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A fully mechanistic dynamical model for plant nitrate uptake is presented. Based on physiological and regulatory pathways and based on physical laws, we form a dynamic system mathematically described by seven differential equations. The model evidences the presence of a short‐term positive feedback on the high‐affinity nitrate uptake, triggered by the presence of nitrate around the roots, which induces its intaking. In the long run, this positive feedback is overridden by two long‐term negative feedback loops which drastically reduces the nitrate uptake capacity. These two negative feedbacks are due to the generation of ammonium and amino acids, respectively, and inhibit the synthesis and the activity of high‐affinity nitrate transporters. This model faithfully predicts the typical spiking behavior of the nitrate uptake, in which an initial strong increase of nitrate absorption capacity is followed by a drop, which regulates the absorption down to the initial value. The model outcome was compared with experimental data and they fit quite nicely. The model predicts that after the initial exposure of the roots with nitrate, the absorption of the anion strongly increases and that, on the contrary, the intensity of the absorption is limited in presence of ammonium around the roots. [ABSTRACT FROM AUTHOR]

A fully mechanistic dynamical model for plant nitrate uptake is presented. Based on physiological and regulatory pathways and based on physical laws, we form a dynamic system mathematically described by seven differential equations. The model evidences the presence of a short‐term positive feedback on the high‐affinity nitrate uptake, triggered by the presence of nitrate around the roots, which induces its intaking. In the long run, this positive feedback is overridden by two long‐term negative feedback loops which drastically reduces the nitrate uptake capacity. These two negative feedbacks are due to the generation of ammonium and amino acids, respectively, and inhibit the synthesis and the activity of high‐affinity nitrate transporters. This model faithfully predicts the typical spiking behavior of the nitrate uptake, in which an initial strong increase of nitrate absorption capacity is followed by a drop, which regulates the absorption down to the initial value. The model outcome was compared with experimental data and they fit quite nicely. The model predicts that after the initial exposure of the roots with nitrate, the absorption of the anion strongly increases and that, on the contrary, the intensity of the absorption is limited in presence of ammonium around the roots. [ABSTRACT FROM AUTHOR]

The simultaneous presence of different N‐forms in the rhizosphere leads to beneficial effects on nitrogen (N) nutrition in plants. Although widely used as fertilizers, the occurrence of cross connection between urea and ammonium nutrition has been scarcely studied in plants. Maize fed with a mixture of urea and ammonium displayed a better N‐uptake efficiency than ammonium‐ or urea‐fed plants (Buoso et al., Plant Physiol Biochem, 2021a; 162: 613–623). Through multiomic approaches, we provide the molecular characterization of maize response to urea and ammonium nutrition. Several transporters and enzymes involved in N‐nutrition were upregulated by all three N‐treatments (urea, ammonium, or urea and ammonium). Already after 1 day of treatment, the availability of different N‐forms induced specific transcriptomic and metabolomic responses. The combination of urea and ammonium induced a prompt assimilation of N, characterized by high levels of some amino acids in shoots. Moreover, ZmAMT1.1a, ZmGLN1;2, ZmGLN1;5, ZmGOT1, and ZmGOT3, as well transcripts involved in glycolysis‐TCA cycle were induced in roots by urea and ammonium mixture. Depending on N‐form, even changes in the composition of phytohormones were observed in maize. This study paves the way to formulate guidelines for the optimization of N fertilization to improve N‐use efficiency in maize and therefore limit N‐losses in the environment. [ABSTRACT FROM AUTHOR]

Despite the wide use of urea and ammonium as N-fertilizers, no information is available about the proper ratio useful to maximize the efficiency of their acquisition by crops. Ionomic analyses of maize seedlings fed with five different mixes of urea and ammonium indicated that after 7 days of treatment, the elemental composition of plant tissues was more influenced by ammonium in the nutrient solution than by urea. Within 24 h, similar high affinity influx rates of ammonium were measured in ammonium-treated seedlings, independently from the amount of the cation present in the nutrient solution (from 0.5 to 2.0 mM N), and it was confirmed by the similar accumulation of 15N derived from ammonium source. After 7 days, some changes in ammonium acquisition occurred among treatments, with the highest ammonium uptake efficiency when the urea-to-ammonium ratio was 3:1. Gene expression analyses of enzymes and transporters involved in N nutrition highlight a preferential induction of the cytosolic N-assimilatory pathway (via GS, ASNS) when both urea and ammonium were supplied in conjunction, this response might explain the higher N-acquisition efficiency when both sources are applied. In conclusion, this study provides new insights on plant responses to mixes of N sources that maximize the N-uptake efficiency by crops and thus could allow to adapt agronomic practices in order to limit the economic and environmental impact of N-fertilization. • The simultaneous presence of urea and ammonium in the nutrient solution promotes the acquisition of ammonium. • The multielement composition of maize seedlings is more influenced by ammonium in the external media than by urea. • This study provides useful indication about the proper ratio urea to ammonium useful to maximize the uptake efficiency of ammonium in maize. [ABSTRACT FROM AUTHOR]

Humic substances are known to positively influence plant growth and nutrition. In particular, the water-extractable fraction of humic substances (WEHS) has been shown to enhance nitrate acquisition, increasing the activity of high affinity nitrate uptake system. However, molecular bases of this physiological response are not clarified so far. Thus, in the present work, the physiological effect of WEHS on nitrate acquisition in maize roots was correlated with changes in the root transcriptomic profile. Results confirmed that WEHS caused a faster induction of a higher capacity to take up nitrate in maize roots. Comparing the root transcriptomic profile of Nitrate - and Nitrate   +   WEHS -treated plants with Control (-N) ones, more than 2000 transcripts appeared to be modulated only in the presence of WEHS. Among these, genes involved in nitrate transport and assimilation ( NRT1s, NRT2s, NAR2.1, NR, GS, GOGAT, CNX, UPM ) were strongly modulated by WEHS. Furthermore, also some genes known to be linked to the nitrogen limitation responses were affected by WEHS, as transcripts coding for transcription factors (as LBD37, NIN-like protein, NFY-A, GRF5 ) and enzymes of hormones’ metabolism. The modulation of these transcripts might play a crucial role in coordinating the induction to nitrate, favouring its uptake and assimilation in WEHS-treated plants. The overexpression of nitrogen assimilatory genes by WEHS might led to an early feedback regulation of the high affinity nitrate transport system, as being operated by N-metabolites. Results of the present work shed further light on the contribution of the organic soil component to the nitrogen use efficiency in crops. [ABSTRACT FROM AUTHOR]

