A complete understanding of ionome homeostasis requires a thorough investigation of the dynamics of the nutrient networks in plants. This review focuses on the complexity of interactions occurring ...between S and other nutrients, and these are addressed at the level of the whole plant, the individual tissues, and the cellular compartments. With regards to macronutrients, S deficiency mainly acts by reducing plant growth, which in turn restricts the root uptake of, for example, N, K, and Mg. Conversely, deficiencies in N, K, or Mg reduce uptake of S. TOR (target of rapamycin) protein kinase, whose involvement in the co-regulation of C/N and S metabolism has recently been unravelled, provides a clue to understanding the links between S and plant growth. In legumes, the original crosstalk between N and S can be found at the level of nodules, which show high requirements for S, and hence specifically express a number of sulfate transporters. With regards to micronutrients, except for Fe, their uptake can be increased under S deficiency through various mechanisms. One of these results from the broad specificity of root sulfate transporters that are up-regulated during S deficiency, which can also take up some molybdate and selenate. A second mechanism is linked to the large accumulation of sulfate in the leaf vacuoles, with its reduced osmotic contribution under S deficiency being compensated for by an increase in Cl uptake and accumulation. A third group of broader mechanisms that can explain at least some of the interactions between S and micronutrients concerns metabolic networks where several nutrients are essential, such as the synthesis of the Mo co-factor needed by some essential enzymes, which requires S, Fe, Zn and Cu for its synthesis, and the synthesis and regulation of Fe-S clusters. Finally, we briefly review recent developments in the modelling of S responses in crops (allocation amongst plant parts and distribution of mineral versus organic forms) in order to provide perspectives on prediction-based approaches that take into account the interactions with other minerals such as N.
Human mineral malnutrition or hidden hunger is considered a global challenge, affecting a large proportion of the world’s population. The reduction in the mineral content of edible plant products is ...frequently found in cultivars bred for higher yields, and is probably increased by intensive agricultural practices. The filling of grain with macro and micronutrients is partly the result of a direct allocation from root uptake and remobilization from vegetative tissues. The aim of this bibliographic review is to focus on recent knowledge obtained from ionomic analysis of plant tissues in order to build a global appraisal of the potential remobilization of all macro and micronutrients, and especially those from leaves. Nitrogen is always remobilized from leaves of all plant species, although with different efficiencies, while nutrients such as K, S, P, Mg, Cu, Mo, Fe and Zn can be mobilized to a certain extent when plants are facing deficiencies. On the opposite, there is few evidence for leaf mobilization of Ca, Mn, Ni and B. Mechanisms related to the remobilization process (remobilization of mineral forms from vacuolar and organic compounds associated with senescence, respectively) are also discussed in the context of drought, an abiotic stress that is thought to increase and known to modulate the ionic composition of grain in crops.
Higher plants have to cope with fluctuating mineral resource availability. However, strategies such as stimulation of root growth, increased transporter activities, and nutrient storage and ...remobilization have been mostly studied for only a few macronutrients. Leaves of cultivated crops (Zea mays, Brassica napus, Pisum sativum, Triticum aestivum, Hordeum vulgare) and tree species (Quercus robur, Populus nigra, Alnus glutinosa) grown under field conditions were harvested regularly during their life span and analyzed to evaluate the net mobilization of 13 nutrients during leaf senescence. While N was remobilized in all plant species with different efficiencies ranging from 40% (maize) to 90% (wheat), other macronutrients (K-P-S-Mg) were mobilized in most species. Ca and Mn, usually considered as having low phloem mobility were remobilized from leaves in wheat and barley. Leaf content of Cu-Mo-Ni-B-Fe-Zn decreased in some species, as a result of remobilization. Overall, wheat, barley and oak appeared to be the most efficient at remobilization while poplar and maize were the least efficient. Further experiments were performed with rapeseed plants subjected to individual nutrient deficiencies. Compared to field conditions, remobilization from leaves was similar (N-S-Cu) or increased by nutrient deficiency (K-P-Mg) while nutrient deficiency had no effect on Mo-Zn-B-Ca-Mn, which seemed to be non-mobile during leaf senescence under field conditions. However, Ca and Mn were largely mobilized from roots (-97 and -86% of their initial root contents, respectively) to shoots. Differences in remobilization between species and between nutrients are then discussed in relation to a range of putative mechanisms.
