Fe Nutrition Of Plants Research Articles

Silicon ameliorates iron deficiency in sour orange seedlings grown under different pHs

Iron (Fe) is an essential element for all plants while silicon (Si) is non-essential but has some beneficial roles for reducing iron deficiency stresses in plants. Soil pH affect iron solubility and availability and thus plant Fe nutrition however, little is known on how high or low pHs affect Si efficiency in alleviating Fe deficiency stress. In this research the interactive effects of root zone pH and Si nutrition on the alleviation of Fe deficiency in sour orange (Citrus aurantium) seedlings were investigated. Citrus seedlings were grown for three months in Fe-sufficient (control), Fe-deficient and Fe-zero media supplied with or without Si (0.7 mM), adjusted to pHs 6.5 and 8.0 and analyzed subsequently. Fe deficiency resulted in declined tissue Fe, proteins, chlorophylls, carotenoids and the activities of catalase, soluble and cell wall peroxidase associated with increased hydrogen peroxide and lipid peroxidation. Higher pH compromised plant growth associated with depleted tissue Fe in both Fe-deficient and Fe-sufficient plants and imposed chlorophylls and soluble proteins reduction. Si nutrition increased plant growth in Fe-deficient and Fe-sufficient plants under both pH treatments. It also improved tissue Fe content, soluble proteins, chlorophylls, carotenoids and the activities of antioxidant enzymes while decreased tissue hydrogen peroxide and lipid peroxidation levels. These results suggest that Si nutrition can reduce the harmful effects of soil Fe deficiency and higher pHs in sour orange plants via increase of Fe content and reduction of oxidative stress; however, in the absence of Fe, the applied Si cannot mitigate plant nutrient stress symptoms.

Iron Nutrition in Plants: Towards a New Paradigm?

Iron (Fe) is an essential micronutrient for plant growth and development. Fe availability affects crops' productivity and the quality of their derived products and thus human nutrition. Fe is poorly available for plant use since it is mostly present in soils in the form of insoluble oxides/hydroxides, especially at neutral to alkaline pH. How plants cope with low-Fe conditions and acquire Fe from soil has been investigated for decades. Pioneering work highlighted that plants have evolved two different strategies to mine Fe from soils, the so-called Strategy I (Fe reduction strategy) and Strategy II (Fe chelation strategy). Strategy I is employed by non-grass species whereas graminaceous plants utilize Strategy II. Recently, it has emerged that these two strategies are not fully exclusive and that the mechanism used by plants for Fe uptake is directly shaped by the characteristics of the soil on which they grow (e.g., pH, oxygen concentration). In this review, recent findings on plant Fe uptake and the regulation of this process will be summarized and their impact on our understanding of plant Fe nutrition will be discussed.

Similarities and differences in iron homeostasis strategies between graminaceous and nongraminaceous plants.

Iron (Fe) homeostasis is essential for both plant development and human nutrition. The maintenance of Fe homeostasis involves a complex network in which Fe signaling nodes and circuits coordinate tightly Fe transporters, ferric reductases, H+ -ATPases, low-molecular-mass metal chelators, and transporters of chelators and Fe-chelate complexes. Early-stage studies have revealed different strategies for Fe homeostasis between graminaceous and nongraminaceous plants. Recent progress has refreshed our understanding of previous knowledge, especially on the uptake, phloem transport and systemic signaling of Fe. This review attempts to summarize recent exciting and potentially influential studies on the various routes of Fe uptake and distribution in plants, focusing on breakthroughs that have changed our understanding of plant Fe nutrition.

Global analysis of nitrogen fertilization effects on grain zinc and iron of major cereal crops.