Soilless cultivation systems represent an alternative to traditional agriculture, as they offer the possibility to reduce water use and to design nutrient formulations of the hydroponic solutions in order to maximize yield and quality of the products. Reduction in accumulation of undesired compounds, like e.g. nitrate, and enhanced nutritional value for human consumption, like e.g. Se-rich compounds, could be reasonably achieved. These aspects, together with the extension of the shelf life, would increase the market value of ready-to-cut vegetables. This review discusses how recent advances in understanding the role of plant nutrients and their interactions and the management of nutrient solution composition, including the addition of non-essential elements, micronutrients’ biofortification, biostimulants, together with the control of some crucial environmental conditions, like temperature, can help accomplishing this task. [ABSTRACT FROM AUTHOR]

There is increasing interest in the hydroponic technology to produce leafy vegetables for ready-to-eat salads. Optimisation of the growing system can lead to higher yield and/or improved nutritional value of the product. Selenium (Se) is an essential element for animal and humans, with quite a narrow range between deficiency and toxicity, whereas it is assumed beneficial for plants. In the present study, two cultivars (Gala and Baron) of corn salad (Valerianella locusta (L.) Laterr.) were used to test the possibility to increase Se content in the edible parts (leaves). Effects on yield, nitrate content, and accumulation of sulfur (S) and S-containing amino acids and Se and Se-containing amino acids were studied. Results showed that corn salad tolerates selenate (Na[sub 2]SeO[sub 4]) concentrations ranging from 10 to 40 μM in the nutrient solution, with plants accumulating Se at levels compatible with the need in human diets at 10 μM selenate. Se-treated plants showed some benefits with respect to a decrease of nitrate concentration and increase of pigment contents (chlorophylls and carotenoids). At 10 μM selenate, Se-cysteine and Se-methionine were produced, without affecting non-protein thiols or cysteine and methionine contents. At the higher Se supply, sulfate accumulated in the leaves with a parallel decrease in the amount of S-amino acids and a rise in the relative amount of Se-amino acids. Based on the chemical analyses, cv. Gala showed better tolerance than cv. Baron to moderate selenate supply (40 μM). [ABSTRACT FROM AUTHOR]

Iron (Fe) sources available for plants in the rhizospheric solution are mainly a mixture of complexes between Fe and organic ligands, including phytosiderophores (PS) and water-extractable humic substances (WEHS). In comparison with the other Fe sources, Fe-WEHS are more efficiently used by plants, and experimental evidences show that Fe translocation contributes to this better response. On the other hand, very little is known on the mechanisms involved in Fe allocation in leaves. In this work, physiological and molecular processes involved in Fe distribution in leaves of Fe-deficient Cucumis sativus supplied with Fe-PS or Fe-WEHS up to 5 days were studied combining different techniques, such as radiochemical experiments, synchrotron micro X-ray fluorescence, real-time reverse transcription polymerase chain reaction and in situ hybridization. In Fe-WEHS-fed plants, Fe was rapidly (1 day) allocated into the leaf veins, and after 5 days, Fe was completely transferred into interveinal cells; moreover, the amount of accumulated Fe was much higher than with Fe-PS. This redistribution in Fe-WEHS plants was associated with an upregulation of genes encoding a ferricIII-chelate reductase (FRO), a Fe2+ transporter (IRT1) and a natural resistance-associated macrophage protein (NRAMP). The localization of FRO and IRT1 transcripts next to the midveins, beside that of NRAMP in the interveinal area, may suggest a rapid and efficient response induced by the presence of Fe-WEHS in the extra-radical solution for the allocation in leaves of high amounts of Fe. In conclusion, Fe is more efficiently used when chelated to WEHS than PS and seems to involve Fe distribution and gene regulation of Fe acquisition mechanisms operating in leaves. [ABSTRACT FROM AUTHOR]

Background Despite its extensive use as a nitrogen fertilizer, the role of urea as a directly accessible nitrogen source for crop plants is still poorly understood. So far, the physiological and molecular aspects of urea acquisition have been investigated only in few plant species highlighting the importance of a high-affinity transport system. With respect to maize, a worldwide-cultivated crop requiring high amounts of nitrogen fertilizer, the mechanisms involved in the transport of urea have not yet been identified. The aim of the present work was to characterize the high-affinity urea transport system in maize roots and to identify the high affinity urea transporter. Results Kinetic characterization of urea uptake (<300 μM) demonstrated the presence in maize roots of a high-affinity and saturable transport system; this system is inducible by urea itself showing higher Vmax and Km upon induction. At molecular level, the ORF sequence coding for the urea transporter, ZmDUR3, was isolated and functionally characterized using different heterologous systems: a dur3 yeast mutant strain, tobacco protoplasts and a dur3 Arabidopsis mutant. The expression of the isolated sequence, ZmDUR3-ORF, in dur3 yeast mutant demonstrated the ability of the encoded protein to mediate urea uptake into cells. The subcellular targeting of DUR3/GFP fusion proteins in tobacco protoplasts gave results comparable to the localization of the orthologous transporters of Arabidopsis and rice, suggesting a partial localization at the plasma membrane. Moreover, the overexpression of ZmDUR3 in the atdur3-3 Arabidopsis mutant showed to complement the phenotype, since different ZmDUR3-overexpressing lines showed either comparable or enhanced 15[N]-urea influx than wild-type plants. These data provide a clear evidence in planta for a role of ZmDUR3 in urea acquisition from an extra-radical solution. Conclusions This work highlights the capability of maize plants to take up urea via an inducible and highaffinity transport system. ZmDUR3 is a high-affinity urea transporter mediating the uptake of this molecule into roots. Data may provide a key to better understand the mechanisms involved in urea acquisition and contribute to deepen the knowledge on the overall nitrogenuse efficiency in crop plants. [ABSTRACT FROM AUTHOR]

Plants mainly rely on a mixture of Fe complexes with different organic ligands, like carboxylates and soluble fractions of water-extractable humic substances (WEHSs), to sustain the supply of this micronutrient. It has been demonstrated that the Fe-WEHS complex is more efficiently acquired by plant roots as it enhances functionality of the mechanisms involved in Fe acquisition at the root and leaf levels, allowing a faster recovery of the Fe-deficiency symptoms. The aim of this work is to verify whether this recovery involves also the allocation and accumulation of nutrients other than Fe to and within the leaf tissues. Iron-deficient plants treated with Fe-WEHS recovered more quickly the functionality both to uptake nitrate at the root level and to fixate CO in the leaves than those supplied with Fe-citrate. Concomitantly, Fe-WEHS-treated plants also accumulated other cationic nutrients faster and at a higher extent. Synchrotron 2D-scanning μ-X-ray fluorescence analyses of the leaves revealed that the recovery promotes a change in the allocation of these nutrients from the vascular system (K, Cu, and Zn) or trichomes (Ca and Mn) to the entire leaf blade. Fe-WEHS treatment efficiently promotes the recovery from Fe-deficiency-induced chlorosis with an enhanced allocation of other nutrients into the leaves and promoting their distribution into the entire leaf blade. [ABSTRACT FROM AUTHOR]

