Does high bicarbonate supply to roots change availability of iron in the leaf apoplast?

The role of the leaf apoplast in iron (Fe) uptake into the leaf symplast is insufficiently understood, particularly in relation to the supposed inactivation of Fe in leaves caused by elevated bicarbonate in calcareous soils. It has been supposed that high bicarbonate supply to roots increases the pH of the leaf apoplast which decreases the physiological availability of Fe in leaf tissues. The study reported here has been carried out with sunflower plants grown in nutrient solution and with grapevine plants grown on calcareous soil under field conditions. The data obtained clearly show that the pH of the leaf apoplastic fluid was not affected by high bicarbonate supply in the root medium (nutrient solution and field experiments). The concentrations of total, symplastic and apoplastic Fe were decreased in chlorotic leaves of both sunflower (nutrient solution experiment) and grapevine plants in which leaf expansion was slightly inhibited (field experiment). However, in grapevine showing severe inhibition of leaf growth, total Fe concentration in chlorotic leaves was the same or even higher than in green ones, indicative to the so-called `chlorosis paradox'. The findings do not support the hypothesis of Fe inactivation in the leaf apoplast as the cause of Fe deficiency chlorosis since no increase was found in the relative amount of apoplastic Fe (% of total leaf Fe) either in the leaves of sunflower or grapevine plants. It is concluded that high bicarbonate concentration in the soil solution does not decrease Fe availability in the leaf apoplast.     

Iron availability in plant tissues-iron chlorosis on calcareous soils

The article describes factors and processes which lead to Fe chlorosis (lime chlorosis) in plants grown on calcareous soils. Such soils may contain high HCO₃ ^- concentrations in their soil solution, they are characterized by a high pH, and they rather tend to accumulate nitrate than ammonium because due to the high pH level ammonium nitrogen is rapidly nitrified and/or even may escape in form of volatile NH₃. Hence in these soils plant roots may be exposed to high nitrate and high bicarbonate concentrations. Both anion species are involved in the induction of Fe chlorosis.Physiological processes involved in Fe chlorosis occur in the roots and in the leaves. Even on calcareous soils and even in plants with chlorosis the Fe concentration in the roots is several times higher than the Fe concentration in the leaves. This shows that the Fe availability in the soil is not the critical process leading to chlorosis but rather the Fe uptake from the root apoplast into the cytosol of root cells. This situation applies to dicots as well as to monocots. Iron transport across the plasmamembrane is initiated by Fe^III reduction brought about by a plasmalemma located Fe^III reductase. Its activity is pH dependent and at alkaline pH supposed to be much depressed. Bicarbonate present in the root apoplast will neutralize the protons pumped out of the cytosol and together with nitrate which is taken up by a H⁺/nitrate cotransport high pH levels are provided which hamper or even block the Fe^III reduction.Frequently chlorotic leaves have higher Fe concentrations than green ones which phenomenon shows that chlorosis on calcareous soils is not only related to Fe uptake by roots and Fe translocation from the roots to the upper plant parts but also dependent on the efficiency of Fe in the leaves. It is hypothesized that also in the leaves Fe^III reduction and Fe uptake from the apoplast into the cytosol is affected by nitrate and bicarbonate in an analogous way as this is the case in the roots. This assumption was confirmed by the highly significant negative correlation between the leaf apoplast pH and the degree of iron chlorosis measured as leaf chlorophyll concentration. Depressing leaf apoplast pH by simply spraying chlorotic leaves with an acid led to a regreening of the leaves.     

