Accumulation mechanisms and subcellular distribution of Cu in maize grown on soil treated with [S, S]-ethylenediamine disuccinic acid

Aims

Many studies have proved that EDTA (ethylenediaminetetraacetic acid), EDDS ([S, S’]-ethylenediamine disuccinic acid), and other chelating agents significantly enhance phyto-extraction of copper (Cu) from soil. However, some key factors, such as changes in membrane permeability of root cells and subcellular distribution of Cu and Cu-EDDS complex in leaves and roots, remain unresolved.

Methods

A pot-culture experiment was conducted using soil artificially contaminated with Cu to different degrees to compare its effect on the above factors and the relationship between them in maize (Zea mays L.).

Results

Treatment with 0.5–6.0?mmol?kg^?1 (soil) EDDS increased membrane permeability in root cells significantly (p?<?0.05). Chelated Cu accounted for 14.6%–17.4% of the total Cu content of roots and 77.7%–78.8% of that of leaves and was distributed mainly in cell walls in both.

Conclusions

EDDS increases Cu accumulation in shoots mainly by increasing the content of soluble Cu in soil and membrane permeability of root cells. Cu in soil may be absorbed through the apoplastic pathway into the root xylem, translocated to the shoots, and accumulated there as a Cu-EDDS complex.     

Regulation of sulfur nutrition in wild-type and transgenic poplar over-expressing gamma-glutamylcysteine synthetase in the cytosol as affected by atmospheric H2S

This study with poplar (Populus tremula x Populus alba) cuttings was aimed to test the hypothesis that sulfate uptake is regulated by demand-driven control and that this regulation is mediated by phloem-transported glutathione as a shoot-to-root signal. Therefore, sulfur nutrition was investigated at (a) enhanced sulfate demand in transgenic poplar over-expressing gamma-glutamylcysteine (gamma-EC) synthetase in the cytosol and (b) reduced sulfate demand during short-term exposure to H2S. H(2)S taken up by the leaves increased cysteine, gamma-EC, and glutathione concentrations in leaves, xylem sap, phloem exudate, and roots, both in wild-type and transgenic poplar. The observed reduced xylem loading of sulfate after H2S exposure of wild-type poplar could well be explained by a higher glutathione concentration in the phloem. In transgenic poplar increased concentrations of glutathione and gamma-EC were found not only in leaves, xylem sap, and roots but also in phloem exudate irrespective of H(2)S exposure. Despite enhanced phloem allocation of glutathione and its accumulation in the roots, sulfate uptake was strongly enhanced. This finding is contradictory to the hypothesis that glutathione allocated in the phloem reduces sulfate uptake and its transport to the shoot. Correlation analysis provided circumstantial evidence that the sulfate to glutathione ratio in the phloem may control sulfate uptake and loading into the xylem, both when the sulfate demand of the shoot is increased and when it is reduced.     

Seasonal dynamics of nutrients in leaves and xylem sap of coffee plants as related to different soil compartments

The northwestern province of Costa Rica is a marginal coffee growing area. At the onset of the rainy season low redox potentials probably induce the mobilization of soil Mn resulting in enhanced plant uptake of Mn. To test this hypothesis we monitored from April to the end of June 1995 the mobile Mn in the soil and nutrient and Mn concentrations in leaves and xylem sap of coffee plants. Every 2 weeks we took aggregate and bulk soil samples. The aggregates were mechanically separated into interior and exterior, air-dried and all soil samples were extracted with 1 M NH₄NO₃. We also extracted the field moist soil with distilled water. In addition, the 3rd and the youngest pair of coffee leaves and xylem sap were sampled and analyzed. According to the results of leaf analyses the nutrient supply of the coffee plants in general seemed to be balanced. However, Mn concentrations of 223 mg kg^-1 in the 3rd leaf pair at 18 April were above the optimum and the youngest leaves indicated Fe deficiency, but senescent leaves accumulated Fe and overcame the deficiency. Manganese concentrations in the xylem sap showed a pronounced maximum 2 weeks prior to a similar maximum of mobile Mn in the aggregate exterior. But in general the temporal variation of nutrient concentrations (especially Ca and Mg) in the plants are well correlated with the easily extractable nutrient concentrations in bulk soil. Probably due to its specific absorption and high rates of redistribution within the plant, K in the soil extracts did not correlate with plant concentrations. Element concentrations of youngest leaves could not be correlated with soil concentrations and are not considered to be an adequate tool for monitoring current nutrient uptake. Since plant element concentrations did not correlate with the aggregate interior, plants probably cannot use that nutrient source efficiently.     

