SummarySilicon (Si) is not considered to be an essential element for higher plants and is believed to have no effect on primary metabolism in unstressed plants. In rice (Oryza sativa), Si nutrition improves grain production; however, no attempt has been made to elucidate the physiological mechanisms underlying such responses.Here, we assessed crop yield and combined advanced gas exchange analysis with carbon isotope labelling and metabolic profiling to measure the effects of Si nutrition on rice photosynthesis, together with the associated metabolic changes, by comparing wild-type rice with the low-Si rice mutant lsi1 under unstressed conditions.Si improved the harvest index, paralleling an increase in nitrogen use efficiency. Higher crop yields associated with Si nutrition exerted a feed-forward effect on photosynthesis which was fundamentally associated with increased mesophyll conductance. By contrast, Si nutrition did not affect photosynthetic gas exchange during the vegetative growth phase or in de-grained plants. In addition, Si nutrition altered primary metabolism by stimulating amino acid remobilization.Our results indicate a stimulation of the source capacity, coupled with increased sink demand, in Si-treated plants; therefore, we identify Si nutrition as an important target in attempts to improve the agronomic yield of rice.

Silicon (Si) has been recognized as a beneficial element to improve rice (Oryza sativa L.) grain yield. Despite some evidence suggesting that this positive effect is observed when Si is supplied along the reproductive growth stage (from panicle initiation to heading), it remains unclear whether its supplementation during distinct growth phases can differentially impact physiological aspects of rice and its yield and the underlying mechanisms. Here, we investigated the effects of additions/removals of Si at different growth stages and their impacts on rice yield components, photosynthetic performance, and expression of genes (Lsi1, Lsi2 and Lsi6) involved in Si distribution within rice shoots. Positive effects of Si on rice production and photosynthesis were manifested when it was specifically supplied during the reproductive growth stage, as demonstrated by: (1) a high crop yield associated with higher grain number and higher 1000-grain weight, whereas the leaf area and whole-plant biomass remained unchanged; (2) an increased sink strength which, in turn, exerted a feed-forward effect on photosynthesis that was coupled with increases in both stomatal conductance and biochemical capacity to fix CO; (3) higher Si amounts in the developing panicles (and grain husks) in good agreement with a remarkable up-regulation of Lsi6 (and to a lesser extent Lsi1). We suggest that proper levels of Si in these reproductive structures seem to play an as yet unidentified role culminating with higher grain number and size.

Silicon (Si) plays important roles in alleviating various abiotic stresses. In rice (Oryza sativa), arsenic (As) is believed to share the Si transport pathway for entry into roots, and Si has been demonstrated to decrease As concentrations. However, the physiological mechanisms through which Si might alleviate As toxicity in plants remain poorly elucidated. We combined detailed gas exchange measurements with chlorophyll fluorescence analysis to examine the effects of Si nutrition on photosynthetic performance in rice plants [a wild-type (WT) cultivar and its lsi1 mutant defective in Si uptake] challenged with As (arsenite). As treatment impaired carbon fixation (particularly in the WT genotype) that was unrelated to photochemical or biochemical limitations but, rather, was largely associated with decreased leaf conductance at the stomata and mesophyll levels. Indeed, regardless of the genotypes, in the plants challenged with As, photosynthetic rates correlated strongly with both stomatal (r(2)  = 0.90) and mesophyll (r(2)  = 0.95) conductances, and these conductances were, in turn, linearly correlated with each other. The As-related impairments to carbon fixation could be considerably reverted by Si in a time- and genotype-dependent manner. In conclusion, we identified Si nutrition as an important target in an attempt to not only decrease As concentrations but also to ameliorate the photosynthetic performance of rice plants challenged with As.

Malate is a central metabolite involved in a multiplicity of plant metabolic pathways, being associated with mitochondrial metabolism and playing significant roles in stomatal movements. Vacuolar malate transport has been characterized at the molecular level and is performed by at least one carrier protein and two channels in Arabidopsis () vacuoles. The absence of the Arabidopsis tonoplast Dicarboxylate Transporter (tDT) in the knockout mutant was associated previously with an impaired accumulation of malate and fumarate in leaves. Here, we investigated the consequences of this lower accumulation on stomatal behavior and photosynthetic capacity as well as its putative metabolic impacts. Neither the stomatal conductance nor the kinetic responses to dark, light, or high CO were highly affected in plants. In addition, we did not observe any impact on stomatal aperture following incubation with abscisic acid, malate, or citrate. Furthermore, an effect on photosynthetic capacity was not observed in the mutant lines. However, leaf mitochondrial metabolism was affected in the plants. Levels of the intermediates of the tricarboxylic acid cycle were altered, and increases in both light and dark respiration were observed. We conclude that manipulation of the tonoplastic organic acid transporter impacted mitochondrial metabolism, while the overall stomatal and photosynthetic capacity were unaffected.

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