Nitrate does not result in iron inactivation in the apoplast of sunflower leaves

It has been hypothesized that nitrate (NO(3)(-)) nutrition might induce iron (Fe) deficiency chlorosis by inactivation of Fe in the leaf apoplast (H.U. Kosegarten, B. Hoffmann, K. Mengel [1999] Plant Physiol 121: 1069-1079). To test this hypothesis, sunflower (Helianthus annuus L. cv Farnkasol) plants were grown in nutrient solutions supplied with various nitrogen (N) forms (NO(3)(-), NH(4)(+) and NH(4)NO(3)), with or without pH control by using pH buffers [2-(N-morpholino)ethanesulfonic acid or 4-(2-hydroxyethyl)-1-piperazineethanesulfonic acid]. It was shown that high pH in the nutrient solution restricted uptake and shoot translocation of Fe independently of N form and, therefore, induced Fe deficiency chlorosis at low Fe supply [1 micro M ferric ethylenediaminedi(O-hydroxyphenylacetic acid)]. Root NO(3)(-) supply (up to 40 mM) did not affect the relative distribution of Fe between leaf apoplast and symplast at constant low external pH of the root medium. Although perfusion of high pH-buffered solution (7.0) into the leaf apoplast restricted (59)Fe uptake rate as compared with low apoplastic solution pH (5.0 and 6.0, respectively), loading of NO(3)(-) (6 mM) showed no effect on (59)Fe uptake by the symplast of leaf cells. However, high light intensity strongly increased (59)Fe uptake, independently of apoplastic pH or of the presence of NO(3)(-) in the apoplastic solution. Finally, there are no indications in the present study that NO(3)(-) supply to roots results in the postulated inactivation of Fe in the leaf apoplast. It is concluded that NO(3)(-) nutrition results in Fe deficiency chlorosis exclusively by inhibited Fe acquisition by roots due to high pH at the root surface.     

Apoplastic pH and FeIII reduction in young sunflower (Helianthus annuus) roots

The relationship between the apoplastic pH in young sunflower roots ( Helianthus annuus L.) and the plasmalemma ferric chelate reductase (FC-R; EC 1.16.1.7) activity in roots was investigated. The hypothesis was tested that a high apoplastic pH depresses FC-R activity, thereby restricting the uptake of Fe²⁺ into the cytosol. Until recently, little has been known about this relationship, because pH and redox reaction measurements are difficult to perform within the confines of the root apoplast. We recorded the apoplastic pH by means of the fluorescence ratio in conjunction with video microscopy by covalently tagging fluorescein boronic acid to OH groups of the root cell wall. Fe^III reduction was measured using a similar approach by tagging ferrozine diboronic acid with OH groups of the cell wall. Ferrozine forms an Fe²⁺ complex, thus indicating the reduction of ferric iron. In roots bathing in buffered outer solutions of different pH, a high pH sensitivity of apoplastic Fe^III reduction was found, with the highest ferric iron reduction rates at an apoplastic pH of 4.9; above an apoplastic pH of 5.3, no reduction was observed. Nitrate in the bathing solution increased the apoplastic pH and hence depressed the Fe^III reduction; ammonium had the reverse effect. Nitrate together with HCO₃^–, a combination which is typical of calcareous soils, had the strongest depressing effect. From the results, it can be concluded that the main reason for the frequently occurring iron deficiency chlorosis of plants grown on calcareous soils is the inhibition of Fe^III reduction in the apoplast, and hence Fe²⁺ uptake into the cytosol.     

A comparison of some populations of Eucalyptus viminalis Labill. growing on calcareous and acid soils in Victoria,Australia