Human zinc (Zn) and iron (Fe) deficiencies can partly be alleviated by enhancing cereal concentrations of these micronutrients. Soil nitrogen (N) levels codetermine cereal grain yields and Zn and Fe nutrition of plants and grains. Grain Zn and Fe concentrations have been reported to be affected by both yield dilution and enhanced acquisition and grain allocation of Zn and Fe. A global meta-analysis of 100 publications concerning wheat, maize, and rice providing 785 records of Zn and 506 records of Fe allowed us to assess their relative importance and quantify the concentrations and bioavailability of Zn and Fe in major cereal grains over a wide range of N fertilization levels. Compared with the no N controls, N application significantly increased grain Zn and Fe concentrations in all crops except maize Zn. The increase in grain protein concentration correlated positively with the increases in Zn and Fe concentrations in all cereals except Zn in maize. In rice, the grain Zn and Fe concentration increase was independent of the rate of N applied. Grain concentrations of Zn and Fe in wheat and Fe in maize were positively correlated with N rate but were only higher than those in the controls above 40–60 kg N ha−1. At lower N rates, the dilution effect was thus stronger than the enhancement effect. N supply had a larger effect on Zn and Fe concentrations in loamy textured soils or at lower soil available N and phosphorus (P), or higher soil organic matter and available Zn contents or with P and Zn fertilization, but the effect sizes differed among crops. Reductions in phytic acid concentration after N fertilization occurred in wheat, potentially improving micronutrient bioavailability. Thus, our findings indicate that N fertilization could be managed in ways that simultaneously support high grain yields and enhance nutritional quality for major cereals.

Several Yeast Species Induce Iron Deficiency Responses in Cucumber Plants (Cucumis sativus L.).

Iron (Fe) deficiency is a first-order agronomic problem that causes a significant decrease in crop yield and quality. Paradoxically, Fe is very abundant in most soils, mainly in its oxidized form, but is poorly soluble and with low availability for plants. In order to alleviate this situation, plants develop different morphological and physiological Fe-deficiency responses, mainly in their roots, to facilitate Fe mobilization and acquisition. Even so, Fe fertilizers, mainly Fe chelates, are widely used in modern agriculture, causing environmental problems and increasing the costs of production, due to the high prices of these products. One of the most sustainable and promising alternatives to the use of agrochemicals is the better management of the rhizosphere and the beneficial microbial communities presented there. The main objective of this research has been to evaluate the ability of several yeast species, such as Debaryomyces hansenii, Saccharomyces cerevisiae and Hansenula polymorpha, to induce Fe-deficiency responses in cucumber plants. To date, there are no studies on the roles played by yeasts on the Fe nutrition of plants. Experiments were carried out with cucumber plants grown in a hydroponic growth system. The effects of the three yeast species on some of the most important Fe-deficiency responses developed by dicot (Strategy I) plants, such as enhanced ferric reductase activity and Fe2+ transport, acidification of the rhizosphere, and proliferation of subapical root hairs, were evaluated. The results obtained show the inductive character of the three yeast species, mainly of Debaryomyces hansenii and Hansenula polymorpha, on the Fe-deficiency responses evaluated in this study. This opens a promising line of study on the use of these microorganisms as Fe biofertilizers in a more sustainable and environmentally friendly agriculture.

Plant iron nutrition: the long road from soil to seeds.

Iron (Fe) is an essential plant micronutrient since many cellular processes including photosynthesis, respiration, and the scavenging of reactive oxygen species depend on adequate Fe levels; however, non-complexed Fe ions can be dangerous for cells, as they can act as pro-oxidants. Hence, plants possess a complex homeostatic control system for safely taking up Fe from the soil and transporting it to its various cellular destinations, and for its subcellular compartmentalization. At the end of the plant's life cycle, maturing seeds are loaded with the required amount of Fe needed for germination and early seedling establishment. In this review, we discuss recent findings on how the microbiota in the rhizosphere influence and interact with the strategies adopted by plants to take up iron from the soil. We also focus on the process of seed-loading with Fe, and for crop species we also consider its associated metabolism in wild relatives. These two aspects of plant Fe nutrition may provide promising avenues for a better comprehension of the long pathway of Fe from soil to seeds.

Does the exudation of coumarins from Fe-deficient, soil-grown Brassicaceae species play a significant role in plant Fe nutrition?

