Mg deficiency induces photo-oxidative stress primarily by limiting CO2 assimilation and not by limiting photosynthetic light utilization.

Photosynthetic processes within chloroplasts require substantial amounts of magnesium (Mg). It is suggested that the minimum Mg concentration for yield and dry matter (DM) formation is 1.5 mg g-1 DM. Yet, it was never clarified whether this amount is required for photosynthetic processes as well. The aim of this study was to determine how varying Mg concentrations affect the photosynthetic efficiency and photoprotective responses. Barley (Hordeum vulgare L.) was grown under four different Mg supplies (1, 0.05, 0.025 and 0.015 mM Mg) for 21 days to investigate the photosynthetic and photoprotective responses to Mg deficiency. Leaf Mg concentrations, CO2 assimilation, photosystem II efficiency, electron transport rate, photochemical and non-photochemical quenching, expression of reactive oxygen species (ROS) scavengers, and the pigment composition were analyzed. Our data indicate that CO2 assimilation is more sensitive to the reduction of tissue Mg concentrations than photosynthetic light reactions. Moreover, supply with the two lowest Mg concentrations induced photo-oxidative stress, as could be derived from increased expression of ROS scavengers and an increased pool size of the xanthophyll cycle pigments. We hypothesize, that the reduction of CO2 assimilation is a critical determinant for the increase of photo-oxidative stress under Mg deficiency.

Photosynthetic Cost Associated With Induced Defense to Plasmopara viticola in Grapevine.

Downy mildew caused by Plasmopara viticola is one of the most destructive diseases of Vitis vinifera worldwide. Grapevine breeding programs have introgressed P. viticola-resistant traits into cultivated V. vinifera genotypes and launched interspecific hybrids with resistance against downy mildew. In general, pathogen infection affects primary metabolism, reduces plant growth and development and modifies the secondary metabolism toward defense responses, which are costly in terms of carbon production and utilization. The objective of this work was to evaluate the photosynthesis impairment by inducible defenses at the leaf level in V. vinifera cultivars resistant to P. viticola. Photosynthetic limitations imposed by P. viticola in susceptible and resistant grapevine cultivars were evaluated. Histochemical localization of hydrogen peroxide and superoxide and the activity of ascorbate peroxidase were assessed. Measurements of leaf gas exchange, chlorophyll fluorescence and the response of leaf CO2 assimilation to increasing air CO2 concentrations were taken, and photosynthetic limitations determined in cultivars Solaris (resistant) and Riesling (susceptible). The net photosynthetic rates were reduced (-25%) in inoculated Solaris plants even before the appearance of cell death-like hypersensitive reactions ("HR"). One day after "HR" visualization, the net photosynthetic rate of Solaris was reduced by 57% compared with healthy plants. A similar pattern was noticed in resistant Cabernet Blanc and Phoenix plants. While the susceptible cultivars did not show any variation in leaf gas exchange before the appearance of visual symptoms, drastic reductions in net photosynthetic rate and stomatal conductance were found in diseased plants 12 days after inoculation. Decreases in the maximum Rubisco carboxylation rate and photochemical impairment were noticed in Riesling after inoculation with P. viticola, which were not found in Solaris. Damage to the photochemical reactions of photosynthesis was likely associated with the oxidative burst found in resistant cultivars within the first 24 h after inoculation. Both chlorophyll degradation and stomatal closure were also noticed in the incompatible interaction. Taken together, our data clearly revealed that the defense response against P. viticola causes a photosynthetic cost to grapevines, which is not reversible even 12 days after the pathogen infection.

Diagnosis of Nitrogen Nutrition in Rice Leaves Influenced by Potassium Levels.

Evaluation of nitrogen (N) status by leaf color is a kind of classic nutritional diagnostic method. However, the color of leaves is influenced not only by N, but also by other nutrients such as potassium (K). Two-year field trials with a factorial combination of N and K were conducted to investigate the effects of different N and K rates on soil plant analysis development (SPAD) readings and leaf N, K, magnesium (Mg), and chlorophyll concentrations. Visual inspections in leaf greenness revealed darker green leaves with increasing N rates, while paler green leaves with increasing K rates. Data showed that SPAD readings, chlorophyll, N and Mg concentrations, and the chloroplast area increased significantly with raising N rates, while declined sharply with the increase in K rates due to the antagonistic relationships between K+ and NH4 + as well as Mg2+. It was also probable that the increase in K promoted the growth of leaves and diluted their N and Mg concentrations. The paler leaf appearance resulting from the application of K may overestimate the actual demand for N in the diagnosis of rice N status. The strong antagonistic relationships between K+, NH4 +, and Mg2+ should be considered in rice production and fertilization.

Minimum magnesium concentrations for photosynthetic efficiency in wheat and sunflower seedlings.