A study of populations of Eucalyptus viminalis found on both acid and alkaline soils showed that seedlings differ in their tolerance of calcareous soils and their susceptibility to lime chlorosis. Seedlings from an open-forest population on calcareous dunes in the Otway Ranges, Victoria, averaged a significantly greater yield than seedlings of three other populations found on acid soils when all were grown on calcareous soil of pH 6.8. A tall ribbon gum form of E. viminalis, from Paradise gully in the Otways, was the least tolerant of alkaline conditions and showed severe signs of chlorosis and reddening of leaf margins. The latter appeared to be related to high uptake of phosphorus. This population showed rapid growth on a fertile acid loam. The data presented also emphasize the need to consider the variation in soils which can occur at any one site. Population differences were most marked when seedlings were grown on soils of pH near 7.0. Yield was reduced in the Otways calcareous population when seedlings were grown on the more leached soil from swales (pH 6.5) between dunes and on the highly calcareous soil from the dune crests (pH 7.8–8.0). The failure of any of the populations of E. viminalis studied to grow well on soils of high pH suggest that this species is not tolerant of highly calcareous soils. This may in part explain the absence of this species from the drier calcareous areas on the Mornington Peninsula and the Yanakie Isthmus, Victoria. Since seedlings had reduced root development on the soil collected from the Peninsula, E. viminalis may be restricted, in effect because of drought stress, to non-calcareous sites in the eastern part of Victoria.     

High apoplastic solute concentrations in leaves alter water relations of the halophytic shrub, Sarcobatus vermiculatus

Predawn plant water potential (Psi(w)) is used to estimate soil moisture available to plants because plants are expected to equilibrate with the root-zone Psi(w). Although this equilibrium assumption provides the basis for interpreting many physiological and ecological parameters, much work suggests predawn plant Psi(w) is often more negative than root-zone soil Psi(w). For many halophytes even when soils are well-watered and night-time shoot and root water loss eliminated, predawn disequilibrium (PDD) between leaf and soil Psi(w) can exceed 0.5 MPa. A model halophyte, Sarcobatus vermiculatus, was used to test the predictions that low predawn solute potential (Psi(s)) in the leaf apoplast is a major mechanism driving PDD and that low Psi(s) is due to high Na+ and K+ concentrations in the leaf apoplast. Measurements of leaf cell turgor (Psi(p)) and solute potential (Psi(s)) of plants grown under a range of soil salinities demonstrated that predawn symplast Psi(w) was 1.7 to 2.1 MPa more negative than predawn xylem Psi(w), indicating a significant negative apoplastic Psi(s). Measurements on isolated apoplastic fluid indicated that Na+ concentrations in the leaf apoplast ranged from 80 to 230 mM, depending on salinity, while apoplastic K+ remained around 50 mM. The water relations measurements suggest that without a low apoplastic Psi(s), predawn Psi(p) may reach pressures that could cause cell damage. It is proposed that low predawn apoplastic Psi(s) may be an efficient way to regulate Psi(p) in plants that accumulate high concentrations of osmotica or when plants are subject to fluctuating patterns of soil water availability.     

Interactions Between Nitrogen and Manganese Nutrition of Barley Genotypes Differing in Manganese Efficiency

Ammonium-fed plants may acidify the rhizosphere and thus increaseavailability of Mn in calcareous alkaline soils. The importanceof N nutrition in the differential expression of tolerance toMn deficiency among cereal genotypes is not yet clear. Two factorialexperiments testing effects of the NH₄-N/NO₃-N ratio and Mnfertilization on growth of barley genotypes differing in toleranceto Mn deficiency were conducted in two calcareous alkaline soilsin pots in a controlled environment. In the soil containing80% CaCO₃at pH 8.5, better root and shoot growth and highershoot Mn concentrations were achieved with nitrate supply, especiallyat lower rates of Mn fertilization. The Mn-efficient genotypeWeeah (tolerant of Mn deficiency) achieved better root and shootgrowth than Mn-inefficient Galleon barley (sensitive to Mn deficiency)regardless of experimental treatment. Fertilization with Mndid not influence total N concentration in barley roots andshoots. In the soil containing 5% CaCO₃at pH 7.8, ammonium-fedplants had better root and shoot growth and, at shoot Mn concentrationsabove the critical level, Mn-inefficient Galleon performed betterthan Mn-efficient Weeah barley. It appears that differentialexpression of Mn efficiency among barley genotypes is not associatedwith differences in Mn availability expected to be producedby differential rhizosphere acidification as a response to differentforms of N supply. There is an apparent preference of locallyselected barley genotypes for nitrate nutrition when grown onthe highly calcareous alkaline soils of southern Australia. Ammonium; calcareous soil; Hordeum vulgare ; manganese; nitrate; nitrogen form; nutrient efficiency; rhizosphere     

