased metal mobility in soils, at 0.5 mg kg^?1 treatment, the translocation factor of cadmium and lead from soil to iron plaque decreased by 71 and 33 % respectively.

Conclusions

The mechanism of selenium mitigating of heavy metal accumulation in rice could be decreasing metal bioavailability in soil. Selenium fertilization could be an effective and feasible method to enrich selenium and reduce cadmium levels in brown rice.     

Screening of plant growth-promoting traits in arsenic-resistant bacteria isolated from the rhizosphere of soybean plants from Argentinean agricultural soil

Aims

The purpose of this study was to investigate plant-growth promoting traits in native and arsenic (As) highly-resistant bacterial strains isolated from the rhizosphere of soybean (Glycine max) plants grown in an Argentinean agricultural field.

Methods

Determination of MICs (Minimum inhibitory concentration) was carried out on solid media supplemented with arsenite (As 3+) or arsenate (As 5+). Morphological, cultural, physiological, biochemical and molecular characterization, and in vitro determination of plant growth promoting (PGP) properties of As resistant isolates were carried out. Arsenic in soil samples was determined by ICP-OES while residual arsenic on post-removal culture medium and accumulation in cells were estimated by GF-AAS after wet acid digestion.

Results

Isolated strains included γ-proteobacteria such as Enterobacter sp. and Pseudomonas sp., and actinobacteria as Rhodococcus sp. All bacterial strains grew in presence of very high arsenite -over 24?mM- and arsenate –over 400?mM- concentrations. Pseudomonas sp. strains presented simultaneously several in vitro PGP traits, although Rhodococcus erythropolis AW3 did not display PGP traits. However, R. erythropolis AW3 was the most As resistant strain and removed and accumulated the greatest amounts of the metalloid.

Conclusion

The presence of As resistant and plant-growth promoting bacterial strains in the rhizosphere of Glycine max, in arsenic containing agricultural soil, suggest that they could potentially play an important role in plant-growth promotion in stressed conditions. These strains were able to remove and accumulate As from liquid media, thus they could be beneficial for sustainable crop production.     

Biochar from Miscanthus: a potential silicon fertilizer

Background and aims

Silicon (Si) is largely recognized to improve plant growth subjected to various biotic and abiotic stresses. As plants accumulate Si in the form of readily-soluble phytolith, we examine the possibility of using phytolith-rich biochar as a bio-available Si source for increasing the agronomical productivity of Si high-accumulator plants while augmenting soil fertility and C sequestration.

Methods

By adding three different biochars (Miscanthus x giganteus straws, coffee husks and woody material) at two different concentrations (1 % and 3 %; w/w) to soil samples, we investigated the effects on the soil respiration, the chemical characteristics and the kinetic release of bio-available Si (CaCl₂-extractable Si).

Results

Here we show that the biochar from Miscanthus straws was the most attractive amendment. Its incorporation at a 3 % rate improved the soil fertility parameters (pH and available cations) and combined the highest mean residence time of carbon (C) in soil (MRT?=?50 years) with the highest rate of release of bio-available Si. We attribute this result to the presence of phytoliths in this biochar, as revealed by SEM-EDS analysis.

Conclusions

Not only did the biochar from Miscanthus enhance both soil C sequestration and fertility, but the results of this study suggest that it can also be considered as a potential source of bio-available Si. Although our conclusions should be substantiated in the field, we suggest that Miscanthus biochar could be used as a potential source of bio-available silicon for the culture of such crop as Si-accumulator plants growing, for instance, in highly weathered tropical soils with low content in carbon, nutrients and bio-available Si.     