Photosynthetic processes in the chloroplast depend on the abundance of magnesium (Mg) in relatively high amounts; hence chloroplasts might react more sensitive to Mg-deficiency than other physiological processes within other organelles. Most authors suggest a critical Mg concentration to be 1.5 mg g-1 DM for biomass and yield formation. However, it is not yet elucidated whether this value also applies to photosynthetic processes. The present study focused on the response of photosynthetic processes to different Mg tissue concentrations. Wheat (Triticum aestivum) and sunflower (Helianthus annuus) plants were grown hydroponically for 10 days with 8 different levels of Mg supply (1.0, 0.5, 0.25, 0.1, 0.075, 0.05, 0.025, 0.01 mM Mg). Specific leaf mass, SPAD values, assimilation rate, Fv/Fm, electron transport rate and photochemical and non-photochemical quenching parameters were determined on youngest mature leaves. Tissue Mg concentrations decreased with lowering Mg supply to lowest concentrations of 0.7 mg g-1 DM in wheat leaves, but photosynthetic capacity was not affected. In sunflower leaves, lowest Mg concentrations of 0.56 mg g-1 DM were achieved and a diminished photosynthetic capacity was observed. The study shows that a Mg tissue concentration of 1.5 mg g-1 DM did not induce a negative effect on the photosynthetic capacity of wheat and sunflower leaves under our experimental conditions and hence, the critical Mg concentration for photosynthetic processes might be lower than for biomass and yield formation.

Critical Leaf Magnesium Thresholds and the Impact of Magnesium on Plant Growth and Photo-Oxidative Defense: A Systematic Review and Meta-Analysis From 70 Years of Research.

Magnesium (Mg) deficiency in plants is a widespread problem affecting productivity and quality in agricultural systems and forestry. Although numerous studies addressed the effect of Mg deficiency on biomass and photosynthetic CO2 assimilation, a summary evaluation of the effect of Mg supply on plant growth and photosynthesis is so far missing. We performed a systematic review and meta-analysis to collect and combine all relevant scientifically published data on the relationship between Mg nutrition and parameters that can be related to plant growth such as root and shoot biomass, harvestable yield, net CO2 assimilation and antioxidant enzyme activities. Moreover, this data pool was used to calculate critical Mg leaf concentrations for biomass and net CO2 assimilation for various plant species. Summarizing all studies included in our analysis, adequate Mg supply enhances net CO2 assimilation by 140%, leading to a biomass increase of 61% compared to Mg deficient control plants. Biomass partitioning between shoot and root is not only sensitive to Mg nutrition, but highly affected by the experimental cultivation technique. If plants are grown under adequate Mg supply during initial growth stages before exposing them to Mg deficiency, the shoot-root ratio was not affected. Otherwise, the shoot-root ratio significantly decreased in contrast to Mg deficient control plants. Concentration of reactive oxygen species decreased under adequate Mg supply by 31% compared to Mg deficient plants, resulting in decreased activities of most antioxidant enzymes and metabolites under adequate Mg supply. We combined all published data relating leaf Mg concentrations to growth and found a critical leaf Mg range for dry weight between 0.1 and 0.2% which was valid for numerous crop species such as wheat, potato, rice, maize, sorghum and barley. Critical leaf Mg concentrations for net CO2 assimilation were higher than for biomass for most species, e.g., potato, rice, citrus, and cotton. In conclusion, our evaluation can be used to identify Mg nutritional status in plants and may help to optimize fertilization strategies. It quantifies the demand of Mg for various crop and tree species for maintaining important physiological processes such as net CO2 assimilation that is required for optimal plant growth and yield.

Interactive effects of nitrogen and potassium on photosynthesis and photosynthetic nitrogen allocation of rice leaves.