Casparian strip development and its potential function in salt tolerance

The root system is particularly affected by unfavourable conditions because it is in direct contact with the soil environment. Casparian strips, a specialised structure deposited in anticlinal walls, are characterised by the impregnation of the primary wall pores with lignin and suberin. The Casparian strips in the endo- and exodermis of vascular plant roots appear to play an important role in preventing the non-selective apoplastic bypass of salts into the stele along the apoplast under salt stress. However, only a few investigations have examined the deposition and function of these apoplastic barriers in response to salt stress in higher plants.     

Copper concentration in plants and in the rhizosphere as influenced by the iron status of tomato (Lycopersicon esculentum L.)

Changes of metal concentration that occur in the rhizosphere may arise from several processes including variation in the concentration of complexing ligands, pH or redox potential that can be influenced by the Fe status of the plant. The aim of this study was to assess for both acidic and calcareous, Cu-contaminated soils how Cu concentration in plants and in the rhizosphere was affected by the Fe status of a strategy I plant species. The change of soil solution pH, total solution Cu concentration and soil redox potential was monitored for 8 days in the rhizosphere of tomato (Lycopersicon esculentum L.) in response to contrasting Fe supply. The concentration of Cu in roots was enhanced under Fe deficiency in the acidic soils. Shoot Cu however did not vary with the Fe status of the plant. The plant Fe status had little effect on rhizosphere pH, redox potential or Cu concentration in solution in either acidic or calcareous soils. Marked differences in pH and solution Cu concentration were observed between rhizosphere and uncropped soils. Roots induced an increase in pH of acidic soils and a decrease in solution Cu concentration in all soils. The decrease in solution Cu concentration in acidic soils may be explained by the increase in rhizosphere pH. The proposed device provided new data on the fate of Cu in the rhizosphere and showed a positive correlation for the four soils considered together between the total Cu concentration in soil solution and root Cu concentration.     

Overcoming Fe deficiency by a transgenic approach in rice

Iron (Fe) is an essential microelement for plant growth. Fe availability is particularly limited on calcareous soils, which have high pH. Approximately 30% of the world's soils are considered calcareous with low Fe availability, which results in extensive areas of Fe deficiency in plants. Some graminaceous plants are known to secrete high amounts of mugineic acid family phytosiderophores (MAs) under Fe deficiency. This Fe acquisition system is called the Strategy-II mechanism. Tolerance to Fe deficiency in graminaceous plants is thought to depend on the quantity of MAs secreted by plants under Fe deficiency stress. This system was utilized to enhance the tolerance of rice to low Fe availability. Transgenic rice expressing the barley naat genes, one of the genes for the enzymes on the biosynthetic pathway of MAs, showed tolerance to low Fe availability when grown in a calcareous soil.     

Cd accumulation in roots and shoots of durum wheat: the roles of transpiration rate and apoplastic bypass