The aim of this work is to evaluate the capability of tomato plants to use different Fe sources, such as Fe citrate, Fe phytosiderophores, and Fe complexed by a water-extractable humic substances (Fe-WEHS) also in relation to physiological and molecular adaptations induced by these complexes at the root level. Tomato plants acquired higher amounts of Fe from Fe-WEHS than from the other two sources and this phenomenon occurred only when the treatment lasted 24 h. The higher acquisition of Fe from Fe-WEHS than other sources depended on a reductive mechanism and on rhizosphere acidification and appeared to be due neither to a higher apoplastic loading nor to a higher resistance of WEHS to microbial degradation. Supply of the different Fe complexes to deficient plants induced a transient upregulation of Fe(III)-chelate reductase ( LeFRO1) and Fe transporter genes, LeIRT1 and LeIRT2. In Fe-WEHS-fed plants, where a quicker and higher upregulation of these genes was evident, a coordination in the expression of LeFRO1, LeIRT1, and LeIRT2 genes occurred already after 1 h treatment when the amount of Fe acquired by the plants from the three sources was similar. Iron from Fe-WEHS could be efficiently acquired in a mixture of natural Fe complexes possibly occurring in the rhizosphere. This phenomenon is due to an altered expression of Fe uptake-related genes and to the root capacity to create favorable conditions for the micronutrient uptake into the rhizosphere. [ABSTRACT FROM AUTHOR]

Soil temperature has a crucial impact on physiological processes and growth of plants with important consequences for plant productivity and food safety including nitrate accumulation in leaf blades of leaf vegetables. Consumer demand for high-quality, fresh-cut vegetables has increased rapidly in the last decades, and temperature modulation can help control nitrate concentration in fresh vegetables, an important trait of product safety. Corn salad plants IValerianella Iocusta (L.) Laterr., cultivar Gala] were grown at three root temperatures (15, 20, and 25 °C) in a floating system. This experimental setup allowed to directly evaluate the effect of root temperature on yield and plant quality excluding the effect on soil processes and properties. Nutrient solution was renewed weekly and kept aerated while air temperature was maintained constant at 20 °C for all treatments during the entire time of experiments. At harvest, plants were collected, the shelf life evaluated, and the nutrient uptake [NO3-, iron (Fe) from 59Fe-o,oEDDHA, and 35SO42-] and mineral content were determined. Results showed that growing conditions at 20 °C of the nutrient solution led to the best plant performance in terms of yield, nitrate content at leaf level, root biomass, leaf area, and greenness with positive effects on postharvest quality, i.e., less rapid leaf loss of greenness and leaf fresh weight (FW) loss during conservation at 4 °C. At this temperature condition of the nutrient solution, it has also been observed an enhanced functionality of mechanisms involved in the acquisition of nutrients like NO3-, Fe, and SO42-, which are known to play an important role in nitrate level in leaf tissues of crops. Plants grown at 15 °C showed minor growth, whereas the nutrient solution at 25 °C caused stress for the plants affecting negatively the quality and yield. Overall, the results obtained showed that root temperature plays a fundamental role in several plant processes that affect yield and its quality; for hydroponic system cultivations, a level of growing-medium temperature close to that of the surrounding air seems suitable. [ABSTRACT FROM AUTHOR]

It is well known that in the rhizosphere soluble Fe sources available for plants are mainly a mixture of complexes between the micronutrient and organic ligands such as organic acids and phytosiderophores (PS) released by roots, microbial siderophores as well as fractions of humified organic matter. In the present work, mechanisms of Fe acquisition operating at the leaf level of plants fed with different Fe-complexes were investigated at the micro-analytical, physiological and molecular levels. Fe-deficient tomato plants ( Solanum Lycopersicum L., cv. ‘Marmande’) were fed for 24 h with a solution (pH 7.5) containing 1 µM Fe as Fe-PS, Fe-citrate or Fe-WEHS. Thereafter, leaf tissue was used for the visualization of Fe distribution, measurements of Fe content, reduction and uptake, and evaluation of expression of Fe-chelate reductase ( LeFRO1), Fe-transporter ( LeIRT1) and Ferritin ( Ferritin2) genes. Leaf discs isolated from Fe-deficient plants treated for 24 h with Fe-WEHS developed higher rates of translocation, Fe-chelate reduction and 59Fe uptake as compared to plants supplied with Fe-citrate or Fe-PS. Leaves of plants treated with Fe-WEHS also showed higher transcript levels of LeFRO1, LeIRT1 and Ferritin2 genes with respect to plants fed with the other Fe-sources. Data obtained support the idea that the efficient use of Fe complexed to WEHS-like humic fractions involves, at least in part, also the activation of Fe-acquisition mechanisms operating at the leaf level. [ABSTRACT FROM AUTHOR]

Iron chlorosis is a wide-spread limiting factor of production in agriculture. To cope with this problem, synthetic chelates (like EDTA or EDDHA) of Fe are used in foliar-spray or in soil treatments; however, these products are very expensive. Therefore paper-production byproducts, like Lignosulfonates (LS), with varying content of carboxylate and sulfonate groups, were tested with respect to their ability to maintain Fe in the solution of soils and to feed plants grown in hydroponics with Fe through foliar sprays or application to the nutrient solution. Results show that LS had a low capability to solubilize 59Fe-hydroxide and that preformed 59Fe(III)-LS complexes had poor mobility through a soil column (pH 7.5) and scarce stability when interacting with soils compared to 59Fe(III)-EDDHA. However when 59Fe(III)-LS were supplied to roots in a hydroponic system, they demonstrated an even higher capability to fed Fe-deficient tomato plants than 59Fe(III)-EDDHA. Hence, data here presented indicate that the low Fe use efficiency from Fe-LS observed in soil-applications is due to interactions of these Fe-sources with soil colloids rather than to the low capability of roots to use them. Foliar application experiments of 59Fe(III)-LS or 59Fe(III)-EDTA to Fe-deficient cucumber plants show that uptake and reduction rates of Fe were similar between all these complexes; on the other hand, when 59Fe(III)-LS were sprayed on Fe-deficient tomato leaves, they showed a lower uptake rate, but a similar reduction rate, than 59Fe(III)-EDTA did. In conclusion, Fe-LS may be a valid, eco-compatible and cheap alternative to synthetic chelates in dealing with Fe chlorosis when applied foliarly or in the nutrient solution of hydroponically grown plants. [ABSTRACT FROM AUTHOR]