The availability of iron (Fe) is highly limited for plant uptake in calcareous soils. To overcome this challenge, plants have developed different strategies of Fe mobilization. Non-gramineous species employ a reduction-based strategy (strategy I), which is assisted by the release of Fe(III)-chelating and -reducing coumarins. In this study, the coumarin release by three strategy-I plant species of the Brassicaceae family, i.e. Brassica napus L., Raphanus sativus L. and Sinapis alba L., on calcareous soil was examined using the RHIZOtest approach. With this experimental setup, the plants were first grown hydroponically in a cylindrical growth unit for 12 days, which was then subsequently brought in contact with calcareous, Fe deficient soil for another 9 days. Root exudates were collected in both experimental phases. Total carbon (C) release, coumarin exudation rates, trace metal plant uptake as well as trace metal mobilization were determined in response to Fe deficiency after 3, 6 and 9 days of soil exposure. High total C exudation rates indicated a clear stress response upon Fe deficiency during the soil contact period. Six different coumarins were detected, with scopoletin being predominantly released by all three plant species during the hydroponic stage with zero Fe supply. The exudation rate of all detected coumarins was significantly lower during soil exposure and represented only a very small fraction of the total organic compounds released. Although the role of coumarins in Fe mobilization is very complex and still not completely understood, it seems that coumarins are less important in Fe mobilization by the three investigated plant species when grown on soil compared to phytosiderophore-releasing strategy-II plants. Our data further indicate that plants growing on a Fe-deficient soil release less coumarins than in an artificial nutrient solution setup with zero Fe supply, which highlights the importance of determining root exudation rates from soil-grown plants, since data from hydroponic studies are likely not representative for soil-grown plants.

Adsorption of humic substances on ferrihydrite affects its use as iron source by plants

Poorly crystalline Fe oxides are sources of Fe to plants. The adsorption of humic substances (HS) on these oxides alters its reactivity and stability in soils, and thus may affect Fe mobilization and uptake by plants from these compounds. This work aimed at studying how the adsorption of HS on Fe oxides affects its use as Fe source by two plant species with different Fe acquisition strategies, white lupin (Strategy I) and wheat (Strategy II). To this end, two completely randomized experiments, one with each plant, were carried out using a calcareous growing media and involving increasing amounts of HS adsorbed on ferrihydrite (0, 16, 60, and 97 mg C g–1) which was used as Fe source. The highest HS rate was the only treatment that significantly increased Fe uptake in wheat relative to control without HS. This was related to a decreased concentration of Fe in poorly crystalline oxides in the growing media. On the contrary, HS did not affect significantly Fe uptake by lupin. However, in this crop, the highest HS rate decreased the concentration of Fe in oxides relative to the lowest HS rate, without significant differences with other treatments. Thus, the effect of adsorbed HS on Fe uptake differed in two plants with different Fe acquisition strategies. The increased Fe uptake in wheat at the highest HS rate can be explained at least in part by an increased Fe mobilization from oxides by plant roots. These findings provide new insights on the role of soil organic matter on plant Fe nutrition.

Application of different foliar iron fertilizers for enhancing the growth and antioxidant capacity of rice and minimizing cadmium accumulation.

Iron (Fe) fertilizer can reduce cadmium (Cd) uptake and toxicity in rice, but the underlying mechanisms of Cd mitigation by different fertilizers are poorly understood. Here, pot experiments in rice were conducted to characterize the effects of four types of foliar-applied Fe fertilizer (chelated ferrous Fe, ferric Fe, ionic ferrous Fe, and ferric Fe) at three doses (20, 50, and 100mgL-1) on photosynthetic capacity, antioxidant ability, yield, and Cd accumulation in Cd-contaminated soil. The results showed that foliar Fe application increased the net photosynthesis rate by 19.3%, peroxidase (POD) by 18.2%, superoxide dismutase (SOD) by 26.9%, and catalase (CAT) by 19.6%, and led to a 7.2% increase in grain yield compared with the control. Moreover, foliar Fe application significantly reduced Cd accumulation by 15.9% in brown rice and decreased the translocation of Cd from roots to other plant tissues. Overall, application of moderate doses (50mgL-1) of chelated ferrous Fe was the most effective method for reducing Cd uptake (decreasing the Cd concentration in brown rice by 29.0%) and toxicity in rice (decreasing malondialdehyde by 23.2% and increasing POD, SOD, and CAT by 54.4%, 51.6%, and 45.7%, respectively), which may stem from the fact that chelated ferrous Fe was a more stable and bioavailable source of Fe for rice. The Cd concentration in rice had negative relationship with Fe concentration, and the translocation of Cd from root to the other tissues was reduced by the higher Fe nutrition status in leaf, suggesting that a high Fe supply may decrease Cd content by inhibiting the expression of the Fe transport system. These results indicate that foliar application of chelated ferrous Fe provides a promising alternative approach for enhancing growth and controlling Cd accumulation in rice plants. Furthermore, these results advance our understanding of the associations between plant Fe nutrition status and Cd accumulation.