Cadmium is readily taken up from soils by plants, depending on soil chemistry, and variably among species and cultivars; altered transpiration and xylem transport and/or translocation in the phloem could cause this variation in Cd accumulation, some degree of which is heritable. Using Triticum turgidum var. durum cvs Kyle and Arcola (high and low grain Cd accumulating, respectively), the objectives of this study were to determine if low-concentration Cd exposure alters transpiration, to relate transpiration to accumulation of Cd in roots and shoots at several life stages, and to evaluate the role of apoplastic bypass in the accumulation of Cd in shoots. The low abundance isotope (106)Cd was used to probe Cd translocation in plants which had been exposed to elemental Cd or were Cd-na?ve; apoplastic bypass was monitored using the fluorescent dye PTS (8-hydroxy-1,3,6-pyrenetrisulphonate). Differential accumulation of Cd by 'Kyle' and 'Arcola' could be partially attributed to the effect of Cd on transpiration, as exposure to low concentrations of Cd increased mass flow and concomitant Cd accumulation in 'Kyle'. Distinct from this, exposure of 'Arcola' to low concentrations of Cd reduced translocation of Cd from roots to shoots relative to root accumulation of Cd. It is possible, but not tested here, that sequestration mechanisms (such as phytochelatin production, as demonstrated by others) are the genetically controlled difference between these two cultivars that results in differential Cd accumulation. These results also suggest that apoplastic bypass was not a major pathway of Cd transport from the root to the shoot in these plants, and that most of the shoot Cd resulted from uptake into the stele of the root via the symplastic pathway.     

The effect of high pH and P on the development of lime-chlorosis in two seedling populations ofEucalyptus obliqua L'Hérit

Summary Glasshouse experiments have shown that the application of an acidulating agent to a calcareous soil can increase growth and alleviate severe chlorosis in an acidic population ofE. obliqua. In contrast, a calcareous population showed only a slight response to this treatment and maintained adequate growth and a low frequency of chlorosis on both control and treated calcareous soils. Foliar analyses of seedlings of the acidic population showed that alleviation of chlorosis was concomitant with a reduction in the levels of P, Ca and K, and an increase in uptake of Fe. However, the total Fe content of foliage was poorly correlated with the occurrence of severe chlorosis. Although this evidence suggested that the differential susceptibility ofE. obliqua to lime-chlorosis can be reduced by increasing the availability of Fe, the greater concentration of Fe in chlorotic seedlings indicated that lime-chlorosis may also be due to an inactivation of Fe within the plant (i.e. by P). This hypothesis was partly confirmed by a water culture experiment which showed that a combination of relatively high pH and high external levels of P could induce severe chlorosis in seedlings of the acidic population. In contrast, it appears that the calcareous population has a more efficient mechanism for absorbing Fe and holding it in an available form, even when external concentrations of P are high. It is suggested that plants which have an efficient mechanism for the uptake of Fe at relatively high pH and are less susceptible to the detrimental effects of P have been selected for on these alkaline calcareous soils.     

Fe Accumulation and Mobilization in Root Apoplast of Soybean Seedlings Under Fe deficiency Condition

Fe accumulation and mobilization in root apoplast of soybean ( Clycine max (L.) Merr. ) seedlings grown under Fe supplement of 10-6 mol/L FeEDTA, or Fe-deficient condition were studied. Under Fe sufficient condition, the content of the root apoplastic Fe pools of soybean seedlings changed in a rhythmic fashion of alternative accumulation and mobilization in every three-day cycling, together with orchestrated rhythmic changes of root Fe ( Ⅲ )-reduction capacity and peroxidase activity. In Fe deficient condition, Fe content in the root apoplastic pools deceased steadily almost to nil, and the root Ye( Ⅲ )-reduction capacity and peroxidase activity, ahhrough under went rhythmic changes but only in 2-day cycling. The results showed an important relation among root apoplastic Ye pools, Fe( Ⅲ )-reduction capacity and peroxidase activity.     

Seed treatment with growth regulators and crop productivity. I. 2,4-D as an inducer of salinity-tolerance in wheat (Triticum aestivum L.)