Alfalfa genotypes differ in their ability to tolerate zinc deficiency

Response of 13 alfalfa (Medicago sativa L.) genotypes to varied Zn supply (+Zn: 2 mg kg⁻¹ soil, −Zn: no added Zn) was studied in a pot experiment under controlled environmental conditions. Plants were grown for four weeks in a Zn-deficient siliceous sandy soil. Plants grown at no added Zn showed typical Zn deficiency symptoms i.e. interveinal chlorosis of leaves, yellowish-white necrotic lesions on leaf blades, necrosis of leaf margins, smaller leaves and a marked reduction in growth. There was solute leakage from the leaves of Zn-deficient plants, while no solute leakage from Zn-sufficient plants. The ratios of P:Zn, Fe:Zn, Cu:Zn and Mn:Zn in Zn-deficient plants were extremely high compared with Zn-sufficient plants indicating disturbance of P:Zn, Fe:Zn, Cu:Zn and Mn:Zn balance within plant system by Zn deficiency. Genotypes differed markedly in Zn efficiency based on shoot dry matter production. Alfalfa genotypes also differed markedly in P:Zn ratio, Cu:Zn ratio and Fe:Zn ratio under —Zn treatment. The shoot dry weight, shoot:root ratio, chlorophyll content of fresh leaf tissue, solute leakage from the leaves, Zn uptake and distribution of Zn in shoots and roots were the most sensitive parameters of Zn efficiency. Zn-efficient genotypes had less solute leakage but higher shoot:root ratio and higher Zn uptake compared with Zn-inefficient genotypes. Under —Zn treatment, Zn-inefficient genotypes had less Zn partitioning to shoots (33–37%) and more Zn retained in roots (63–67%), while Zn-efficient genotypes had about equal proportions of Zn in roots (50%) and shoots (50%). This revised version was published online in June 2006 with corrections to the Cover Date.     

Transpiration, and nitrogen uptake and flow in two maize (Zea mays L.) inbred lines as affected by nitrogen supply

BACKGROUND AND AIMS: The influence of two nitrogen (N) levels on growth, water relations, and N uptake and flow was investigated in two different inbred lines of maize (N-efficient Zi330 and N-inefficient Chen94-11) to analyse the differences in N uptake and cycling within a plant. METHODS: Xylem sap from different leaves of the inbred lines cultured in quartz sand was collected by application of pressure to the root system. Plant transpiration was measured on a daily basis by weighing five pots of each of the treatments. KEY RESULTS: N-efficient Zi330 had a higher relative growth rate and water-use efficiency at both high (4 mm) and low (0.08 mm) N levels. At a high N level, the amount of N taken up was similar for the two inbred lines; the amount of N transported in the xylem and retranslocated in the phloem was slight greater in Chen94-11 than in Zi330. At a low N level, however, the total amount of N taken up, transported in the xylem and retranslocated in the phloem of Zi330 was 2.2, 2.7 and 2.7 times more, respectively, than that of Chen94-11. Independent of inbred line and N level, the amounts of N transported in the xylem and cycled in the phloem were far more than that taken up by roots at the same time. Low N supply shifted NO(3)(-1) reduction towards the roots. The major nitrogenous compound in the xylem sap was NO(3)(-1), when plants grew at the high N level, while amino acid-N was predominant when plants grew at the low N level. CONCLUSIONS: The N-efficient maize inbred line Zi330 had a higher ability to take up N and cycle N within the plant than N-inefficient Chen94-11 when grown under N-deficiency.     

Dynamics of concentrations and nutrient fluxes in the xylem of Ricinus communis– diurnal course, impact of nutrient availability and nutrient uptake

Diurnal courses of nutrient transport in the xylem and their response to external availability of nutrients were studied. In soil culture, maximal concentrations in all analysed substances were observed during night‐time. Over experimental periods of up to 20 d, concentrations of some ions increased, most by accumulation in the soil. Stringent nutrient conditions were established in a novel pressure chamber. An aeroponic nutrient delivery system inside allows the sampling of xylem sap from intact plants under full control of the nutrient conditions at the root. Analysis of xylem transport under these highly defined conditions established that (1) diurnal variations in concentrations and fluxes in the xylem are dominated by plant‐internal processes; (2) concentrations of nutrients in the xylem sap are highly but specifically correlated with each other; (3) nitrate uptake and nitrate flux to the shoot are largely uncoupled; and (4) in continuous light, diurnal variations of xylem sap concentrations vanish. Step changes in nitrate concentrations of the nutrient solution established that (5) the concomitant increase in nitrate concentration and flux in the xylem is delayed by 2–3 h and is only transient. Diurnal variations of xylem sap composition and use of the new technique to elucidate xylem‐transport mechanisms are discussed.     