White lupin ( Lupinus albus L.) is able to grow on soils with sparingly available phosphate (P) by producing specialized structures called cluster roots. To mobilize sparingly soluble P forms in soils, cluster roots release substantial amounts of carboxylates and concomitantly acidify the rhizosphere. The relationship between acidification and carboxylate exudation is still largely unknown. In the present work, we studied the linkage between organic acids (malate and citrate) and proton exudations in cluster roots of P-deficient white lupin. After the illumination started, citrate exudation increased transiently and reached a maximum after 5 h. This effect was accompanied by a strong acidification of the external medium and alkalinization of the cytosol, as evidenced by in vivo nuclear magnetic resonance (NMR) analysis. Fusicoccin, an activator of the plasma membrane (PM) H+-ATPase, stimulated citrate exudation, whereas vanadate, an inhibitor of the H+-ATPase, reduced citrate exudation. The burst of citrate exudation was associated with an increase in expression of the LHA1 PM H+-ATPase gene, an increased amount of H+-ATPase protein, a shift in pH optimum of the enzyme and post-translational modification of an H+-ATPase protein involving binding of activating 14-3-3 protein. Taken together, our results indicate a close link in cluster roots of P-deficient white lupin between the burst of citrate exudation and PM H+-ATPase-catalysed proton efflux. [ABSTRACT FROM AUTHOR]

Abstract: The impact of flavonoids released by phosphorus-deficient white lupin roots on inorganic P and soil microorganisms is largely unknown. We report that flavonoids isolated from white lupin roots mobilized inorganic phosphorus and decreased soil microbial respiration, citrate mineralization, and soil phosphohydrolase activities, but did not reduce the soil ATP content. The results suggest that white lupin''s release of flavonoids into the rhizosphere plays a significant role in its efficient P-acquisition strategy by solubilizing Fe-bound P and by limiting the microbial mineralization of citrate. [Copyright &y& Elsevier]

Background: In viticulture, iron (Fe) chlorosis is a common abiotic stress that impairs plant development and leads to yield and quality losses. Under low availability of the metal, the applied N form (nitrate and ammonium) can play a role in promoting or mitigating Fe deficiency stresses. However, the processes involved are not clear in grapevine. Therefore, the aim of this study was to investigate the response of two grapevine rootstocks to the interaction between N forms and Fe uptake. This process was evaluated in a hydroponic experiment using two ungrafted grapevine rootstocks Fercal (Vitis berlandieri x V. vinifera) tolerant to deficiency induced Fe chlorosis and Couderc 3309 (V. riparia x V. rupestris) susceptible to deficiency induced Fe chlorosis. Results: The results could differentiate Fe deficiency effects, N-forms effects, and rootstock effects. Interveinal chlorosis of young leaves appeared earlier on 3309 C from the second week of treatment with NO3−/NH4+ (1:0)/-Fe, while Fercal leaves showed less severe symptoms after four weeks of treatment, corresponding to decreased chlorophyll concentrations lowered by 75% in 3309 C and 57% in Fercal. Ferric chelate reductase (FCR) activity was by trend enhanced under Fe deficiency in Fercal with both N combinations, whereas 3309 C showed an increase in FCR activity under Fe deficiency only with NO3−/NH4+ (1:1) treatment. With the transcriptome analysis, Gene Ontology (GO) revealed multiple biological processes and molecular functions that were significantly regulated in grapevine rootstocks under Fe-deficient conditions, with more genes regulated in Fercal responses, especially when both forms of N were supplied. Furthermore, the expression of genes involved in the auxin and abscisic acid metabolic pathways was markedly increased by the equal supply of both forms of N under Fe deficiency conditions. In addition, changes in the expression of genes related to Fe uptake, regulation, and transport reflected the different responses of the two grapevine rootstocks to different N forms. Conclusions: Results show a clear contribution of N forms to the response of the two grapevine rootstocks under Fe deficiency, highlighting the importance of providing both N forms (nitrate and ammonium) in an appropriate ratio in order to ease the rootstock responses to Fe deficiency. [ABSTRACT FROM AUTHOR]

Aim: The rate of nitrate (NO3−) uptake and changes in rhizosphere properties were studied growing seedlings of two maize inbred lines differing in nitrogen use efficiency (NUE) in rhizoboxes.Results: Changes in NO3− uptake rates occurred in response to anion addition (induction) in seedlings grown both in hydroponic culture and in soil in rhizoboxes. The characterization of root exudate composition showed a line-specific metabolite profile, which was also affected by NO3− availability. The induction affected respiration, nitrification, ammonification and enzyme activities of the rhizosphere. Furthermore, the composition of rhizosphere bacterial communities of the two maize lines differed suggesting the selective capacity of plants.Conclusions: Overall, results showed a strong and fast modification of rhizospheric soil properties in response to physiological changes in plants caused by fluctuating NO3− availability. [ABSTRACT FROM AUTHOR]

Background: It is well known that in the rhizosphere soluble Fe sources available for plants are mainly represented by a mixture of complexes between the micronutrient and organic ligands such as carboxylates and phytosiderophores (PS) released by roots, as well as fractions of humified organic matter. The use by roots of these three natural Fe sources (Fe-citrate, Fe-PS and Fe complexed to water-extractable humic substances, Fe-WEHS) have been already studied at physiological level but the knowledge about the transcriptomic aspects is still lacking. Results: The 59Fe concentration recorded after 24 h in tissues of tomato Fe-deficient plants supplied with 59Fe complexed to WEHS reached values about 2 times higher than those measured in response to the supply with Fe-citrate and Fe-PS. However, after 1 h no differences among the three Fe-chelates were observed considering the 59Fe concentration and the root Fe(III) reduction activity. A large-scale transcriptional analysis of root tissue after 1 h of Fe supply showed that Fe-WEHS modulated only two transcripts leaving the transcriptome substantially identical to Fe-deficient plants. On the other hand, Fe-citrate and Fe-PS affected 728 and 408 transcripts, respectively, having 289 a similar transcriptional behaviour in response to both Fe sources. Conclusions: The root transcriptional response to the Fe supply depends on the nature of chelating agents (WEHS, citrate and PS). The supply of Fe-citrate and Fe-PS showed not only a fast back regulation of molecular mechanisms modulated by Fe deficiency but also specific responses due to the uptake of the chelating molecule. Plants fed with Fe-WEHS did not show relevant changes in the root transcriptome with respect to the Fe-deficient plants, indicating that roots did not sense the restored cellular Fe accumulation. [ABSTRACT FROM AUTHOR]