The paper investigates how the apoplastic route of ion transfer is affected by the outermost cortex cell layers of a primary root. Staining of hand-made cross sections with aniline blue in combination with berberine sulfate demonstrated the presence of casparian bands in the endo- and exodermis, potentially being responsible for hindering apoplastic ion movement. The use of the apoplastic dye Evan's Blue allowed viewing under a light microscope of potential sites of uncontrolled solute entry into the apoplast of the root cortex which mainly consisted of injured rhizodermis and/or exodermis cells. The distribution of the dye after staining was highly comparable to EDX analyses on freeze-dried cryosectioned roots. Here, we used Rb+ as a tracer for K+ in a short-time application on selected regions of intact roots from intact plants. After subsequent quench-freezing with liquid propane the distribution of K+ and Rb+ in cell walls was detected on freeze-dried cryosections by their specific X-rays resulting from the incident electrons in a SEM. All such attempts led to a single conclusion, namely, that the walls of the two outermost living cell sheaths of the cortex largely restrict passive solute movements into the apoplast. The ring of turgescent living rhizodermis cells in the root tip region forms the first barrier. With increasing distance to the root tip, in the course of their maturation resp. degradation, this particular function of the rhizodermis cells is replaced by the hypodermis resp. exodermis. Furthermore, the restriction of apoplastic ion flow by the outermost cortex cell layers is rather effective but not complete. Thus, the solute transfer into the stele is mainly restricted by the casparian bands of the endodermis. The overall conclusion is that the resistances of the rhizodermis and exodermis are additive to the endodermis in their role of regulating the apoplastic solute movement across roots. This revised version was published online in August 2006 with corrections to the Cover Date.     

Effects of soil temperature on root and shoot growth traits and iron deficiency chlorosis in sorghum genotypes grown on a low iron calcareous soil

Iron deficiency chlorosis (FeDC) is a common disorder for sorghum [Sorghum bicolor (L.) Moench] grown on alkaline calcareous soils. Four sorghum genotypes were grown in growth chambers on a low Fe (1.3 g/g DTPA-extractable), alkaline (pH 8.0), calcareous (3.87% CaCO₃ equivalent) Aridic Haplustoll to determine effects of different soil temperatures (12, 17, 22 and 27°C at a constant 27°C air temperature) on various root and shoot growth traits and development of FeDC. As soil temperature increased, leaf chlorosis became more severe, and shoot and root dry weights, root lengths, and leaf areas increased markedly. Shoot/root ratios, shoot weight/root length, leaf area/shoot weight and leaf area/root weight and root length also increased while root length/root weight decreased as soil temperature increased. Severe FeDC developed in all genotypes even though genotypes had previously shown different degrees of resistance to FeDC. Genotypes differed in most growth traits, especially dry matter yields, root lengths, and leaf areas, but most traits did not appear to be related to genotype resistance to FeDC. The most FeDC resistant genotype had the slowest growth rate and this may be a mechanism for its greater resistance to FeDC.     

LaALMT1 mediates malate release from phosphorus-deficient white lupin root tips and metal root to shoot translocation

Under phosphorus (P) deficiency, Lupinus albus (white lupin) releases large amounts of organic acid anions from specialized root structures, so-called cluster or proteoid roots, to mobilize and acquire sparingly soluble phosphates from a restricted soil volume. The molecular mechanisms underlying this release and its regulation are, however, poorly understood. Here, we identified a gene belonging to the aluminium (Al)-activated malate transporter (ALMT) family that specifically contributes to malate, but not citrate release. This gene, LaALMT1, was most prominently expressed in the root apices under P deficiency, including those of cluster roots and was also detected in the root stele. Contrary to several ALMT homologs in other species, the expression was not stimulated, but moderately repressed by Al. Aluminium-independent malate currents were recorded from the plasma membrane localized LaALMT1 expressed in Xenopus oocytes. In composite lupins with transgenic roots, LaALMT1 was efficiently mutated by CRISPR-Cas9, leading to diminished malate efflux and lower xylem sap malate concentrations. When grown in an alkaline P-deficient soil, mutant shoot phosphate concentrations were similar, but iron and potassium concentrations were diminished in old leaves, suggesting a role for ALMT1 in metal root to shoot translocation, a function that was also supported by growth in hydroponics.     