The Rice Aquaporin Lsi1 Mediates Uptake of Methylated Arsenic Species

Pentavalent methylated arsenic (As) species such as monomethylarsonic acid [MMA(V)] and dimethylarsinic acid [DMA(V)] are used as herbicides or pesticides, and can also be synthesized by soil microorganisms or algae through As methylation. The mechanism of MMA(V) and DMA(V) uptake remains unknown. Recent studies have shown that arsenite is taken up by rice (Oryza sativa) roots through two silicon transporters, Lsi1 (the aquaporin NIP2;1) and Lsi2 (an efflux carrier). Here we investigated whether these two transporters also mediate the uptake of MMA(V) and DMA(V). MMA(V) was partly reduced to trivalent MMA(III) in rice roots, but only MMA(V) was translocated to shoots. DMA(V) was stable in plants. The rice lsi1 mutant lost about 80% and 50% of the uptake capacity for MMA(V) and DMA(V), respectively, compared with the wild-type rice, whereas Lsi2 mutation had little effect. The short-term uptake kinetics of MMA(V) can be described by a Michaelis-Menten plus linear model, with the wild type having 3.5-fold higher V_max than the lsi1 mutant. The uptake kinetics of DMA(V) were linear with the slope being 2.8-fold higher in the wild type than the lsi1 mutant. Heterologous expression of Lsi1 in Xenopus laevis oocytes significantly increased the uptake of MMA(V) but not DMA(V), possibly because of a very limited uptake of the latter. Uptake of MMA(V) and DMA(V) by wild-type rice was increased as the pH of the medium decreased, consistent with an increasing proportion of the undissociated species. The results demonstrate that Lsi1 mediates the uptake of undissociated methylated As in rice roots.Arsenic (As) contamination affects millions of people worldwide, particularly in South Asia where As-contaminated groundwater has been extracted for drinking (Chakraborti et al., 2002; Nordstrom, 2002). Recent studies have shown that foods, especially rice (Oryza sativa), are an important source of inorganic As to populations dependent on a rice diet (Kile et al., 2007; Ohno et al., 2007; Mondal and Polya, 2008). Paddy rice is more efficient than other cereal crops in accumulating As (Williams et al., 2007). This is because anaerobic conditions in submerged paddy soils lead to mobilization of arsenite [As(III); Takahashi et al., 2004; Xu et al., 2008], which is then taken up by rice roots mainly through the highly efficient transport pathway for silicon (Si; Ma et al., 2008). The relatively high accumulation of As in rice is of concern, as it may pose a significant health risk (Zhu et al., 2008; Meharg et al., 2009).A number of As species may be present in soil depending on soil conditions and the history of As contamination. These include arsenate [As(V)], As(III), and methylated As species such as monomethylarsonic acid [MMA(V): CH₃AsO(OH)₂] and dimethylarsinic acid [DMA(V): (CH₃)₂AsO(OH)]. As(V) is the main species in aerobic soils, while As(III) dominates in anaerobic environments such as flooded paddy soils. Both MMA(V) and DMA(V) have been found in paddy soils (Takamatsu et al., 1982), which may have been derived from microbial and algal biomethylation and/or past uses of methylated As compounds. MMA(V), as sodium or calcium salt, and DMA(V), as sodium salt or free acid (also called cacodylic acid), are herbicides widely used for weed control on cotton (Gossypium hirsutum), orchards, and lawns, or as a defoliant of cotton (U.S. Environmental Protection Agency, 2006). Conversion of cotton fields for the production of paddy rice in the United States may be a reason for the high levels of methylated As reported for the U.S. rice (Meharg et al., 2009).The mechanism of As(V) uptake by plants through the phosphate transport system has been well established (for review, see Zhao et al., 2009). In contrast, As(III) is taken up into the cells by aquaglyceroporins in Escherichia coli, yeast (Saccharomyces cerevisiae), and mammalian tissues (for review, see Bhattacharjee and Rosen, 2007). Recent studies have shown that several plant aquaporin channels belonging to the Nodulin 26-like Intrinsic Protein (NIP) subfamily are permeable to As(III) when expressed heterologously in yeast (Bienert et al., 2008; Isayenkov and Maathuis, 2008; Ma et al., 2008). The rice Si transporter Lsi1 (OsNIP2;1; Ma et al., 2006) is also permeable to As(III) when expressed in yeast or Xenopus laevis oocytes (Ma et al., 2008). Furthermore, the lsi1 rice mutant lost 57% of the influx capacity for As(III) compared to the wild type in short-term assays, suggesting that Lsi1 is an important entry route for As(III) (Ma et al., 2008). In rice roots, a second Si transporter, Lsi2, functions as an efflux carrier to transport Si efflux from the exodermis and endodermis cells toward the stele for xylem loading (Ma et al., 2007). This transporter also mediates As(III) efflux; two independent lsi2 mutants had 73% to 91% lower concentrations of As(III) in the xylem sap than their wild types (Ma et al., 2008). The shared uptake pathway between Si (silicic acid) and As(III) (arsenous acid) is consistent with their physiochemical properties; both are present predominantly as undissociated neutral molecules at the normal environmental and physiological pH range (pK_a = 9.2, >99% undissociated at pH ≤ 7.0), and the two molecules have similar sizes.The uptake mechanisms of methylated As species by plant roots are not known. Previous studies showed that both MMA(V) and DMA(V) can be taken up by roots and translocated to shoots in a number of plant species (Marin et al., 1992; Carbonell-Barrachina et al., 1998, 1999; Burló et al., 1999). Marin et al. (1992) found that uptake by rice followed the order of As(III) > MMA(V) > As(V) > DMA (V), although DMA(V) was more efficiently translocated from roots to shoots than other As species. Raab et al. (2007) reported large variations in the absorption and translocation efficiencies for As(V), MMA(V), and DMA(V) among the 46 plant species tested. On average, root absorption of As(V) was 2.5- and 5-times higher than MMA(V) and DMA(V), respectively. The translocation efficiency, defined as the shoot-to-root concentration ratio after 24-h exposure, was highest for DMA(V) (0.8), followed by MMA(V) (0.3) and As(V) (0.09). The concentration-dependent uptake kinetics of MMA(V) in rice roots could be described by the Michaelis-Menten equation, whereas the limited uptake of DMA(V) appeared to be linear in relation to the increasing concentration in the uptake medium (Abedin et al., 2002). Abbas and Meharg (2008) showed that DMA(V) uptake by maize (Zea mays) seedlings was enhanced by more than 10-fold by a pretreatment of phosphorus starvation; this compared with only 2-fold increase in As(V) uptake. They thought that DMA(V) might be taken up by the phosphate transporters, or that phosphorus starvation altered expression of a range of membrane transporters or even membrane permeability itself.In addition to the root uptake of methylated As species, some plants appear to be able to biomethylate As, but the pathway and enzymology remains unclear (Wu et al., 2002; Zhao et al., 2009). In microbes, As methylation follows the Challenger pathway involving repeated steps of As reduction and oxidative methylation (Bentley and Chasteen, 2002). As(V) is first reduced to As(III), which is methylated by S-adenosylmethyltransferase using S-adenosyl-l-Met as the methyl donor. The product of this reaction is pentavalent MMA(V), which is reduced by a reductase to trivalent MMA(III) with thiols (e.g. glutathione). Methylation and reduction steps continue to produce di- and trimethyl As compounds. MMA(III) and DMA(III) are intermediates in the As methylation pathway, which is not very stable (Gong et al., 2001). In rice grain, DMA(V) is the main form of methylated As, and can account for up to 80% of the total As (Zavala et al., 2008; Meharg et al., 2009). In light of the significant presence of methylated As in rice, it is important to elucidate the transport and assimilation pathways of these As species in plants.In this study, we present evidence that MMA(V) and DMA(V) are taken up by rice roots, at least partly, through the NIP aquaporin channel Lsi1, and that this process is strongly pH dependent. We also show that MMA(V) can be reduced to MMA(III) in planta.     