Including more grain legumes in cropping systems is important for the development of agroecological practices and the diversification of protein sources for human and animal consumption. Grain legume ...yield and quality is impacted by abiotic stresses resulting from fluctuating availabilities in essential nutrients such as iron deficiency chlorosis (IDC). Promoting plant iron nutrition could mitigate IDC that currently impedes legume cultivation in calcareous soils, and increase the iron content of legume seeds and its bioavailability. There is growing evidence that plant microbiota contribute to plant iron nutrition and might account for variations in the sensitivity of pea cultivars to iron deficiency and in fine to seed nutritional quality. Pyoverdine (pvd) siderophores synthesized by pseudomonads have been shown to promote iron nutrition in various plant species (Arabidopsis, clover and grasses). This study aimed to investigate the impact of three distinct ferripyoverdines (Fe-pvds) on iron status and the ionome of two pea cultivars (cv.) differing in their tolerance to IDC, (cv. S) being susceptible and (cv. T) tolerant. One pvd came from a pseudomonad strain isolated from the rhizosphere of cv. T (pvd1T), one from cv. S (pvd2S), and the third from a reference strain C7R12 (pvdC7R12). The results indicated that Fe-pvds differently impacted pea iron status and ionome, and that this impact varied both according to the pvd and the cultivar. Plant iron concentration was more increased by Fe-pvds in cv. T than in cv. S. Iron allocation within the plant was impacted by Fe-pvds in cv. T. Furthermore, Fe-pvds had the greatest favorable impact on iron nutrition in the cultivar from which the producing strain originated. This study evidences the impact of bacterial siderophores on pea iron status and pea ionome composition, and shows that this impact varies with the siderophore and host-plant cultivar, thereby emphasizing the specificity of these plant-microorganisms interactions. Our results support the possible contribution of pyoverdine-producing pseudomonads to differences in tolerance to IDC between pea cultivars. Indeed, the tolerant cv. T, as compared to the susceptible cv. S, benefited from bacterial siderophores for its iron nutrition to a greater extent.
Legume plants, such as peas, are of significant nutritional interest for both humans and animals. However, plant nutrition and thus, seed composition, depends on soil mineral nutrient availability. ...Understanding the impact of their deprivation on the plant mineral nutrient content, net uptake, and remobilization is of key importance but remains complex as the elements of the plant ionome are linked in intricate networks, one element deprivation impacting uptake and remobilization of other nutrients. To get a better insight into pea mineral nutrition, the transitory deprivations of 13 mineral nutrients were imposed during the vegetative growth phase. Thereafter, plants were grown under optimal mineral conditions until physiological maturity. Plant nutritional status and seed quality impacts caused by the deprivations were characterized using measurement of mineral nutrient concentration and plant biomass allocation. Our results highlight: (i) the preferential allocation of dry weight and elements to shoots at the expense of the roots under non-limiting conditions, and more particularly to the tendrils in comparison to the other shoot organs, (ii) the positive and/or negative impact of one mineral nutrient deprivation on other elements of the ionome, (iii) four different remobilization strategies for eight mineral nutrients, and (iv) possible strategies to improve seed quality
fine control of fertilization during a period of mineral nutrient deficiency.
To investigate the regulatory interactions between S assimilation and N metabolism in Brassica napus, de novo synthesis of amino acids and proteins was quantified by 15N and 34S tracing, and the ...responses of transporter genes, assimilatory enzymes and metabolites pool involving in nitrate and sulfate metabolism were assessed under continuous sulfur supply, sulfur deprivation and sulfate resupply after 3 days of sulfur (S) deprivation. S-deprived plants were characterized by a strong induction of sulfate transporter genes, ATP sulfurylase (ATPS) and adenosine 5′-phosphosulfate reductase (APR), and by a repressed activity of nitrate reductase (NR) and glutamine synthetase (GS). Sulfate resupply to the S-deprived plants strongly increased cysteine, amino acids and proteins concentration. The increase in sulfate and cysteine concentration caused by sulfate resupply was not matched with the expression of sulfate transporters and the activity of ATPS and APR which were rapidly decreased by sulfate resupply. A strong induction of O-acetylserine(thiol)lyase (OASTL), NR and GS upon sulfate resupply was accompanied with the increase in cysteine, amino acids and proteins pool. Sulfate resupply resulted in a strong increase in de novo synthesis of amino acids and proteins, as evidenced by the increases in N and S incorporation into amino acids (1.8- and 2.4-fold increase) and proteins (2.2-and 6.3-fold increase) when compared to S-deprived plants. The results thus indicate that sulfate resupply followed by S-deprivation accelerates nitrate assimilation for protein synthesis.
•N and S assimilation was assessed by SO42− resupply followed by S-deprivation.•S-resupply activated rapidly the activity of NR, GS and OASTL.•ATPS and APR activity does not match with the proteins increased by S-resupply.•S-resupply increased the incorporation of 15N and 34S to amino acids and proteins.•S-resupply accelerates nitrate assimilation mainly for protein synthesis.