Iron Deficiency Prolongs Seed Dormancy in Arabidopsis Plants.

The understanding of seed dormancy, germination and longevity are important goals in plant biology, with relevant applications for agriculture, food industry and also human nutrition. Reactive Oxygen Species (ROS) are key molecules involved in the release of dormancy, when their concentrations fall within the so called ‘oxidative window.’ The mechanisms of ROS distribution and sensing in seeds, from dormant to germinating ones, still need elucidation. Also, the impact of iron (Fe) deficiency on seed dormancy is still unexplored; this is surprising, given the known pro-oxidant role of Fe when in a free form. We provide evidence of a link between plant Fe nutrition and dormancy of progeny seeds by using different Arabidopsis ecotypes and mutants with different dormancy strengths grown in control soil or under severe Fe deficiency. The latter condition extends the dormancy in several genotypes. The focus on the mechanisms involved in the Fe deficiency-dependent alteration of dormancy and longevity promises to be a key issue in seed (redox) biology.

Synthesis of [13C4]-labeled 2'-deoxymugineic acid.

The phytosiderophore 2'-deoxymugineic acid (DMA) is exuded via the root system by all grasses (including important crop plants like rice, wheat and barley) to mobilize Fe(III) from soil and improve plant Fe nutrition, crucial for high crop yields. Elucidation of the biogeochemistry of 2'-deoxymugineic acid in the rhizosphere requires its quantification in minute amounts. To this end, (13)C4-DMA was synthesized for the first time, from cheap isotopically labeled starting materials. The synthetic route utilizes L-allyl((13)C2)glycine and L-(2-(13)C)azetidine ((13)C)carboxylic acid as versatile labeled building blocks. The title compound was recently used as an internal standard for analysis of soil and plant samples allowing the first accurate quantification of DMA in these matrices by means of LC-MS/MS. It is furthermore used in tracer experiments investigating biodegradation of DMA in soil.

Involvement of the ABCG37 transporter in secretion of scopoletin and derivatives by Arabidopsis roots in response to iron deficiency

Studies of Iron (Fe) uptake mechanisms by plant roots have focussed on Fe(III)-siderophores or Fe(II) transport systems. Iron deficency also enhances root secretion of flavins and phenolics. However, the nature of these compounds, their transport outside the roots and their role in Fe nutrition are largely unknown. We used HPLC/ESI-MS (TOF) and HPLC/ESI-MS/MS (ion trap) to characterize fluorescent phenolic-type compounds accumulated in roots or exported to the culture medium of Arabidopsis plants in response to Fe deficiency. Wild-type and mutant plants altered either in phenylpropanoid biosynthesis or in the ABCG37 (PDR9) ABC transporter were grown under standard or Fe-deficient nutrition conditions and compared. Fe deficiency upregulates the expression of genes encoding enzymes of the phenylpropanoid pathway and leads to the synthesis and secretion of phenolic compounds belonging to the coumarin family. The ABCG37 gene is also upregulated in response to Fe deficiency and coumarin export is impaired in pdr9 mutant plants. Therefore it can be concluded that: Fe deficiency induces the secretion of coumarin compounds by Arabidopsis roots; the ABCG37 ABC transporter is required for this secretion to take place; and these compounds improved plant Fe nutrition.

Effects of different Fe supplies on mineral partitioning and remobilization during the reproductive development of rice (Oryza sativa L.).

