Differential modulation of Target of Rapamycin activity under single and combined iron and sulfur deficiency in tomato plants.

Over the past few decades, a close relationship between sulfur (S) and iron (Fe) in terms of functionality and nutrition was demonstrated in the tomato. However, very little is known about the regulatory mechanisms underlying S/Fe interactions. Recently, the potential role of citrate in plant adaptation to Fe deficiency and combined S and Fe deficiency has been described. It is known that an impaired organic acid metabolism may stimulate a retrograde signal, which has been proven to be linked to the Target of Rapamycin (TOR) signaling in yeast and animal cells. Recent reports provided evidence of TOR involvement in S nutrient sensing in plants. This suggestion prompted us to investigate whether TOR may play a role in the cross-talk of signaling pathway occurring during plant adaptation to combined nutrient deficiency of Fe and S. Our results revealed that Fe deficiency elicited an increase of TOR activity associated with enhanced accumulation of citrate. In contrast, S deficiency resulted in decreased TOR activity and citrate accumulation. Interestingly, citrate accumulated in shoots of plants exposed to combined S/Fe deficiency to values between those found in Fe- and S-deficient plants, again correlated with TOR activity level. Our results suggest that citrate might be involved in establishing a link between plant response to combined S/Fe deficiency and the TOR network.

Interaction between Sulfate and Selenate in Tetraploid Wheat (Triticum turgidum L.) Genotypes.

Selenium (Se) is an essential micronutrient of fundamental importance to human health and the main Se source is from plant-derived foods. Plants mainly take up Se as selenate (SeO42-), through the root sulfate transport system, because of their chemical similarity. The aims of this study were (1) to characterize the interaction between Se and S during the root uptake process, by measuring the expression of genes coding for high-affinity sulfate transporters and (2) to explore the possibility of increasing plant capability to take up Se by modulating S availability in the growth medium. We selected different tetraploid wheat genotypes as model plants, including a modern genotype, Svevo (Triticum turgidum ssp. durum), and three ancient Khorasan wheats, Kamut, Turanicum 21, and Etrusco (Triticum turgidum ssp. turanicum). The plants were cultivated hydroponically for 20 days in the presence of two sulfate levels, adequate (S = 1.2 mM) and limiting (L = 0.06 mM), and three selenate levels (0, 10, 50 µM). Our findings clearly showed the differential expression of genes encoding the two high-affinity transporters (TdSultr1.1 and TdSultr1.3), which are involved in the primary uptake of sulfate from the rhizosphere. Interestingly, Se accumulation in shoots was higher when S was limited in the nutrient solution.

Single and Combined Fe and S Deficiency Differentially Modulate Root Exudate Composition in Tomato: A Double Strategy for Fe Acquisition?

Fe chlorosis is considered as one of the major constraints on crop growth and yield worldwide, being particularly worse when associated with S shortage, due to the tight link between Fe and S. Plant adaptation to inadequate nutrient availabilities often relies on the release of root exudates that enhance nutrients, mobilization from soil colloids and favour their uptake by roots. This work aims at characterizing the exudomic profile of hydroponically grown tomato plants subjected to either single or combined Fe and S deficiency, as well as at shedding light on the regulation mechanisms underlying Fe and S acquisition processes by plants. Root exudates have been analysed by untargeted metabolomics, through liquid chromatography-mass spectrometry as well as gas chromatography-mass spectrometry following derivatization. More than 200 metabolites could be putatively annotated. Venn diagrams show that 23%, 10% and 21% of differential metabolites are distinctively modulated by single Fe deficiency, single S deficiency or combined Fe-S deficiency, respectively. Interestingly, for the first time, a mugineic acid derivative is detected in dicot plants root exudates. The results seem to support the hypothesis of the co-existence of the two Fe acquisition strategies in tomato plants.

The potential of two different Avena sativa L. cultivars to alleviate Cu toxicity.