Growth and phosphorus nutrition of rice when inorganic fertiliser application is partly replaced by straw under varying moisture availability in sandy and clay soils

Background and aims

The combined effects of (1) reduced soil moisture availability, (2) reduced application of inorganic fertilisers while incorporating straw, (3) soil type, and their effects on growth, root system plasticity, phosphorus (P) nutrition of rice, and soil P dynamics are poorly known, but very important when aiming to increase the efficiency of water and P use.

Methods

Using large pots a three-factor factorial experiment was conducted with two moisture treatments (i.e. continuous flooding, and draining of top soil after flowering while subsoil was kept moist through capillary action), three fertilisation treatments; with (P1) and without (P0) applications of inorganic P fertilisers, and 25 % of inorganic fertilisers reduced while incorporating straw (5 t ha^?1), and soil type (i.e. clay and sandy soils with 15 and 9 mg P kg^?1 soil, respectively in P0). Shoot and root growth, root system plasticity, P nutrient status and soil P dynamics were measured.

Key results

Straw incorporation with reduced inorganic fertiliser application ensured a higher shoot dry weight and yield only in flooded clay soil as compared with P0 and P1, and a similar shoot dry weight and yield to P1 under drained clay soil. A positive growth response was facilitated by an increased water-use efficiency and rate of photosynthesis in shoots, and increased root system plasticity through the production of greater root length, more roots in deep soil layers, and an increased fraction of fine roots. Straw enhanced P extractability in soil. Drained soil reduced P uptake (15–45 %) and increased P-use efficiency. In addition to the re-translocation of P from senescing leaves and stems under both moisture conditions, the P concentration in green leaves under drained condition was also reduced (41–72 %).

Conclusion

Growth benefits of straw incorporation were observed in clay soil under both moisture conditions, and this was facilitated by the improved P availability, increased P uptake, and greater root system plasticity with the production of deeper and finer roots, compared with that in sandy soil, and inorganic fertiliser applications alone. As P uptake was reduced under drained soil, P re-translocation and % P allocated to panicles increased.     