BackgroundMinimal information exists on whole-plant dynamics of mineral flow through rice plants and on the source tissues responsible for mineral export to developing seeds. Understanding these phenomena in a model plant could help in the development of nutritionally enhanced crop cultivars. A whole-plant accumulation study, using harvests during reproductive development under different Fe supplies, was conducted to characterize mineral accumulation in roots, non-flag leaves, flag leaves, stems/sheaths, and panicles of Kitaake rice plants.ResultsLow Fe supply promoted higher accumulation of Zn, Cu and Ni in roots, Mn, Ca, Mg and K in leaves and Zn in stems/sheaths and a smaller accumulation of Fe, Mn and Ca in roots and Zn and Ni in leaves. High Fe supply promoted higher accumulation of Fe in roots and Zn in leaves and a smaller accumulation of Fe in leaves and stems/sheaths and Zn, Cu and K in roots. Correlation analyzes indicated that fluctuations in Mn-Ca, Zn-Cu, Zn-Ni, Cu-Ni, Mo-S, Ca-Mg, Cu-Mn and Cu-Mg concentrations in response to different Fe supplies were positively correlated in at least four of the five organs analyzed.ConclusionsMineral content loss analysis indicated that mineral remobilization from vegetative organs can occur in rice plants; however, for seeds to acquire minerals, vegetative remobilization is not absolutely required. Also, mineral remobilization from vegetative tissues in rice was greatly dependent of plant Fe nutrition. Remobilization was observed for several minerals from flag leaves and stems/sheaths, but the amounts were generally far below the total mineral accretion observed in panicles, suggesting that continued uptake and translocation of minerals from the roots during seed fill are probably more important than mineral remobilization.Electronic supplementary materialThe online version of this article (doi:10.1186/1939-8433-5-27) contains supplementary material, which is available to authorized users.

Identification and characterization of the major mitochondrial Fe transporter in rice

The uptake, translocation, and compartmentalization of Fe are essential for plant cell function and life cycle. Despite rapid progress in our understanding of Fe homeostasis in plants, Fe transport from the cytoplasm to mitochondria was, until recently, poorly understood. The screening of 3,993 mutant lines for symptoms of Fe deficiency resulted in the identification and characterization of a major mitochondrial Fe transporter (MIT) in rice. MIT was found to localize to mitochondria and to complement the growth of a yeast strain defective in mitochondrial Fe transport. The knock-out of MIT resulted in a lethal phenotype, and in knock-down plants, several agronomic characteristics were compromised, such as plant height, average number of tillers, days to flower, fertility, and yield. Changes in the expression of genes involved in Fe transport suggested a disturbance of cellular Fe transport. Furthermore, the mitochondrial Fe concentration and the activity of the mitochondrial Fe-S enzyme aconitase were significantly reduced compared with wild-type plants. The identification of MIT is a significant advance in the field of plant Fe nutrition and should facilitate the cloning of paralogs from other plant species.

Plant Fe status affects the composition of siderophore-secreting microbes in the rhizosphere

Soil microbes have been demonstrated to play an important role in favouring plant iron (Fe) uptake under Fe-limiting conditions. However, the mechanisms involved are still unclear. This present study reported the effects of plant Fe status on the composition of siderophore-secreting microbes in the rhizosphere, and their potential function in improving plant Fe nutrition. An Fe-efficient plant, red clover (Trifolium pratense 'Kenland') was cultured in a calcareous soil to obtain rhizosphere soils with (Fe-sufficient) or without (Fe-stressed) foliar FeEDTA spraying. The siderophore-producing ability of rhizospheric microbes was measured. The bioavailability of the siderophore-solubilized Fe from iron oxides/hydroxides was tested in hydroponic culture. In rhizosphere soil, the number of microbes that secreted siderophores quickly was more in the Fe-stressed treatment than in the Fe-sufficient one, while the number of microbes that did not secret siderophores was the opposite. A significantly higher concentration of phenolics was detected in the rhizosphere soil of Fe-stressed plants. Moreover, after the soil was incubated with phenolic root exudates, the composition of the siderophore-secreting microbial community was similar with that of the rhizosphere of Fe-stressed plant. Additionally, the siderophores produced by a rhizospheric microbe isolated from the Fe-stressed treatment can well solubilize iron oxides/hydroxides, and the utilization of the siderophore-solubilized Fe by plant was even more efficient than EDTA-Fe. Iron-deficiency stress of red clover would alter the composition of siderophore-secreting microbes in the rhizosphere, which is probably due to the phenolics secretion of the root, and may in turn help to improve the solubility of Fe in soils and plant Fe nutrition via elevated microbial siderophore secretion.