White lupin is very often used as a model plant for root exudation studies due to its capability to release huge amounts of organic acids and flavonoids. The complex nature of these organic compounds makes not only their analytical determination difficult but also their extraction from soil samples. For these reasons simplified approaches, as hydroponic-based systems are widely used to study the root exudation. Therefore, the composition of a trap solution is crucial to limit artefacts causing over/underestimation of exudation rates and/or a biased molecular composition of the collected compounds. The present study was aimed at assessing the influence of different trap solutions and collection times on the quali- and quantitative root exudation pattern of white lupin ( Lupinus albus L.) grown under phosphorus (P) and iron (Fe) deficiency. Our results suggest that, in works aimed at studying root exudation processes, water is the most effective trap solution to collect the exudates like organic acids and flavonoids, especially in short time (e.g. 2 h). For longer times, low concentrations of Ca could be helpful to limit osmotic stress and possible passive leakage and/or diffusion. The use of bacteriostatic compounds as NaN and Micropur bias the results, due to interferences either with the metabolism or inhibition of the exudation processes, especially in the case of flavonoids such as quercetin. Also, the use of a pH buffer solution like 2-(N-morpholino)ethanesulfonic acid (MES) should be avoided for its undesired interferences with the release. [ABSTRACT FROM AUTHOR]

Nitrogen (N) is the nutrient most applied in agriculture as fertilizer (as nitrate, Nit; ammonium, A; and/or urea, U, forms) and its availability strongly constrains the crop growth and yield. To investigate the early response (24 h) of N-deficient tomato plants to these three N forms, a physiological and molecular study was performed. In comparison to N-deficient plants, significant changes in the transcriptional, metabolomic and ionomic profiles were observed. As a probable consequence of N mobility in plants, a wide metabolic modulation occurred in old leaves rather than in young leaves. The metabolic profile of U and A-treated plants was more similar than Nit-treated plant profile, which in turn presented the lowest metabolic modulation with respect to N-deficient condition. Urea and A forms induced some changes at the biosynthesis of secondary metabolites, amino acids and phytohormones. Interestingly, a specific up-regulation by U and down-regulation by A of carbon synthesis occurred in roots. Along with the gene expression, data suggest that the specific N form influences the activation of metabolic pathways for its assimilation (cytosolic GS/AS and/or plastidial GS/GOGAT cycle). Urea induced an up-concentration of Cu and Mn in leaves and Zn in whole plant. This study highlights a metabolic reprogramming depending on the N form applied, and it also provide evidence of a direct relationship between urea nutrition and Zn concentration. The understanding of the metabolic pathways activated by the different N forms represents a milestone in improving the efficiency of urea fertilization in crops. [Display omitted] • Ammonium induce an up concentration of Gln and a down concentration of Glu in leaves. • Asn is accumulated in leaves and roots irrespective from the nitrogen-form applied. • Plant secondary metabolism is particularly modulated by ammonium and urea. • Zinc concentration have a direct relationship with the nitrogen source supplied. [ABSTRACT FROM AUTHOR]

Plant growth-promoting rhizobacteria (PGPR) are soil bacteria that are able to colonize rhizosphere and to enhance plant growth by means of a wide variety of mechanisms like organic matter mineralization, biological control against soil-borne pathogens, biological nitrogen fixation, and root growth promotion. A very interesting feature of PGPR is their ability of enhancing nutrient bioavailability. Several bacterial species have been characterized as P-solubilizing microorganisms while other species have been shown to increase the solubility of micronutrients, like those that produce siderophores for Fe chelation. The enhanced amount of soluble macro- and micronutrients in the close proximity of the soil-root interface has indeed a positive effect on plant nutrition. Furthermore, several pieces of evidence highlight that the inoculation of plants with PGPR can have considerable effects on plant at both physiological and molecular levels (e.g., induction of rhizosphere acidification, up- and downregulation of genes involved in ion uptake, and translocation), suggesting the possibility that soil biota could stimulate plants being more efficient in retrieving nutrients from soil and coping with abiotic stresses. However, the molecular mechanisms underlying these phenomena, the signals involved as well as the potential applications in a sustainable agriculture approach, and the biotechnological aspects for possible rhizosphere engineering are still matters of discussion. [ABSTRACT FROM AUTHOR]

The low bioavailability of nutrients, especially nitrogen (N) and phosphorus (P), is one of the most limiting factors for crop production. In this study, under N- and P-free nutrient solution (-N-P), nodulating white lupin plants developed some nodules and analogous cluster root structures characterized by different morphological, physiological, and molecular responses than those observed upon single nutrient deficiency (strong acidification of external media, a better nutritional status than -N+P and +N-P plants). The multi-elemental analysis highlighted that the concentrations of nutrients in white lupin plants were mainly affected by P availability. Gene-expression analyses provided evidence of interconnections between N and P nutritional pathways that are active to promote N and P balance in plants. The root exudome was mainly characterized by N availability in nutrient solution, and, in particular, the absence of N and P in the nutrient solution triggered a high release of phenolic compounds, nucleosides monophosphate and saponines by roots. These morphological, physiological, and molecular responses result from a close interplay between N and P nutritional pathways. They contribute to the good development of nodulating white lupin plants when grown on N- and P-free media. This study provides evidence that limited N and P availability in the nutrient solution can promote white lupin-Bradyrhizobium symbiosis, which is favourable for the sustainability of legume production. [ABSTRACT FROM AUTHOR]

Abstract: Soil-less cultivation of horticultural crops represents a fairly recent innovation to traditional agriculture which has several advantages including higher water-use efficiency. When plants are grown with this system, their roots come in contact with nutrients solely via the hydroponic solution. Although its beneficial effects have been widely demonstrated, silicon (Si) is mostly omitted from the composition of nutrient solutions. Therefore, the objective of this study was to assess the beneficial effect of Si addition to hydroponic solution on quali-quantitative aspects of edible production of two cultivars of corn salad (Valerianella locusta (L.) Laterr.) grown in soil-less floating system. Impacts on shelf life of this food were also studied. Results show that the supply of Si increased the edible yield and the quality level reducing the nitrate concentration in edible tissues. This result might be attributed to changes either in the metabolism (such as the nitrate assimilation process) or to the functionality of root mechanisms involved in the nutrient acquisition from the outer medium. In fact, our results show for the first time the ability of Si to modulate the root activity of nitrate and Fe uptake through, at least in part, a regulation of gene expression levels of the proteins involved in this phenomenon. In addition, the presence of Si decreased the levels of polyphenoloxidase gene expression at harvest and, in post-harvest, slowed down the chlorophyll degradation delaying leaf senescence and thus prolonging the shelf life of these edible tissues. In conclusion, data showed that the addition of Si to the nutrient solution can be a useful tool for improving quali-quantitatively the yield of baby leaf vegetable corn salad as well as its shelf life. Since the amelioration due to the Si has been achieved only with one cultivar, the recommendation of its inclusion in the nutrient solution does not exclude the identification of cultivars suitable for this cultivation system and the comprehension of agronomical and environmental factors which could limit the Si benefits. [Copyright &y& Elsevier]