BACKGROUND: Nitrogen (N) and potassium (K) are two important mineral nutrients in regulating leaf photosynthesis. Studying the interactive effects of N and K on regulating N allocation and photosynthesis (Pn) of rice leaves will be of great significance for further increasing leaf Pn, photosynthetic N use efficiency (PNUE) and grain yield. We measured the gas exchange of rice leaves in a field experiment and tested different kinds of leaf N based on N morphology and function, and calculated the interactive effects of N and K on N allocation and the PNUE. RESULTS: Compared with N0 (0 kg N ha- 1) and K0 (0 kg K2O ha- 1) treatments, the Pn was increased by 17.1 and 12.2% with the supply of N and K. Compared with N0K0 (0 kg N and 0 kg K2O ha- 1), N0K120 (0 kg N and 120 kg K2O ha- 1) and N0K180 (0 kg N and 180 kg K2O ha- 1), N supply increased the absolute content of photosynthetic N (Npsn) by 15.1, 15.5 and 10.5% on average, and the storage N (Nstore) was increased by 32.7, 64.9 and 72.7% on average. The relative content of Npsn was decreased by 5.6, 12.1 and 14.5%, while that of Nstore was increased by 8.7, 27.8 and 33.8%. Supply of K promoted the transformation of Nstore to Npsn despite the leaf N content (Na) was indeed decreased. Compared with N0K0, N180K0 (180 kg N and 0 kg K2O ha- 1) and N270K0 (270 kg N and 0 kg K2O ha- 1), K supply increased the relative content of Npsn by 17.7, 8.8 and 7.3%, and decreased the relative content of Nstore by 24.2, 11.4 and 8.7% respectively. CONCLUSIONS: This study indicated the mechanism that K supply decreased the Na but increased the Npsn content and then increased leaf Pn and PNUE from a new viewpoint of leaf N allocation. The supply of K promoted the transformation of Nstore to Npsn and increased the PNUE. The decreased Nstore mainly resulted from the decrease of non-protein N. Combined use of N and K could optimize leaf N allocation and maintain a high leaf Npsn content and PNUE.

Functioning of potassium and magnesium in photosynthesis, photosynthate translocation and photoprotection.

Potassium (K) and magnesium (Mg) are mineral nutrients that are required in large quantities by plants. Both elements critically contribute to the process of photosynthesis and the subsequent long-distance transport of photoassimilates. If K or Mg is not present in sufficient quantities in photosynthetic tissues, complex interactions of anatomical, physiological and biochemical responses result in a reduction of photosynthetic carbon assimilation. As a consequence, excessive production of reactive oxygen species causes photo-oxidation of the photosynthetic apparatus and causes an up-regulation of photoprotective mechanisms. In this article, we review the functioning of K and Mg in processes directly or indirectly associated with photosynthesis. Focus is given to chloroplast ultrastructure, light-dependent and -independent reactions of photosynthesis and the diffusion of CO2  - a major substrate for photosynthesis - into chloroplasts. We further emphasize their contribution to phloem-loading and long-distance transport of photoassimilates and to the photoprotection of the photosynthetic apparatus.

Quantitative limitations to photosynthesis in K deficient sunflower and their implications on water-use efficiency.

Potassium (K) is crucial for crop growth and is strongly related to stress tolerance and water-use efficiency (WUE). A major physiological effect of K deficiency is the inhibition of net CO2 assimilation (AN) during photosynthesis. Whether this reduction originates from limitations either to photochemical energy conversion or biochemical CO2 fixation or from a limitation to CO2 diffusion through stomata and the leaf mesophyll is debated. In this study, limitations to photosynthetic carbon gain of sunflower (Helianthus annuus L.) under K deficiency and PEG- induced water deficit were quantified and their implications on plant- and leaf-scale WUE (WUEP, WUEL) were evaluated. Results show that neither maximum quantum use efficiency (Fv/Fm) nor in-vivo RubisCo activity were directly affected by K deficiency and that the observed impairment of AN was primarily due to decreased CO2 mesophyll conductance (gm). K deficiency additionally impaired leaf area development which, together with reduced AN, resulted in inhibition of plant growth and a reduction of WUEP. Contrastingly, WUEL was not affected by K supply which indicated no inhibition of stomatal control. PEG-stress further impeded AN by stomatal closure and resulted in enhanced WUEL and high oxidative stress. It can be concluded from this study that reduction of gm is a major response of leaves to K deficiency, possibly due to changes in leaf anatomy, which negatively affects AN and contributes to the typical symptoms like oxidative stress, growth inhibition and reduced WUEP.

The activity of nodules of the supernodulating mutant Mtsunn is not limited by photosynthesis under optimal growth conditions.

Legumes match the nodule number to the N demand of the plant. When a mutation in the regulatory mechanism deprives the plant of that ability, an excessive number of nodules are formed. These mutants show low productivity in the fields, mainly due to the high carbon burden caused through the necessity to supply numerous nodules. The objective of this study was to clarify whether through optimal conditions for growth and CO2 assimilation a higher nodule activity of a supernodulating mutant of Medicago truncatula (M. truncatula) can be induced. Several experimental approaches reveal that under the conditions of our experiments, the nitrogen fixation of the supernodulating mutant, designated as sunn (super numeric nodules), was not limited by photosynthesis. Higher specific nitrogen fixation activity could not be induced through short- or long-term increases in CO2 assimilation around shoots. Furthermore, a whole plant P depletion induced a decline in nitrogen fixation, however this decline did not occur significantly earlier in sunn plants, nor was it more intense compared to the wild-type. However, a distinctly different pattern of nitrogen fixation during the day/night cycles of the experiment indicates that the control of N2 fixing activity of the large number of nodules is an additional problem for the productivity of supernodulating mutants.