Effects of humic substances on iron nutrition of lupin

Poorly crystalline Fe oxides and organic matter are two important factors affecting Fe nutrition of plants. The main objective of this work was to study the contribution of humic substances to Fe nutrition of a typical Fe-chlorosis sensitive plant (white lupin, Lupinus albus L.). An experiment was performed involving two growing media (siliceous and calcareous) and different Fe sources: control without additional Fe added to the growing media, ferrihydrite (FH), FH + humic substances (HS, at two rates, 0.1 and 0.3 g C kg−1 growing media), Fe complexed to humic substances (Fe–HS), and Fe–EDDHA (as an effective Fe source in calcareous media). Chlorophyll meter readings and dry matter production (DM) were significantly greater with Fe–EDDHA and Fe–HS when compared with the other treatments in calcareous media. The positive effect of complexed Fe (to EDDHA or HS) on Fe nutrition can be, at least partially, related to an increase in Fe(III)-reducing capacity by roots, which seems to be improved by an enhanced root development. No positive effect on Fe nutrition was observed with the application of HS in a ferrihydrite enriched growing media (FH + HS) at 4 weeks, particularly with the application of HS at 0.3 g C kg−1 in calcareous media. Thus, the effect of HS on Fe availability was only positive if some Fe is complexed to HS. The efficiency of HS with Fe complexed (Fe–HS) in preventing Fe chlorosis was found to be similar to Fe–EDDHA. This is important not only for the knowledge of factors affecting Fe availability in soils, but also with a view of using Fe–HS complexes as effective products to correct this nutritional problem.

Iron Deficiency-Induced Secretion of Phenolics Facilitates the Reutilization of Root Apoplastic Iron in Red Clover

Phenolic compounds are frequently reported to be the main components of root exudates in response to iron (Fe) deficiency in Strategy I plants, but relatively little is known about their function. Here, we show that removal of secreted phenolics from the root-bathing solution almost completely inhibited the reutilization of apoplastic Fe in roots of red clover (Trifolium pratense). This resulted in much lower levels of shoot Fe and significantly higher root Fe compared with control and also resulted in leaf chlorosis, suggesting this approach stimulated Fe deficiency. This was supported by the observation that phenolic removal significantly enhanced root ferric chelate reductase activity, which is normally induced by plant Fe deficiency. Furthermore, root proton extrusion, which also is normally increased during Fe deficiency, was found to be higher in plants exposed to the phenolic removal treatment too. These results indicate that Fe deficiency-induced phenolics secretion plays an important role in the reutilization of root apoplastic Fe, and this reutilization is not mediated by proton extrusion or the root ferric chelate reductase. In vitro studies with extracted root cell walls further demonstrate that excreted phenolics efficiently desorbed a significant amount of Fe from cell walls, indicating a direct involvement of phenolics in Fe remobilization. All of these results constitute the first direct experimental evidence, to our knowledge, that Fe deficiency-induced secretion of phenolics by the roots of a dicot species improves plant Fe nutrition by enhancing reutilization of apoplastic Fe, thereby improving Fe nutrition in the shoot.

Ammonium improves iron nutrition by decreasing leaf apoplastic pH of sunflower plants (Helianthus annuus L. cv. Frankasol)

The effect of nitrogen form on pH and concentration of soluble iron (Fe) in leaf apoplast was investigated in hydrophonically grown sunflower plants (Helianthus annuus L. cv. Frankasol), and the mechanism underlying the improved Fe nutrition by ammonium (NH4) supply was also elucidated. Ammonium supply ameliorated Fe nutrition of plants grown without Fe through decreasing apoplastic pH and increasing soluble Fe concentration in apoplastic fluid of young leaves. The soluble Fe concentration in apoplastic fluid and cell sap of young leaves, and xylem exudates of NH4 fed-plants was higher than that of nitrate (NO3) fed-plants, and no typical Fe-deficiency chlorosis in young leaves was observed in NH4, fed plant without Fe supply. The apoplastic pH was 6.15 and 5.94 in young leaves of Fe-deficient plants fed respectively with NO3 and NH4, while in Fe-sufficient plants, the apoplastic pH was 6.43 with NO3, and 5.50 with NH4 supply. In primary leaves, the apoplastic pH was around 6.25 irrespective of nitrogen form and Fe supply. The pH of xylem exudate was 5.72 in Fe-deficient plants fed with NO3 and 5.49 with NH4. Iron nutrition increased the pH of xylem exudate by 0.27 and 0.16 unit under NO3 and NH4 supply respectively.