White lupin (Lupinus albusL. cv. Amiga) reacts to phosphate deficiency by producing cluster roots which exude large amounts of organic acids. The detailed knowledge of the excretion physiology of the different root parts makes it a good model plant to study plant-bacteria interaction. Since the effect of the organic acid exudation by cluster roots on the rhizosphere microflora is still poorly understood, we investigated the abundance, diversity and functions of bacteria associated with the cluster roots of white lupin, with special emphasis on the influence of root proximity (comparing root, rhizosphere soil and bulk soil fractions) and cluster root growth stages, which are characterized by different excretion activities. Plants were grown for five weeks in microcosms, in the presence of low phosphate concentrations, on acidic sand inoculated with a soil suspension from a lupin field. Plate counts showed that bacterial abundance decreased at the stage where the cluster root excretes high amounts of citrate and protons.In vitrotests on isolates showed that the frequencies of auxin producers were highest in juvenile and mature cluster roots and significantly decreased in senescent cluster roots. However, no significant difference in the frequency of auxin producers was found between cluster and non cluster roots. The diversity and structure of bacterial communities were investigated by DGGE of 16S rDNA and 16S rRNA. The diversity and community structure were mostly influenced by root proximity and, to a lesser extent, by cluster root stage. The richness of bacterial communities decreased with root proximity, whereas the proportion of active populations increased. The high citrate and proton excretion occurring at the mature stage of cluster roots had a strong impact on the structure and richness of the bacterial communities, both in the root and in the rhizosphere soil. [ABSTRACT FROM AUTHOR]

In plants the communication between organs is mainly carried out via the xylem and phloem. The concentration and the molecular species of some phytohormones, assimilates and inorganic ions that are translocated in the xylem vessel play a key role in the systemic nutritional signaling in plants. In this work the composition of the xylem sap of maize was investigated at the metabolic and ionomic level depending on the N form available in the nutrient solution. Plants were grown up to 7 days in hydroponic system under N-free nutrient solution or nutrient solution containing N in form of nitrate, urea, ammonium or a combination of urea and ammonium. For the first time this work provides evidence that the ureic nutrition reduced the water translocation in maize plants more than mineral N forms. This result correlates with those obtained from the analyses of photosynthetic parameters (stomatal conductance and transpiration rate) suggesting a parsimonious use of water by maize plants under urea nutrition. A peculiar composition in amino acids and phytohormones (i.e. S, Gln, Pro, ABA) of the xylem sap under urea nutrition could explain differences in xylem sap exudation in comparison to plants treated with mineral N forms. The knowledge improvement of urea nutrition will allow to further perform good agronomic strategies to improve the resilience of maize crop to water stress. • Urea reduced the water translocation in maize plants more than mineral N forms. • Results suggest a parsimonious use of water by maize plants under urea nutrition. • N form changes the amino acid and phytohormone composition of the xylem sap. [ABSTRACT FROM AUTHOR]

Nitrogen (N) deficiency is one of the major stresses that crops are exposed to. It is plausible to suppose that a stress condition can induce a memory in plants that might prime the following generations. Here, an experimental setup that considered four successive generations of N-sufficient and N-limited Arabidopsis was used to evaluate the existence of a transgenerational memory. The results demonstrated that the ability to take up high amounts of nitrate is induced more quickly as a result of multigenerational stress exposure. This behavior was paralleled by changes in the expression of nitrate responsive genes. RNAseq analyses revealed the enduring modulation of genes in downstream generations, despite the lack of stress stimulus in these plants. The modulation of signaling and transcription factors, such as NIGTs, NFYA and CIPK23 might indicate that there is a complex network operating to maintain the expression of N-responsive genes, such as NRT2.1, NIA1 and NIR. This behavior indicates a rapid acclimation of plants to changes in N availability. Indeed, when fourth generation plants were exposed to N limitation, they showed a rapid induction of N-deficiency responses. This suggests the possible involvement of a transgenerational memory in Arabidopsis that allows plants to adapt efficiently to the environment and this gives an edge to the next generation that presumably will grow in similar stressful conditions. [ABSTRACT FROM AUTHOR]

• Occurrence of a strong cross-interaction between Fe and P acquisition mechanisms. • white lupin roots upregulated Fe-responsive genes ascribable to Strategy-I response. • The Strategy-I mechanism was upregulated also in P-deficient clusters roots. • Viceversa , some P-responsive genes were upregulated also in Fe deficient roots. • P and Fe deficiency upregulated glycolysis and phenylpropanoid pathway, respectively. In agricultural soil, the bioavailability of iron (Fe) and phosphorus (P) is often below the plant's requirement causing nutritional deficiency in crops. Under P-limiting conditions, white lupin (Lupinus albus L.) activates mechanisms that promote P solubility in the soil through morphological, physiological and molecular adaptations. Similar changes occur also in Fe-deficient white lupin roots; however, no information is available on the molecular bases of the response. In the present work, responses to Fe and P deficiency and their reciprocal interactions were studied. Transcriptomic analyses indicated that white lupin roots upregulated Fe-responsive genes ascribable to Strategy-I response, this behaviour was mainly evident in cluster roots. The upregulation of some components of Fe-acquisition mechanism occurred also in P-deficient cluster roots. Concerning P acquisition, some P-responsive genes (as phosphate transporters and transcription factors) were upregulated by P deficiency as well by Fe deficiency. These data indicate a strong cross-connection between the responses activated under Fe or P deficiency in white lupin. The activation of Fe- and P-acquisition mechanisms might play a crucial role to enhance the plant's capability to mobilize both nutrients in the rhizosphere, especially P from its associated metal cations. [ABSTRACT FROM AUTHOR]