To investigate the varietal difference in sulfur use efficiency (SUE) and drought stress tolerance, Brassica napus 'Mosa' and 'Saturnin' were exposed to polyethylene glycol (PEG)-induced drought ...stress for 72 h. Direct quantification of S uptake, de novo synthesis of amino acids and proteins was performed by tracing (34)S. The responses of photosynthetic activity in relation to SUE were also examined. The total amount of newly absorbed S decreased with drought stress in both cultivars but the decrease rate was significantly higher in Mosa (-64%) than in Saturnin (-41%). Drought stress also decreased the amount of S assimilated into amino acids ((34)S-amino acids) and proteins ((34)S-proteins). The total amount of S incorporated into amino acids and proteins was generally higher in Saturnin (663.7 μg S per plant) than in Mosa (337.3 μg S per plant). The estimation of SUE based on S uptake (SUpE) and S assimilation (SUaE) showed that SUE was much higher in Saturnin than in Mosa. The inhibition of photosynthetic activity including Rubisco protein degradation caused by drought stress was much lower in the cultivar with higher SUE (Saturnin). The present study clearly indicates that the genotype with higher SUE is more tolerant to PEG-induced drought stress.
While it is generally acknowledged that drought is one of the main abiotic factors affecting plant growth, how mineral nutrition is specifically and negatively affected by water deficit has received ...very little attention, other than being analyzed as a consequence of reduced growth. Therefore,
plants were subjected to a gradual onset of water deficits (mild, severe, or severe extended), and leaves were analyzed at the ionomic, transcriptomic and metabolic levels. The number of Differentially Expressed Genes (DEGs) and of the most differentially accumulated metabolites increased from mild (525 DEGs, 57 metabolites) to severe (5454 DEGs, 78 metabolites) and severe extended (9346 DEGs, 95 metabolites) water deficit. Gene ontology enrichment analysis of the 11,747 DEGs identified revealed that ion transport was one of the most significant processes affected, even under mild water deficit, and this was also confirmed by the shift in ionomic composition (mostly micronutrients with a strong decrease in Mo, Fe, Zn, and Mn in leaves) that occurred well before growth reduction. The metabolomic data and most of the transcriptomic data suggested that well-known early leaf responses to drought such as phytohormone metabolism (ABA and JA), proline accumulation, and oxidative stress defense were induced later than repression of genes related to nutrient transport.
Elevated tropospheric ozone concentration (O₃) increases oxidative stress in vegetation and threatens the stability of crop production. Current O₃ pollution in the United States is estimated to ...decrease the yields of maize (
) up to 10%, however, many bioenergy feedstocks including switchgrass (
) have not been studied for response to O₃ stress. Using Free Air Concentration Enrichment (FACE) technology, we investigated the impacts of elevated O₃ (~100 nmol mol
) on leaf photosynthetic traits and capacity, chlorophyll fluorescence, the Ball⁻Woodrow⁻Berry (BWB) relationship, respiration, leaf structure, biomass and nutrient composition of switchgrass. Elevated O₃ concentration reduced net CO₂ assimilation rate (
), stomatal conductance (
), and maximum CO₂ saturated photosynthetic capacity (
), but did not affect other functional and structural traits in switchgrass or the macro- (except potassium) and micronutrient content of leaves. These results suggest that switchgrass exhibits a greater O₃ tolerance than maize, and provide important fundamental data for evaluating the yield stability of a bioenergy feedstock crop and for exploring O₃ sensitivity among bioenergy feedstocks.
Identification of early sulphur (S) deficiency indicators is important for species such as Brassica napus, an S-demanding crop in which yield and the nutritional quality of seeds are negatively ...affected by S deficiency. Because S is mostly stored as SO₄2- in leaf cell vacuoles and can be mobilized during S deficiency, this study investigated the impact of S deprivation on leaf osmotic potential in order to identify compensation processes. Plants were exposed for 28 days to S or to chlorine deprivation in order to differentiate osmotic and metabolic responses. While chlorine deprivation had no significant effects on growth, osmotic potential and nitrogen metabolism, Brassica napus revealed two response periods to S deprivation. The first one occurred during the first 13 days during which plant growth was maintained as a result of vacuolar SO₄2- mobilization. In the meantime, leaf osmotic potential of S-deprived plants remained similar to control plants despite a reduction in the SO₄2- osmotic contribution, which was fully compensated by an increase in NO₃⁻, PO₄3- and Cl⁻ accumulation. The second response occurred after 13 days of S deprivation with a significant reduction in growth, leaf osmotic potential, NO₃⁻ uptake and NO₃⁻ reductase activity, whereas amino acids and NO₃⁻ were accumulated. This kinetic analysis of S deprivation suggested that a (Cl⁻+NO₃⁻+PO₄3-):SO₄2- ratio could provide a relevant indicator of S deficiency, modified nearly as early as the over-expression of genes encoding SO₄2- tonoplastic or plasmalemmal transporters, with the added advantage that it can be easily quantified under field conditions.
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