Agronomic strategies as intercropping might be applied to reduce plant-available copper (Cu) in Cu-contaminated soils. Thus, our aim was to characterize two different oat cultivars, Avena sativa L. cv. Fronteira and cv. Perona for their ability to tolerate and/or phytostabilize Cu. Copper toxicity reduced plant biomass of both cultivars. The exudate analysis revealed the presence of phenolic compounds and phytosiderophores, yet with a different pattern between the cultivars: cv. Fronteira showed a Cu-concentration and time-dependent release of phenolic compounds, while cv. Perona down-regulated this release during the second week of treatment. Copper concentration increased linearly in all the tissues analysed with increasing Cu concentration showing yet a different compartmentalization: cv. Fronteira and cv. Perona preferentially accumulated Cu in the apoplasm and symplast, respectively. This higher accumulation of Cu in the apoplasm strongly reduces the available binding sites, leading to a competitive absorption with other macro-and micronutrients (e.g. Ca, Mn, Zn). Furthermore, in both cultivars Cu toxicity led to a significant reduction of shoot phosphorus content. The ionomic profile and compartmentalization of Cu together with the root activities demonstrate the different tolerance mechanism towards Cu toxicity of the two oat cultivars. In particular, cv. Fronteira seems to adopt an exclusion strategy based on accumulating Cu in the apoplasm and on the exudation of phenolic compounds. Thus, this cultivar could reduce the mobility and the consequent soil bioavailability of Cu playing an important role as phytostabilizer plant in intercropping systems in Cu-contaminated vineyards or orchards.

The Interplay between Sulfur and Iron Nutrition in Tomato.

Plant response mechanisms to deficiency of a single nutrient, such as sulfur (S) or iron (Fe), have been described at agronomic, physiological, biochemical, metabolomics, and transcriptomic levels. However, agroecosystems are often characterized by different scenarios, in which combined nutrient deficiencies are likely to occur. Soils are becoming depleted for S, whereas Fe, although highly abundant in the soil, is poorly available for uptake because of its insolubility in the soil matrix. To this end, earlier reports showed that a limited S availability reduces Fe uptake and that Fe deficiency results in the modulation of sulfate uptake and assimilation. However, the mechanistic basis of this interaction remains largely unknown. Metabolite profiling of tomato (Solanum lycopersicum) shoots and roots from plants exposed to Fe, S, and combined Fe and S deficiency was performed to improve the understanding of the S-Fe interaction through the identification of the main players in the considered pathways. Distinct changes were revealed under the different nutritional conditions. Furthermore, we investigated the development of the Fe deficiency response through the analysis of expression of ferric chelate reductase, iron-regulated transporter, and putative transcription factor genes and plant sulfate uptake and mobilization capacity by analyzing the expression of genes encoding sulfate transporters (STs) of groups 1, 2, and 4 (SlST1.1, SlST1.2, SlST2.1, SlST2.2, and SlST4.1). We identified a high degree of common and even synergistic response patterns as well as nutrient-specific responses. The results are discussed in the context of current models of nutrient deficiency responses in crop plants.

Cadmium exposure affects iron acquisition in barley (Hordeum vulgare) seedlings.

This study addresses the question of the interference between iron (Fe) nutrition and cadmium (Cd) toxicity at the level of growth performance, phytosiderophores (PS) release, micronutrient accumulation and expression of genes involved in Fe homeostasis in barley seedlings, a plant with strategy II-based response to Fe shortage. Cd exposure induced responses similar to those of genuine Fe deficiency also in Fe-sufficient plants. Most genes involved in PS biosynthesis and secretion (HvNAS3, HvNAS4, HvNAS6, HvNAS7, HvNAAT-A, HvDMAS1 and HvTOM1) induced by Fe deprivation were also significantly upregulated in the presence of Cd under Fe sufficient conditions. Accordingly, the enhanced expression of these genes in roots under Cd exposure was accompanied by an increase of PS release. However, induced expression of HvIRO2 and the downregulation of HvIDEF1 and HvIRT1, after Cd exposure, suggested the presence of a pathway that induces HvIRO2-mediated PS biosynthesis under Cd stress, which probably is not simply caused by Fe deficiency. The downregulation of HvIRT1 and HvNramp5 may represent a protective mechanism at transcriptional level against further Cd uptake by these transporters. These results likely indicate that Cd itself may be able to activate Fe acquisition mechanism in an Fe-independent manner.

Iron deprivation results in a rapid but not sustained increase of the expression of genes involved in iron metabolism and sulfate uptake in tomato (Solanum lycopersicum L.) seedlings.