Reduction and transport of Fe from siderophores

Soils contain siderophores produced by bacteria and fungi; however, the role of siderophores in Fe nutrition of plants is uncertain. The Strategy I plant cucumber (Cucumis sativus L.) was used in an investigation of ferric chelate reduction activity and uptake and transport of Fe from ferric hydroxyethylethylenetriacetic acid (FeHEDTA) and ferric N,N′–di–(2–hydroxybenzoyl)–ethylenediamine– N,N′-diacetic acid (FeHBED) and the hydroxamate siderophores, ferric rhodotorulic acid (FeRA) and ferric ferrioxime B (FeFOB). Cucumber seedlings were grown in a hydroponic medium without Fe or supplied with 10 μM FeHEDTA. Iron-deficient cucumber roots readily reduced FeHEDTA, while Fe-sufficient roots had low levels of ferric chelate reduction activity. The siderophore FeRA was reduced by Fe-deficient roots at 8% of the rate of FeHEDTA, while FeFOB was not reduced. The highly stable synthetic chelate FeHBED was reduced at 16% the rate of FeHEDTA. Fe transport to shoots by Fe-deficient seedlings from the slowly reducible complexes 59FeRA and 59FeHBED was, respectively, 74% and 73% of that transported from 59FeHEDTA. The ferrous complexing agent, bathophenanthrolinedisulfonic acid (BPDS), had a strong inhibitory effect on uptake and transport of Fe from 59FeHEDTA or 59FeRA into shoots. An average of 11% as much Fe was transported to shoots of Fe-deficient seedlings from 59FeFOB as from 59FeHEDTA. Neither the Fe nutritional status of the seedlings nor the presence of BPDS influenced the uptake and transport of Fe from 59FeFOB. It is concluded that cucumber roots may take up substantial amounts of Fe from FeRA and FeHBED following reduction, while small amounts of Fe may be taken up from FeFOB by a mechanism not involving reduction of the ferric siderophore at the root surface.

Fusarinines and dimerum acid, mono- and dihydroxamate siderophores from Penicillium chrysogenum, improve iron utilization by strategy I and strategy II plants.

Cucumber, as a strategy I plant, and Maize as a strategy II plant, were cultivated in hydroponic culture in the presence of a ferrated siderophore mixture (1 microM) from a culture of Penicillium chrysogenum isolated from soil. The siderophore mixture significantly improved the iron status of these plants as measured by chlorophyll concentration to the same degree as a 100-fold higher FeEDTA supply. Analysis of the siderophore mixture from P. chrysogenum by HPLC and electrospray mass spectrometry revealed that besides the trihydroxamates, coprogen and ferricrocin, large amounts of dimerum acid and fusarinines were present which represent precursor siderophores or breakdown products of coprogen. In order to prove the iron donor properties of dimerum acid and fusarinines for plants, purified coprogen was hydrolyzed with ammonia and the hydrolysis products consisting of dimerum acid and fusarinine were used for iron uptake by cucumber and maize. In short term experiments radioactive iron uptake and translocation rates were determined using ferrioxamine B, coprogen and hydrolysis products of coprogen. While the trihydroxamates revealed negligible or intermediate iron uptake rates by both plant species, the fungal siderophore mixture and the ammoniacal hydrolysis products of coprogen showed high iron uptake, suggesting that dimerum acid and fusarinines are very efficient iron sources for plants. Iron reduction assays using cucumber roots or ascorbic acid also showed that iron bound to hydrolysis products of coprogen was more easily reduced compared to iron bound to trihydroxamates. Ligand exchange studies with epi-hydroxymugineic acid and EDTA showed that iron was easily exchanged between coprogen hydrolysis products and phytosiderophores or EDTA. The results indicate that coprogen hydrolysis products are an excellent source for Fe nutrition of plants.