Zinc nutrition in rice production systems: a review

Background

Zinc (Zn) deficiency is one of the important abiotic factors limiting rice productivity worldwide and also a widespread nutritional disorder affecting human health. Given that rice is a staple for populations in many countries, studies of Zn dynamics and management in rice soils is of great importance.

Scope

Changing climate is forcing the growers to switch from conventional rice transplanting in flooded soils to water-saving cultivation, including aerobic rice culture and alternate wetting and drying system. As soil properties are changed with altered soil and water management, which is likely to affect Zn solubility and plant availability and should be considered before Zn management in rice. In this review, we critically appraise the role of Zn in plant biology and its dynamics in soil and rice production systems. Strategies and options to improve Zn uptake and partitioning efficiency in rice by using agronomic, breeding and biotechnological tools are also discussed.

Conclusions

Although soil application of inorganic Zn fertilizers is widely used, organic and chelated sources are better from economic and environmental perspectives. Use of other methods of Zn application (such as seed treatment, foliar application etc., in association with mycorrhizal fungi) may improve Zn-use efficiency in rice. Conventional breeding together with modern genomic and biotechnological tools may result in development of Zn-efficient rice genotypes that should be used in conjunction with judicious fertilization to optimize rice yield and grain Zn content.     

Integrated effects of organic,inorganic and biological amendments on methane emission,soil quality and rice productivity in irrigated paddy ecosystem of Bangladesh: field study of two consecutive rice growing seasons

Aims

Effects of different soil amendments were investigated on methane (CH₄) emission, soil quality parameters and rice productivity in irrigated paddy field of Bangladesh.

Methods

The experiment was laid out in a randomized complete block design with five treatments and three replications. The experimental treatments were urea (220 kg ha^?1) + rice straw compost (2 t ha^?1) as a control, urea (170 kg ha^?1) + rice straw compost (2 t ha^?1) + silicate fertilizer, urea (170 kg ha^?1) + sesbania biomass (2 t ha^?1 ) + silicate fertilizer, urea (170 kg ha^?1) + azolla biomass (2 t ha^?1) + cyanobacterial mixture 15 kg ha^?1 silicate fertilizer, urea (170 kg ha^?1) + cattle manure compost (2 t ha^?1) + silicate fertilizer.

Results

The average of two growing seasons CH₄ flux 132 kg ha^?1 was recorded from the conventional urea (220 kg ha^?1) with rice straw compost incorporated field plot followed by 126.7 (4 % reduction), 130.7 (1.5 % reduction), 116 (12 % reduction) and 126 (5 % reduction) kg CH₄ flux ha^?1 respectively, with rice straw compost, sesbania biomass, azolla anabaena and cattle manure compost in combination urea and silicate fertilizer applied plots. Rice grain yield was increased by 15 % and 10 % over the control (4.95 Mg ha^?1) with silicate plus composted cattle manure and silicate plus azolla anabaena, respectively. Soil quality parameters such as soil organic carbon, total nitrogen, microbial biomass carbon, soil redox status and cations exchange capacity were improved with the added organic materials and azolla biofertilizer amendments with silicate slag and optimum urea application (170 kg ha^?1) in paddy field.

Conclusion

Integrated application of silicate fertilizer, well composted organic manures and azolla biofertilizer could be an effective strategy to minimize the use of conventional urea fertilizer, reducing CH₄ emissions, improving soil quality parameters and increasing rice productivity in subtropical countries like Bangladesh.     

Root-induced changes in radial oxygen loss, rhizosphere oxygen profile, and nitrification of two rice cultivars in Chinese red soil regions

Aims

To evaluate the external and internal morphological differences of roots that might influence rice root radial oxygen loss (ROL) and the corresponding rhizosphere nitrification activity, growth characteristics and nitrogen nutrition of rice.

Methods

The root ROL and rhizosphere oxygen profile were determined using a miniaturised Clark-type oxygen microelectrode system, and the rhizosphere nitrification activity was studied with a short-term nitrification activity assay.

Results

The rice biomass, nitrogen accumulation and nitrogen use efficiency (NUE) of ZH (high yield) were significantly higher than those of HS (low yield). The root biomass, number, diameter and porosity of ZH were also much greater than those of HS. The inner and surface oxygen concentrations of the root of ZH were significantly higher than those of HS. The order of paddy soil oxygen penetration depth was ZH?>?HS?>?CK, and the order of the oxygen concentrations detected in the water layer and rhizosphere soil was the same. The rhizosphere nitrification activity and nitrate concentration of ZH were significantly higher than those of HS.