Characterization of the relationship between sulfur and iron in both Strategy I and Strategy II plants, has proven that low sulfur availability often limits plant capability to cope with iron shortage. Here it was investigated whether the adaptation to iron deficiency in tomato (Solanum lycopersicum L.) plants was associated with an increased root sulfate uptake and translocation capacity, and modified dynamics of total sulfur and thiols accumulation between roots and shoots. Most of the tomato sulfate transporter genes belonging to Groups 1, 2, and 4 were significantly upregulated in iron-deficient roots, as it commonly occurs under S-deficient conditions. The upregulation of the two high affinity sulfate transporter genes, SlST1.1 and SlST1.2, by iron deprivation clearly suggests an increased root capability to take up sulfate. Furthermore, the upregulation of the two low affinity sulfate transporter genes SlST2.1 and SlST4.1 in iron-deficient roots, accompanied by a substantial accumulation of total sulfur and thiols in shoots of iron-starved plants, likely supports an increased root-to-shoot translocation of sulfate. Results suggest that tomato plants exposed to iron-deficiency are able to change sulfur metabolic balance mimicking sulfur starvation responses to meet the increased demand for methionine and its derivatives, allowing them to cope with this stress.

Nitrate induction triggers different transcriptional changes in a high and a low nitrogen use efficiency maize inbred line.

In higher plants, NO3(-) can induce its own uptake and the magnitude of this induction is positively related to the external anion concentration. This phenomenon has been characterized in both herbaceous and woody plants. Here, different adaptation strategies of roots from two maize (Zea mays L., ZmAGOs) inbred lines differing in nitrogen use efficiency (NUE) and exhibiting different timing of induction were discussed by investigating NO3(-) -induced changes in their transcriptome. Lo5 line (high NUE) showing the maximum rate of NO3(-) uptake 4 h after the provision of 200 µmol/L NO3(-) treatment modulated a higher number of transcripts relative to T250 (low NUE) that peaked after 12 h. The two inbred lines share only 368 transcripts that are modulated by the treatment with NO3(-) and behaved differently when transcripts involved in anion uptake and assimilation were analyzed. T250 line responded to the NO3(-) induction modulating this group of genes as reported for several plant species. On the contrary, the Lo5 line did not exhibit during the induction changes in this set of genes. Obtained data suggest the importance of exploring the physiological and molecular variations among different maize genotypes in response to environmental clues like NO3(-) provision, in order to understand mechanisms underlying NUE.

Corrigendum: Characterization of Triticum turgidum sspp. durum, turanicum and polonicum grown in Central Italy in relation to technological and nutritional aspects.

[This corrects the article DOI: 10.3389/fpls.2023.1269212.].

Transcriptional and physiological changes in the S assimilation pathway due to single or combined S and Fe deprivation in durum wheat (Triticum durum L.) seedlings.

The effect of iron (Fe) and sulphur (S) deprivation on sulphate uptake and assimilation pathways was investigated in durum wheat by analysing the expression of genes coding for major transporters and enzymes involved in sulphate assimilation and reduction: high-affinity sulphate transporters (TdSultr1.1 and TdSultr1.3), ATP sulphurylase (TdATPSul1 and TdATPSul2), APS reductase (TdAPR), sulphite reductase (TdSiR), O-acetylserine(thiol)lyase (TdOASTL1 and TdOASTL2), and serine acetyltransferase (TdSAT1 and TdSAT2). Further experiments were carried out to detect changes in the activities of these enzymes, together with the evaluation of growth parameters (fresh biomass accumulation, leaf green values, and total S, thiol, and Fe concentrations). Fe shortage in wheat plants under adequate S nutrition resulted in an S deficiency-like response. Most of the genes of the S assimilatory pathway induced by S deprivation (TdATPSul1, TdAPR, TdSir, TdSAT1, and TdSAT2) were also significantly up-regulated after the imposition of the Fe limitation under S-sufficient conditions. However, the differential expression of genes encoding the two high-affinity transporters (TdSultr1.1 and TdSultr1.3) indicates that the mechanisms of sulphate uptake regulation under Fe and S deficiency are different in wheat. Moreover, it was observed that the mRNA level of genes encoding ATPS, APR, and OASTL and the corresponding enzyme activities were often uncoupled in response to Fe and S availability, indicating that most probably their regulation involves a complex interplay of transcriptional, translational, and/or post-translational mechanisms induced by S and/or Fe deficiency.

Characterization of Triticum turgidum sspp. durum, turanicum, and polonicum grown in Central Italy in relation to technological and nutritional aspects.