Background: Under limited iron (Fe) availability maize, a Strategy II plant, improves Fe acquisition through the release of phytosiderophores (PS) into the rhizosphere and the subsequent uptake of Fe-PS complexes into root cells. Occurrence of Strategy-I-like components and interactions with phosphorous (P) nutrition has been hypothesized based on molecular and physiological studies in grasses. Results: In this report transcriptomic analysis (NimbleGen microarray) of Fe deficiency response revealed that maize roots modulated the expression levels of 724 genes (508 up- and 216 down-regulated, respectively). As expected, roots of Fe-deficient maize plants overexpressed genes involved in the synthesis and release of 2'-deoxymugineic acid (the main PS released by maize roots). A strong modulation of genes involved in regulatory aspects, Fe translocation, root morphological modification, primary metabolic pathways and hormonal metabolism was induced by the nutritional stress. Genes encoding transporters for Fe2+ (ZmNRAMP1) and P (ZmPHT1;7 and ZmPHO1) were also up-regulated under Fe deficiency. Fe-deficient maize plants accumulated higher amounts of P than the Fe-sufficient ones, both in roots and shoots. The supply of 1 µM 59Fe, as soluble (Fe-Citrate and Fe-PS) or sparingly soluble (Ferrihydrite) sources to deficient plants, caused a rapid down-regulation of genes coding for PS and Fe(III)-PS transport, as well as of ZmNRAMP1 and ZmPHT1;7. Levels of 32P absorption essentially followed the rates of 59Fe uptake in Fe-deficient plants during Fe resupply, suggesting that P accumulation might be regulated by Fe uptake in maize plants. Conclusions: The transcriptional response to Fe-deficiency in maize roots confirmed the modulation of known genes involved in the Strategy II and revealed the presence of Strategy I components usually described in dicots. Moreover, data here presented provide evidence of a close relationship between two essential nutrients for plants, Fe and P, and highlight a key role played by Fe and P transporters to preserve the homeostasis of these two nutrients in maize plants. [ABSTRACT FROM AUTHOR]

In this study, we provide experimental evidences that in calcareous soils microbial degradation/decomposition of citrate can promote Al-(hydr)oxide precipitation concurrently decreasing copper (Cu) solubility by a coprecipitation process. Citrate is an organic acid anion commonly released by roots to increase nutrient availability or to limit Al toxicity. However, under specific environmental conditions (i.e. high microbial activity of Al-citrate-degrading bacteria, alkaline pH), this organic acid may become ineffective in mobilizing Cu for the plant acquisition process. To demonstrate this, a calcareous soil and an artificial soil system have been treated with citrate solutions; then, changes in Al and Cu solubility and the formation of Cu-containing Al-(hydr)oxides were monitored. Both in experiments with the artificial soil and in those where the soil was inoculated with microbial strains, the formation of Cu-Al coprecipitates not only occurred but was also concurrent with the decrease of Cu and Al solubility. The role of bacteria in metal-citrate complex degradation has been assessed, and the 16S rDNA of bacteria related with these processes has been sequenced for genus identification. Bacteria belonging to Pseudomonas, Sphingomonas, Bradyrhizobium and Sphingopixis have been identified as possible candidates to degrade Al- and Cu-citrate complexes thus triggering the metal precipitation phenomena. [ABSTRACT FROM AUTHOR]

Viticulture represents an important agricultural practice in many countries worldwide. Yet, the continuous use of fungicides has caused copper (Cu) accumulation in soils, which represent a major environmental and toxicological concern. Despite being an important micronutrient, Cu can be a potential toxicant at high concentrations since it may cause morphological, anatomical and physiological changes in plants, decreasing both food productivity and quality. Rhizosphere processes can, however, actively control the uptake and translocation of Cu in plants. In particular, root exudates affecting the chemical, physical and biological characteristics of the rhizosphere, might reduce the availability of Cu in the soil and hence its absorption. In addition, this review will aim at discussing the advantages and disadvantages of agronomic practices, such as liming, the use of pesticides, the application of organic matter, biochar and coal fly ashes, the inoculation with bacteria and/or mycorrhizal fungi and the intercropping, in alleviating Cu toxicity symptoms. [ABSTRACT FROM AUTHOR]

Combinations of different molecules usually co-exuded by plant roots in soil can significantly affect the mobilization of mineral elements from soil. The flavonoids rutin and quercetin appeared to be highly efficient in Fe and Mn mobilization from soil, being rutin about 25 times more effective than citrate in extracting Fe from an alkaline calcareous soil, and 15 times in mobilizing Mn from a slight acidic agricultural soil. Quercetin was particularly effective in mobilizing Mn from the acidic soil and 50 times more efficient than citrate. A significant synergistic effect was detected when either quercetin or rutin was combined with citrate, extracting, respectively, 1.7 and 1.5 times more Mn than what is expected by a simple combination of the two. Sorption processes on soil particles were more relevant for flavonoids than for organic acids for which microbial degradation often prevails, with the only exception of oxalate. Citrate was usually the most efficient organic acid in mobilizing major and trace elements from soil even if oxalate was a better extractant for Cu, Zn, and Ni from the alkaline soil. In the acid soil, a synergistic effect among organic acids was observed only for Mn while in alkaline soil it was observed for Si. The mechanism by which Fe is extracted by rutin in the alkaline soil is a reductive one, with one molecule of rutin being capable of mobilizing two atoms of Fe. Also for Mn in the acid soil, quercetin and rutin solubilize this element by a reductive process. However, when quercetin and rutin are combined with citrate, a complex-forming dissolution mechanism also occurs which increases the mobilization of Mn over the expected rates. [ABSTRACT FROM AUTHOR]

Even though urea and nitrate are the two major nitrogen (N) forms applied as fertilizers in agriculture and occur concomitantly in soils, the reciprocal influence of these two N sources on the mechanisms of their acquisition are poorly understood. Therefore, molecular and physiological aspects of urea and nitrate uptake were investigated in maize (Zea mays), a crop plant consuming high amounts of N. In roots, urea uptake was stimulated by the presence of urea in the external solution, indicating the presence of an inducible transport system. On the other hand, the presence of nitrate depressed the induction of urea uptake and, at the same time, the induction of nitrate uptake was depressed by the presence of urea. The expression of about 60,000 transcripts of maize in roots was monitored by microarray analyses and the transcriptional patterns of those genes involved in nitrogen acquisition were analyzed by real-time reverse transcription-PCR (RT-PCR). In comparison with the treatment without added N, the exposure of maize roots to urea modulated the expression of only very few genes, such as asparagine synthase. On the other hand, the concomitant presence of urea and nitrate enhanced the overexpression of genes involved in nitrate transport (NRT2) and assimilation (nitrate and nitrite reductase, glutamine synthetase 2), and a specific response of 41 transcripts was determined, including glutamine synthetase 1-5, glutamine oxoglutarate aminotransferase, shikimate kinase and arogenate dehydrogenase. Also based on the real-time RT-PCR analysis, the transcriptional modulation induced by both sources might determine an increase in N metabolism promoting a more efficient assimilation of the N that is taken up. [ABSTRACT FROM AUTHOR]