Conclusions

More porous and thicker roots improved the individual root ROL, and more adventitious root numbers enhanced the entire plant ROL and correspondingly improved the rhizosphere nitrification activity, which might influence the growth and nitrogen nutrition of rice.     

Comparison of As sequestration in iron plaque and uptake by different genotypes of rice plants grown in As-contaminated paddy soils

Background and aims

Limited information is available on comparing the iron plaque formation capabilities and their effect on arsenic (As) uptake by different rice plant genotypes grown in As-contaminated soils. This study investigates the effect of iron plaque on As uptake in different rice genotypes grown in As-contaminated soils from the Guandu Plain of northern Taiwan.

Methods

Twenty-eight rice genotypes including 14 japonica and 14 indica genotypes were used in this study. Rice seedlings were grown in As-contaminated soils for 38 days. The iron plaque formed on the rice roots were extracted using dithionite–citrate–bicarbonate. The concentrations of As, Fe, and P in soil solutions, iron plaque, and plants were measured. The speciation of As in the root’s iron plaque was determined by As K-edge X-ray absorption near-edge structure spectroscopy (XANES).

Results

The amounts of iron plaque formation on roots were significantly different among 28 tested rice genotypes, and 75.7–92.8 % of As uptake from soils could be sequestered in iron plaque. However, there were no significant negative correlations between the amounts of Fe or As in the iron plaque and the content of As accumulated in rice plants of tested genotypes. XANES data showed that arsenate was the predominant As species in iron plaque, and there were difference in the distribution of As species among different rice genotypes.

Conclusions

The iron plaque can sequester most of As uptake from soils no matter what rice genotypes used in this study. However, the iron plaque alone did not control the extent of As accumulation in rice plants from As-contaminated soils among 28 tested rice genotypes. Low As uptake genotypes of rice selected from this study can be recommended to be grown in the As-contaminated soils.     

Biogeochemical cycling of soil phosphorus during natural revegetation of Pinus sylvestris on disused sand quarries in Northwestern Russia

Background and aims

Quarrying causes severe degradation of soils and vegetation that can be recovered partially when the quarries are abandoned and re-colonised by plants. To understand the recovery of soil functionality and nutrient cycling, we studied the development of soil phosphorus pools during Scots pine (Pinus sylvestris) revegetation in a disused sand quarry in Northwestern Russia.

Methods

Sites that had been developing for different times since abandonment were compared to the parent sand and an adjacent undisturbed forest. Phosphorus speciation in genetic horizons of soil profiles was determined by sequential fractionation and solution phosphorus-31 nuclear magnetic resonance spectroscopy.

Results

Rapid transformations in soil properties occurred in 40 years, with a marked decline in pH and an accumulation of organic matter. Phosphorus transformations were shaped by geochemical processes, with a rapid release of inorganic phosphorus from primary minerals and accumulation of organic phosphorus to concentrations exceeding those found in the undisturbed site. Adsorbed and/or precipitated phosphorus increased rapidly, despite few reactive mineral colloidal surfaces.

Conclusions

Natural succession of Scots pine in post-mining landscapes promotes ecosystem restoration through the rapid re-establishment of the biogeochemical cycles of organic matter and phosphorus. This study provides an important example of biogeochemical phosphorus cycling during the initial stages of pedogenesis.     

Differential responses of system of rice intensification (SRI) and conventional flooded-rice management methods to applications of nitrogen fertilizer

Background

Rising food demand, slowing productivity growth, poor N-use efficiency in rice, and environmental degradation necessitate the development of more productive, environmentally-sound crop and soil management practices. The system of rice intensification (SRI) has been proposed as a methodology to address these trends. However, it is not known how its modified crop-soil-water management practices affect efficiency of inorganic nitrogen applications.

Methods

Field experiments investigated the impacts of SRI management practices with different N-application rates on grain yield, root growth and activity, uptake of N and its use-efficiency, leaf chlorophyll content, leaf N-concentration, and photosynthetic rate in comparison with standard management practices for transplanted flooded rice (TFR).

Results

Overall, grain yield with SRI was 49 % higher than with TFR, with yield enhanced at every N application dose. N-uptake, use-efficiency, and partial factor productivity from applied N were significantly higher in SRI than TFR. Higher leaf nitrogen and chlorophyll contents during the ripening-stage in SRI plants reflected delayed leaf-senescence, extension of photosynthetic processes, and improved root-shoot activities contributing to increased grain yield.

Conclusions

Rice grown under SRI management used N fertilizer more efficiently due to profuse root development and improved physiological performance resulting in enhanced grain yield compared to traditional flooded rice.