Introduction: Wheat is a staple food, with the two most common species being Triticum aestivum and Triticum turgidum ssp. durum. Moreover, the latter, T. turgidum, includes other tetraploid subspecies, among which the sspp. turanicum (Khorasan wheat) and polonicum (Polish wheat), whose importance has increased in the last decades, representing alternative crops for marginal areas, in addition to being a source of genetic diversity. Methods: In this work, different accessions of these three subspecies of T. turgidum have been grown in 2 years in the same environment and have been characterized for technological properties and factors affecting nutritional quality, such as fiber amount and the content of micro- and macro-nutrients in grains, and for root morphological traits. Results: These analyses allowed the identification, in particular, of a Polish wheat accession showing better technological performances, a higher amount of positive micro- and macro-elements, and a lower amount of toxic cadmium. The modern variety Svevo and the Polish Pol2 showed the lowest and the highest shoot:root ratio, respectively. The high shoot:root ratio in Pol2 was mainly attributable to the decrease in root growth. Although Pol2 had a lower root biomass, its particular root morphology made it more efficient for nutrient uptake, as evident from the greater accumulation of micro- and macro-nutrients. Discussion: These results underline that it is not possible to draw general conclusions about the difference between primitive and modern wheats, but rather a case-by-case approach should be chosen.

The suppression of TdMRP3 genes reduces the phytic acid and increases the nutrient accumulation in durum wheat grain.

Micronutrient malnutrition affects more than half of the world population. Reduced bioavailability of microelements in the raw materials is considered one of the main causes of mineral deficiency in populations whose diet is largely based on the consumption of staple crops. In this context, the production of low phytic acid (lpa) cereals is a main goal of the breeding programs, as phytic acid (PA) binds essential mineral cations such as iron (Fe), zinc (Zn), manganese (Mn), potassium (K), calcium (Ca) and magnesium (Mg) precipitating in the form of phytate salts poorly digested by monogastric animals, including humans, due to the lack of phytases in the digestive tract. Since PA limits the bioavailability of microelements, it is widely recognized as an anti-nutritional compound. A Targeting Induced Local Lesions IN Genomes (TILLING) approach has been undertaken to silence the genes encoding the TdABCC13 proteins, known as Multidrug-Resistance associated Proteins 3 (TdMRP3), transporters involved in the accumulation of PA inside the vacuole in durum wheat. The TdMRP3 complete null genotypes showed a significant reduction in the content of PA and were able to accumulate a higher amount of essential micronutrients (Fe, Zn, Mn) compared to the control. The number of spikelets and seeds per spike, traits associated with the agronomic performances, were reduced compared to the control, but the negative effect was in part balanced by the increased grain weight. The TdMRP3 mutant lines showed morphological differences in the root apparatus such as a significant decrease in the number of root tips, root length, volume and surface area and an increase in root average diameter compared to the control plants. These materials represent a promising basis for obtaining new commercial durum wheats with higher nutritional value.

Multiple linear regression and linear mixed models identify novel traits of salinity tolerance in Olea europaea L.

Olive can be considered as moderately tolerant to salinity, with marked differences among cultivars. In the present study, two olive cultivars with different salt tolerance, 'Leccino' (sensitive) and 'Frantoio' (tolerant), were treated with 120 mM of NaCl for 30 days. We measured the expression of genes involved in the management of sodium in the leaves, such as NHX, SOS1 and H+ ATPase, and the concentration of Na+, K+, Mn2+, Mg2+ and Ca2+ in the roots, bark, xylem and leaves of the olive plants. The results were analyzed with multiple linear models and mixed models. Furthermore, we utilized the analysis of covariance to find significant relationships in our data. Both cultivars significantly reduced net photosynthesis and increased water-use efficiency after 30 days of treatment. Sodium accumulated significantly in the roots of both cultivars, and 'Leccino' plants were also able to translocate it to the leaves and the bark. The NHX and vacuolar ATPase subunit E genes were significantly overexpressed in both the cultivars treated with NaCl. The SOS1, ATPase11 and ATPase8 genes were overexpressed in 'Frantoio'. The covariance between gene expression and element concentrations data was analyzed to identify significant interactions between cultivars and treatments. Na+ accumulation in the roots of 'Frantoio' was positively related to the accumulation of K+, Mn2+, Mg2+ and Ca2+ in the xylem, bark and leaves. 'Frantoio' capability to mobilize elements, especially Ca2+, together with the overexpression of key genes for sodium management, could be crucial for salt tolerance.

Arbuscular Mycorrhizal Symbiosis Differentially Affects the Nutritional Status of Two Durum Wheat Genotypes under Drought Conditions.

