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
BackgroundThe mechanisms by which nitrate is transported into the roots have been characterized both at physiological and molecular levels. It has been demonstrated that nitrate is taken up in an energy-dependent way by a four-component uptake machinery involving high- and low- affinity transport systems. In contrast very little is known about the physiology of nitrate transport towards different plant tissues and in particular at the leaf level.ResultsThe mechanism of nitrate uptake in leaves of cucumber (Cucumis sativus L. cv. Chinese long) plants was studied and compared with that of the root. Net nitrate uptake by roots of nitrate-depleted cucumber plants proved to be substrate-inducible and biphasic showing a saturable kinetics with a clear linear non saturable component at an anion concentration higher than 2 mM. Nitrate uptake by leaf discs of cucumber plants showed some similarities with that operating in the roots (e.g. electrogenic H+ dependence via involvement of proton pump, a certain degree of induction). However, it did not exhibit typical biphasic kinetics and was characterized by a higher Km with values out of the range usually recorded in roots of several different plant species. The quantity and activity of plasma membrane (PM) H+-ATPase of the vesicles isolated from leaf tissues of nitrate-treated plants for 12 h (peak of nitrate foliar uptake rate) increased with respect to that observed in the vesicles isolated from N-deprived control plants, thus suggesting an involvement of this enzyme in the leaf nitrate uptake process similar to that described in roots. Molecular analyses suggest the involvement of a specific isoform of PM H+-ATPase (CsHA1) and NRT2 transporter (CsNRT2) in root nitrate uptake. At the leaf level, nitrate treatment modulated the expression of CsHA2, highlighting a main putative role of this isogene in the process.ConclusionsObtained results provide for the first time evidence that a saturable and substrate-inducible nitrate uptake mechanism operates in cucumber leaves. Its activity appears to be related to that of PM H+-ATPase activity and in particular to the induction of CsHA2 isoform. However the question about the molecular entity responsible for the transport of nitrate into leaf cells therefore still remains unresolved.
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 A mu M 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 Fe-59 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
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 ferric(III) -chelate reductase (FRO), a Fe(2+) 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.
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