应用电子探针对植物根际和根内营养元素微区分布的探讨

用电子探针可检测出玉米、大豆根际和根内含有Na,Mg,Al,Si,P,S,Cl,K,Ca,Ti,Fe,Cu和Zn 13种元素。这些元素在根际土壤、粘液层和根组织内的含量分布有一定的规律性。除Si,Al,Ca,Fe在根际土壤中峰值较高外,Ti仅在土壤中达到可检测量;S,Fe和Zn富集在粘液层,Mg,P,Cl只在根组织内才有较明显的峰。这些规律可作为区分根—土界面的参考指标。K含量在根内明显高于根际土壤,并由表皮层到中柱径向增加;Ca则与K不同,且受植物种类的影响。     

The development of potential screens based on shoot calcium and iron concentrations for the evaluation of tolerance in Egyptian genotypes of white lupin (Lupinus albus L.) to limed soils

European cultivars of white lupin (Lupinus albus L.) grow poorly in limed or calcareous soils. However, Egyptian genotypes are grown successfully in highly calcareous soil and show no stress symptoms. To examine their physiological responses to alkaline soil and develop potential screens for tolerance, three experiments were conducted in limed and non-limed (neutral pH) soil. Measurements included net CO2 uptake, and the partitioning of Fe2+ and Fe3+ and soluble and insoluble Ca in stem and leaf tissue. Intolerant plants showed clear symptoms of stress, whereas stress in the Egyptian genotypes and in L pilosus Murr. (a tolerant species) was less marked. Only the intolerant plants became chlorotic and this contributed to their reduced net CO2 uptake in the limed soil. In contrast, Egyptian genotypes and L pilosus showed no change in net CO2 uptake between the soils. The partitioning of Ca and Fe either resulted from the stress responses, or was itself a stress response. L pilosus and some Egyptian genotypes differed in soluble Ca concentrations compared with the intolerant cultivars, although no significant difference was apparent in the Ca partitioning of the Egyptian genotype Giza 1. In a limed soil, Giza 1 maintained its stem Fe3+ concentration at a level comparable with that of plants grown in non-limed soil, whereas stem [Fe3+] of an intolerant genotype increased. Gizal increased the percentage of plant Fe that was Fe2+ in its leaf tissue under these conditions; that of the intolerant genotype was reduced. The potential tolerance of the Egyptian genotypes through these mechanisms and the possibility of nutritional-based screens are discussed.     

Localization of phytosiderophore release and of iron uptake along intact barley roots

Under iron deficiency the release of so-called phytosiderophores by roots of barley plants ( Hordeum vulgare L. cv. Europa) was greater by a factor of 10 to 50 compared to iron-sufficient plants. This enhanced release occurred particularly in apical zones of the seminal roots and in the lateral root zones. Under iron deficiency, uptake rates for iron, supplied as FeIII phytosiderophore, increased by a factor of ca 5 as compared to iron-sufficient plants. This enhanced uptake rate for iron was also much more pronounced in apical than in basal root zones. In contrast, with supply of the synthetic iron chelate, FelII EDDHA (ferric diaminoethane-N, N-di- o -hydroxyphenyl acetic acid), the Fe deficiency-enhanced uptake rates for iron were only small and similar along the roots, except for the lateral root zones. The high selectivity of barley roots for uptake and translocation of FeIII phytosiderophores compared with FeIII EDDHA is reflected by the fact that, at the same external concentration (2 μ M ), rates of uptake and translocation of iron from FeIII phytosiderophores were between 100 (Fe-sufficient) and 1 000 times higher (Fe-deficient plants) than from FeIII EDDHA. The relatively high rates of uptake and particularly of translocation of iron supplied as FeIII EDDHA in the zone of lateral root formation strongly suggest an apoplastic pathway of radial transport of the synthetic iron chelate into the stele in this root zone.
The results demonstrate that apical root zones are the main sites both for Fe deficiency-enhanced release of phytosiderophores and for uptake and translocation of iron supplied as FeIII phytosiderophores.