The effects of cerium oxide (CeO 2 ) and indium oxide (In 2 O 3 ) nanoparticles (NPs) exposure on Arabidopsis thaliana (L.) Heynh. were investigated. After inoculation in half strength MS medium amended with 0−2000 ppm CeO 2 and In 2 O 3 NPs for 25 days, both physiological and molecular responses were evaluated. Exposure at 250 ppm CeO 2 NPs significantly increased plant biomass, but at 500−2000 ppm, plant growth was decreased by up to 85% in a dose-dependent fashion. At 1000 and 2000 ppm CeO 2 NPs, chlorophyll production was reduced by nearly 60% and 85%, respectively, and anthocyanin production was increased 3−5-fold. Malondialdehyde (MDA) production, a measure of lipid peroxidation, was unaffected by exposure to 250−500 ppm CeO 2 NPs, but at 1000 ppm, MDA formation was increased by 2.5-fold. Exposure to 25−2000 ppm In 2 O 3 NPs had no effect on A. thaliana biomass and only minor effects (15%) on root elongation. Total chlorophyll and MDA production were unaffected by In 2 O 3 NPs exposure. Molecular response to NP exposure as measured by qPCR showed that both types of elements altered the expression of genes central to the stress response such as the sulfur assimilation and glutathione (GSH) biosynthesis pathway, a series of genes known to be significant in the detoxification of metal toxicity in plants. Interestingly, In 2 O 3 NPs exposure resulted in a 3.8−4.6-fold increase in glutathione synthase (GS) transcript production, whereas CeO 2 NPs yielded only a 2-fold increase. It seems likely that the significantly greater gene regulation response upon In 2 O 3 NPs exposure was directly related to the decreased phytotoxicity relative to CeO 2 treatment. The use of NP rare earth oxide elements has increased dramatically, yet knowledge on fate and toxicity has lagged behind. To our knowledge, this is the first report evaluating both physiological and molecular plant response from exposure to these important nanoparticles. KEYWORDS: Arabidopsis thaliana, CeO 2 and In 2 O 3 NPs, Stress response, Gene expression, Anthocyanin, Lipid peroxidation, Sulfur assimilation pathway coli, Bacillus subtilis, and Pseudomonas f luorescens), 5,6 plants (Cucurbita pepo L., Solanum lycopersicum L., and Zea mays L.) 7−9 and animals (Zebrafish: Danio rerio). 10,11 Moreover, the potential hazards of nanoparticles to human health have been discussed in assays using human cells. 4,12 Given these findings, it is clear that a full and mechanistic understanding of the risks Special Issue: Sustainable Nanotechnology
Plants detoxify toxic metals through a GSH-dependent pathway. GSH homeostasis is maintained by the g-glutamyl cycle, which involves GSH synthesis and degradation and the recycling of component amino acids. The enzyme g-glutamyl cyclotransferase (GGCT) is involved in Glu recycling, but the gene(s) encoding GGCT has not been identified in plants. Here, we report that an Arabidopsis thaliana protein with a cation transport regulator-like domain, hereafter referred to as GGCT2;1, functions as g-glutamyl cyclotransferase. Heterologous expression of GGCT2;1 in Saccharomyces cerevisiae produced phenotypes that were consistent with decreased GSH content attributable to either GSH degradation or the diversion of g-glutamyl peptides to produce 5-oxoproline (5-OP). 5-OP levels were further increased by the addition of arsenite and GSH to the medium, indicating that GGCT2;1 participates in the cellular response to arsenic (As) via GSH degradation. Recombinant GGCT2;1 converted both GSH and g-glutamyl Ala to 5-OP in vitro. GGCT2;1 transcripts were upregulated in As-treated Arabidopsis, and ggct2;1 knockout mutants were more tolerant to As and cadmium than the wild type. Overexpression of GGCT2;1 in Arabidopsis resulted in the accumulation of 5-OP. Under As toxicity, the overexpression lines showed minimal changes in de novo Glu synthesis, while the ggct2;1 mutant increased nitrogen assimilation by severalfold, resulting in a very low As/N ratio in tissue. Thus, our results suggest that GGCT2;1 ensures sufficient GSH turnover during abiotic stress by recycling Glu.