Durum wheat is one of the most important agricultural crops, currently providing 18% of the daily intake of calories and 20% of daily protein intake for humans. However, being wheat that is cultivated in arid and semiarid areas, its productivity is threatened by drought stress, which is being exacerbated by climate change. Therefore, the identification of drought tolerant wheat genotypes is critical for increasing grain yield and also improving the capability of crops to uptake and assimilate nutrients, which are seriously affected by drought. This work aimed to determine the effect of arbuscular mycorrhizal fungi (AMF) on plant growth under normal and limited water availability in two durum wheat genotypes (Svevo and Etrusco). Furthermore, we investigated how the plant nutritional status responds to drought stress. We found that the response of Svevo and Etrusco to drought stress was differentially affected by AMF. Interestingly, we revealed that AMF positively affected sulfur homeostasis under drought conditions, mainly in the Svevo cultivar. The results provide a valuable indication that the identification of drought tolerant plants cannot ignore their nutrient use efficiency or the impact of other biotic soil components (i.e., AMF).

Interaction Between Sulfur and Iron in Plants.

It is well known that S interacts with some macronutrients, such as N, P, and K, as well as with some micronutrients, such as Fe, Mo, Cu, Zn, and B. From our current understanding, such interactions could be related to the fact that: (i) S shares similar chemical properties with other elements (e.g., Mo and Se) determining competition for the acquisition/transport process (SULTR transporter family proteins); (ii) S-requiring metabolic processes need the presence of other nutrients or regulate plant responses to other nutritional deficiencies (S-containing metabolites are the precursor for the synthesis of ethylene and phytosiderophores); (iii) S directly interacts with other elements (e.g., Fe) by forming complexes and chemical bonds, such as Fe-S clusters; and (iv) S is a constituent of organic molecules, which play crucial roles in plants (glutathione, transporters, etc.). This review summarizes the current state of knowledge of the interplay between Fe and S in plants. It has been demonstrated that plant capability to take up and accumulate Fe strongly depends on S availability in the growth medium in both monocots and dicot plants. Moreover, providing S above the average nutritional need enhances the Fe content in wheat grains, this beneficial effect being particularly pronounced under severe Fe limitation. On the other hand, Fe shortage induces a significant increase in the demand for S, resulting in enhanced S uptake and assimilation rate, similar to what happens under S deficiency. The critical evaluation of the recent studies on the modulation of Fe/S interaction by integrating old and new insights gained on this topic will help to identify the main knowledge gaps. Indeed, it remains a challenge to determine how the interplay between S and Fe is regulated and how plants are able to sense environmental nutrient fluctuations and then to adapt their uptake, translocation, assimilation, and signaling. A better knowledge of the mechanisms of Fe/S interaction might considerably help in improving crop performance within a context of limited nutrient resources and a more sustainable agriculture.

Plasticity, exudation and microbiome-association of the root system of Pellitory-of-the-wall plants grown in environments impaired in iron availability.

The investigation of the adaptive strategies of wild plant species to extreme environments is a challenging issue, which favors the identification of new traits for plant resilience. We investigated different traits which characterize the root-soil interaction of Parietaria judaica, a wild plant species commonly known as "Pellitory-of-the-wall". P. judaica adopts the acidification-reduction strategy (Strategy I) for iron (Fe) acquisition from soil, and it can complete its life cycle in highly calcareous environments without any symptoms of chlorosis. In a field-to-lab approach, the microbiome associated with P. judaica roots was analyzed in spontaneous plants harvested from an urban environment consisting in an extremely calcareous habitat. Also, the phenolics and carboxylates content and root plasticity and exudation were analyzed in P. judaica plants grown under three different controlled conditions mimicking the effect of calcareous environments on Fe availability: results show that P. judaica differentially modulates root plasticity under different Fe availability-impaired conditions, and that it induces, to a high extent, the exudation of caffeoylquinic acid derivatives under calcareous conditions, positively impacting Fe solubility.

Response of Olive Shoots to Salinity Stress Suggests the Involvement of Sulfur Metabolism.

Global warming has two dangerous global consequences for agriculture: drought, due to water scarcity, and salinization, due to the prolonged use of water containing high concentrations of salts. Since the global climate is projected to continue to change over this century and beyond, choosing salt-tolerant plants could represent a potential paramount last resort for exploiting the secondary saline soils. Olive is considered moderately resistant to soil salinity as compared to other fruit trees, and in the present study, we investigated the influence of NaCl solutions (ranging from 0 to 200 mM) in a salt-tolerant (cv Canino) and two of its transgenic lines (Canino AT17-1 and Canino AT17-2), overexpressing tobacco osmotin gene, and in a salt-sensitive (Sirole) olive cultivar. After four weeks, most of the shoots of both Canino and Sirole plants showed stunted growth and ultimate leaf drop by exposure to salt-enriched media, contrary to transgenic lines, that did not show injuries and exhibited a normal growth rate. Malondialdehyde (MDA) content was also measured as an indicator of the lipid peroxidation level. To evaluate the role of the S assimilatory pathway in alleviating the adverse effects of salt stress, thiols levels as well as extractable activities of ATP sulfurylase (ATPS) and O-acetyl serine(thiol)lyase (OASTL), the first and the last enzyme of the S assimilation pathway, respectively, have been estimated. The results have clearly depicted that both transgenic lines overexpressing osmotin gene coped with increasing levels of NaCl by the induction of S metabolism, and particularly increase in OASTL activity closely paralleled changes of NaCl concentration. Linear correlation between salt stress and OASTL activity provides evidence that the S assimilation pathway plays a key role in adaptive response of olive plants under salt stress conditions.