BACKGROUND Plants differ in their response to high aluminium (Al) concentrations, which typically cause toxicity in plants grown on acidic soils. The response depends on plant species and environmental conditions such as substrate and cultivation system. The present study aimed to assess Al-phosphate (P) dynamics in the rhizosphere of two bean species, Phaseolus vulgaris L. var. Red Kidney and Phaseolus lunatus L., in rhizobox experiments. RESULTS Root activity of the bean species induced up to a sevenfold increase in exchangeable Al and up to a 30-fold decrease in extractable P. High soluble Al concentrations triggered the release of plant-specific carboxylates, which differed between soil type and plant species. The results suggest that P. vulgaris L. mitigates Al stress by an internal defence mechanism and P. lunatus L. by an external one, both mechanisms involving organic acids. CONCLUSION Rhizosphere mechanisms involved in Al detoxification were found to be different for P. vulgaris L. and P. lunatus L., suggesting that these processes are plant species-specific. Phaseolus vulgaris L. accumulates Al in the shoots (internal tolerance mechanism), while P. lunatus L. prevents Al uptake by releasing organic acids (exclusion mechanism) into the growth media. © 2013 Society of Chemical Industry [ABSTRACT FROM AUTHOR]

The development of highly brilliant synchrotron facilities all around the world is opening the way to new research in biological sciences including speciation studies of trace elements in plants. In this paper, for the first time, iron (Fe) speciation in xylem sap has been assessed by X-ray absorption near-edge structure (XANES) spectroscopy at the highly brilliant synchrotron PETRA III, beamline P06. Both standard organic Fe-complexes and xylem sap samples of Fe-deficient tomato plants were analyzed. The high photon flux provided by this X-ray synchrotron source allows on one side to obtain good XANES spectra in a reasonable amount of time (approx. 15 min for 200 eV scan) at low Fe concentrations (sub parts-per-million), while on the other hand may cause radiation damage to the sample, despite the sample being cooled by a stream of liquid nitrogen vapor. Standard Fe-complexes such as Fe(III)-succinate, Fe(III)-α-ketoglutarate, and Fe(III)-nicotianamine are somehow degraded when irradiated with synchrotron X-rays and Fe(III) can undergo photoreduction. Degradation of the organic molecules was assessed by HPLC-UV/Vis analyses on the same samples investigated by X-ray absorption spectroscopy (XAS). Fe speciation in xylem sap samples revealed Fe(III) to be complexed by citrate and acetate. Nevertheless, artifacts created by radiation damage cannot be excluded. The use of highly brilliant synchrotrons as X-ray sources for XAS analyses can dramatically increase the sensitivity of the technique for trace elements thus allowing their speciation in xylem sap. However, great attention must be paid to radiation damage, which can lead to biased results. [Figure not available: see fulltext.] [ABSTRACT FROM AUTHOR]

Iron (Fe) is an essential element for plant growth and development; hence determining Fe distribution and concentration inside plant organs at the microscopic level is of great relevance to better understand its metabolism and bioavailability through the food chain. Among the available microanalytical techniques, synchrotron μ-XRF methods can provide a powerful and versatile array of analytical tools to study Fe distribution within plant samples. In the last years, the implementation of new algorithms and detection technologies has opened the way to more accurate (semi)quantitative analyses of complex matrices like plant materials. In this paper, for the first time the distribution of Fe within tomato roots has been imaged and quantified by means of confocal μ-XRF and exploiting a recently developed fundamental parameter-based algorithm. With this approach, Fe concentrations ranging from few hundreds of ppb to several hundreds of ppm can be determined at the microscopic level without cutting sections. Furthermore, Fe (semi)quantitative distribution maps were obtained for the first time by using two opposing detectors to collect simultaneously the XRF radiation emerging from both sides of an intact cucumber leaf. [Figure not available: see fulltext.] [ABSTRACT FROM AUTHOR]

In the present study, five insoluble Fe-humic substance (HS) complexes (Fe-IHS) extracted from Leonardite were synthesized and characterized by Infrared (IR) spectroscopy, scanning electron microscopy (SEM), and dynamic light scattering (DLS). The efficiency of Fe-IHS complexes in curing Fe chlorosis was compared with two soluble Fe sources (Fe complexed by a water-extractable humic fraction [Fe--WEHS] and the synthetic Fe-chelate, Fe-o,o-EDDHA]) and a poorly soluble mineral form (Ferrihydrite). The IR spectra revealed coordination of the Fe ions with the carboxylic and phenolic groups of the oxygen-rich humic substances. The SEM and DLS data suggest that the Fe-IHS complexes were large aggregated particles with diameter of about 500 to 1500 nm formed by different nanoparticles with very small size ranging from 50 to 200 nm. Chemical extractions show that the Fe pool is dominated by poorly ordered Fe phases strongly associated with IHS. The efficiency of Fe-IHS complexes in curing Fe chlorosis symptoms in Fe-deficient plants has been demonstrated, being, when used in high concentration, comparable to those of Fe-WEHS and Fe-o,oEDDHA. The capability of cucumber (Cucumis sativus L.) plants to acquire Fe from poorly soluble Fe forms was also confirmed when a poorly crystalline radiolabeled Fe-oxide (59 Ferrihydrite) source, used to represent a short-range order Fe phases associated with Fe-IHS complexes, has been used. The results of this work point out the possible use of environmental-friendly Fe insoluble high- molecular weight complexes (Fe-IHS) as an effective product to correct the Fe nutritional disorder. [ABSTRACT FROM AUTHOR]

White lupins ( Lupinus albus L.) respond to phosphate deficiency by producing special root structures called cluster roots. These cluster roots secrete large amounts of carboxylates into the rhizosphere, mostly citrate and malate, which act as phosphate solubilizers and enable the plant to grow in soils with sparingly available phosphate. The success and efficiency of such a P-acquisition strategy strongly depends on the persistence and stability of the carboxylates in the soil, a parameter that is influenced to a large extent by biodegradation through rhizosphere bacteria and fungi. In this study, we show that white lupin roots use several mechanisms to reduce microbial growth. The abundance of bacteria associated with cluster roots was decreased at the mature state of the cluster roots, where a burst of organic acid excretion and a drastic pH decrease is observed. Excretion of phenolic compounds, mainly isoflavonoids, induced fungal sporulation, indicating that vegetative growth, and thus potential citrate consumption, is reduced. In addition, the activity of two antifungal cell wall-degrading enzymes, chitinase and glucanase, were highest at the stage preceding the citrate excretion. Therefore, our results suggest that white lupin has developed a complex strategy to reduce microbial degradation of the phosphate-solubilizing agents. [ABSTRACT FROM AUTHOR]