Rice accumulates high level of arsenic (As) in its edible parts and thus plays an important role in the transfer of As into the food chain. However, the mechanisms of As uptake and its detoxification in rice are not well understood. Recently, members of the Nodulin 26-like intrinsic protein (NIP) subfamily of plant aquaporins were shown to transport arsenite in rice and Arabidopsis. Here we report that members of the rice plasma membrane intrinsic protein (PIP) subfamily are also involved in As tolerance and transport. Based on the homology search with the mammalian AQP9 and yeast Fps1 arsenite transporters, we identified and cloned five rice PIP gene subfamily members. qRT-PCR analysis of PIPs in rice root and shoot tissues revealed a significant down regulation of transcripts encoding OsPIP1;2, OsPIP1;3, OsPIP2;4, OsPIP2;6, and OsPIP2;7 in response to arsenite treatment. Heterologous expression of OsPIP2;4, OsPIP2;6, and OsPIP2;7 in Xenopus laevis oocytes significantly increased the uptake of arsenite. Overexpression of OsPIP2;4, OsPIP2;6, and OsPIP2;7 in Arabidopsis yielded enhanced arsenite tolerance and higher biomass accumulation. Further, these transgenic plants showed no significant accumulation of As in shoot and root tissues in long term uptake assays. Whereas, short duration exposure to arsenite caused both active influx and efflux of As in the roots. The data suggests a bidirectional arsenite permeability of rice PIPs in plants. These rice PIPs genes will be highly useful for engineering important food and biofuel crops for enhanced crop productivity on contaminated soils without increasing the accumulation of toxic As in the biomass or edible tissues.
Boron (B) toxicity is responsible for low cereal crop production in a number of regions worldwide. In this report, we characterized two rice genes, OsPIP2;4 and OsPIP2;7, for their involvement in B permeability and tolerance. Transcript analysis demonstrated that the expression of OsPIP2;4 and OsPIP2;7 were downregulated in shoots and strongly upregulated in rice roots by high B treatment. Expression of both OsPIP2;4 and OsPIP2;7 in yeast HD9 strain lacking Fps1, ACR3, and Ycf1 resulted in an increased B sensitivity. Furthermore, yeast HD9 strain expressing OsPIP2;4 and OsPIP2;7 accumulated significantly higher B as compared to empty vector control, which suggests their involvement in B transport. Overexpression of OsPIP2;4 and OsPIP2;7 in Arabidopsis imparted higher tolerance under B toxicity. Arabidopsis lines overexpressing OsPIP2;4 and OsPIP2;7 showed significantly higher biomass production and greater root length, however there was no difference in B accumulation in long term uptake assay. Short-term uptake assay using tracer B (¹⁰B) in shoots and roots demonstrated increased ¹⁰B accumulation in Arabidopsis lines expressing OsPIP2;4 and OsPIP2;7, compare to wild type control plants. Efflux assay of B in the roots showed that ¹⁰B was effluxed from the Arabidopsis transgenic plants overexpressing OsPIP2;4 or OsPIP2;7 during the initial 1-h of assay. These data indicate that OsPIP2;4 and OsPIP2;7 are involved in mediating B transport in rice and provide tolerance via efflux of excess B from roots and shoot tissues. These genes will be highly useful in developing B tolerant crops for enhanced yield in the areas affected by high B toxicity.
High boron (B) concentration is toxic to plants that limit plant productivity. Recent studies have shown the involvement of the members of major intrinsic protein (MIP) family in controlling B transport. Here, we have provided experimental evidences showing the bidirectional transport activity of rice OsPIP1;3 and OsPIP2;6. Boron transport ability of OsPIP1;3 and OsPIP2;6 were displayed in yeast HD9 mutant strain (∆fps1∆acr3∆ycf1) as a result of increased B sensitivity, influx and accumulation by OsPIP1;3, and rapid efflux activity by OsPIP2;6. RT-PCR analysis showed strong upregulation of OsPIP1;3 and OsPIP2;6 transcripts in roots by B toxicity. Transgenic Arabidopsis lines overexpressing OsPIP1;3 and OsPIP2;6 exhibited enhanced tolerance to B toxicity. Furthermore, B concentration was significantly increased after 2 and 3 hours of tracer boron (10B) treatment. Interestingly, a rapid efflux of 10B from the roots of the transgenic plants was observed within 1 h of 10B treatment. Boron tolerance in OsPIP1;3 and OsPIP2;6 lines was inhibited by aquaporin inhibitors, silver nitrate and sodium azide. Our data proved that OsPIP1;3 and OsPIP2;6 are indeed involved in both influx and efflux of boron transport. Manipulation of these PIPs could be highly useful in improving B tolerance in crops grown in high B containing soils.
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