Supply of sulphur to S-deficient young barley seedlings restores their capability to cope with iron shortage.

The effect of the S nutritional status on a plant's capability to cope with Fe shortage was studied in solution cultivation experiments in barley (Hordeum vulgare L. cv. Europa). Barley is a Strategy II plant and responds to Fe deficiency by secretion of chelating compounds, phytosiderophores (PS). All PS are derived from nicotianamine whose precursor is methionine. This suggests that a long-term supply of an inadequate amount of S could reduce a plant's capability to respond to Fe deficiency by limiting the rate of PS biosynthesis. The responses of barley (Hordeum vulgare L. cv. Europa) plants grown for 12 d on Fe-free nutrient solutions (NS) containing 0 or 1.2 mM SO(4)(2-), was examined after 24 h or 48 h from transfer to NS containing 1.2 mM SO(4)(2-). After the supply of S was restored to S-deprived plants, an increase in PS release in root exudates was evident after 24 h of growth in S-sufficient NS and the increment reached values up to 4-fold higher than the control 48 h after S resupply. When S was supplied to S-deficient plants, leaf ATPS (EC 2.7.7.4) and OASTL (EC 4.2.99.8) activities exhibited a progressive recovery. Furthermore, root HvST1 transcript abundance remained high for 48 h following S resupply and a significant increase in the level of root HvYS1 transcripts was also found after only 24 h of S resupply. Data support the idea that the extent to which the plant is able to cope with Fe starvation is strongly associated with its S nutritional status. In particular, our results are indicative that barley plants fully recover their capability to cope with Fe shortage after the supply of S is restored to S-deficient plants.

Evaluating the potential use of Cu-contaminated soils for giant reed (Arundo donax, L.) cultivation as a biomass crop.

Over the past decades, the important topic of environmental sustainability, impact, and security of the fossil fuel supply has stimulated interest in using lignocellulosic feedstocks as biofuel to partially cover energy demands. Among energy no-food crops, giant reed (Arundo donax, L.), a perennial rhizomatous grass has been identified as a leading candidate crop for lignocellulosic feedstock, due to its positive energy balance, and low ecological/agro-management demands. The aim of the present study was to characterize the physiological response of Arundo donax (L.) to artificial soil contamination with three different Cu levels (200, 400, and 800 ppm), and to assess the relationship between plant Cu tolerance and S assimilation rate. The present study not only confirms the ability of Arundo donax L. to cope with Cu stress and therefore to grow in marginal, degraded lands abandoned by mainstream agricultural, but also shows that plant performance might be likely ascribed to a modulation of sulfate metabolism resulting in increased thiols content.

Root-shoot-root Fe translocation in cucumber plants grown in a heterogeneous Fe provision.

Iron (Fe) is an essential micronutrient for plant life and development. However, in soil, Fe bioavailability is often limited and variable in space and time, thus different regions of the same root system might be exposed to different nutrient provisions. Few studies showed that the response to variable Fe provision is controlled at local and systemic levels, albeit the identity of the signals involved is still elusive. Iron itself was suggested as local mediator, whilst hormones were proposed for the long-distance signalling pathway. Therefore, the aim of this work was to assess whether Fe, when localized in a restricted area of the root system, might be involved in both local and systemic signaling. The combination of resupply experiments in a split-root system, the use of 57Fe isotope and chemical imaging techniques allowed tracing Fe movement within cucumber plants. Soon after the resupply, Fe is distributed to the whole plant, likely to overcome a minimum Fe concentration threshold aimed at repressing the deficiency response. Iron was then preferentially translocated to leaves and, only afterwards, the root system was completely resupplied. Collectively, these observations might thus highlight a root-to-shoot-to-root Fe translocation route in cucumber plants grown on a patchy nutrient substrate.