Plant uptake and the leaching of metals during the hot EDDS-enhanced phytoextraction process

Using pot experiments, the effect of the application of the biodegradable chelating agent S,S-ethylenediaminedisuccinic acid (EDDS) in hot solutions at 90 degrees C on the uptake of Cu, Pb, Zn, and Cd by corn (Zea mays L. cv. Nongda No. 108) and beans (P vulgaris L. white bean), and the potential leaching of metals from soil, were studied. When EDDS was applied as a hot solution at the rate of 1 mmol kg(-1), the concentrations and total phytoextraction of metals in plant shoots exceeded or approximated those in the shoots of plants treated with normal EDDS at the rate of 5 mmol kg(-1). On the other hand, the leaching of Cu, Pb, Zn, and Cd after the application of the hot EDDS solution at the rate of 1 mmol kg(-1) was reduced by 46%, 21%, 57%, and 35% in comparison with that from the application of normal EDDS at 5 mmol kg(-1), respectively. For treatment with 1 mmol kg(-1) of EDDS, the leached metals decreased to the levels of the control group (that without EDDS amendment) 14 d after the application of EDDS. The soil amendment with biodegradable EDDS in hot solutions may provide a good alternative to chelate-enhanced phytoextraction in enhancing metal uptake by plants and limiting metals from leaching out of the soil.     

Water and Nutrient Dynamics in Surface Roots and Soils are not Modified by Short-term Flooding of Phreatophytic Plants in a Hyperarid Desert

Little is known of the mechanisms employed by woody plants to acquire key resources such as water and nutrients in hyperarid environments. For phreatophytic plants, deep roots are necessary to access the water table, but given that most nutrients in many desert ecosystems are stored in the upper soil layers, viable shallow roots may be equally necessary for nutrient uptake. We sought to better understand the interaction between water and nutrient uptake from soil horizons differing in the relative abundance of these resources. To this end, we monitored plant water and nutrient status before and after applying flood irrigation to four phreatophytic perennial plant species in the remote hyperarid Taklamakan desert in western China. Sap flow in the roots of five plants of the perennial desert species Alhagi sparsifolia Shap., Karelina caspica (Pall.) Less., Calligonum caput medusea Schrenk, and Eleagnus angustifolia Hill. was monitored using the heat ratio method (HRM). Additionally we measured predawn and midday water potential, foliar nitrate reductase activity (NRA), xylem sap nutrient concentration and the concentration of total solutes in the leaves before, 12 and 96 h after flooding to investigate possible short-term physiological effects on water and nutrient status. Rates of sap flow measured during the day and at night in the absence of transpiration did not change after flooding. Moderately high rates of sap flow (HRM heat pulse velocity, 5–25 cm h⁻¹) detected during the day in soils that had a near zero water content at the surface indicated that all species had contact to groundwater. There was no evidence from sap flow data that plants had utilised flood water to increase maximum rates of transpiration under similar climatic conditions, and there was no evidence of a process to improve the efficiency of water or nutrient uptake, such as hydraulic redistribution (i.e. the passive movement of water from moist soil to very dry soil via roots). Measurements of plant water status, xylem sap nutrient status, foliar NRA and the concentration of osmotically active substances were also unaffected by flood irrigation. Our results clearly show that groundwater acts as the major source of water and nutrients for these plants. The inability of plants to utilise abundant surface soil–water or newly available nutrients following irrigation was attributed to the absence of fine roots in the topsoil layer.     

Conservation of element concentration in xylem sap of red spruce

We investigated the chemistry of xylem sap as a marker of red spruce metabolism and soil chemistry at three locations in northern New England. A Scholander pressure chamber was used to extract xylem sap from roots and branches cut from mature trees in early June and September. Root sap contained significantly greater concentrations of K, Ca, Mg, Mn, and Al than branch sap. Sap collected in June contained a signficantly greater concentration of Mn than sap collected in September. Sap concentration was related to forest location for N and Mn. Variations in concentrations of N and K were significantly related to the interaction of tree organ and month of collection. Variations in concentrations of P, Cu, Zn, and Fe were not attributable to tree organ, month of collection, or forest location. Patterns of element concentration in xylem sap compared to previously published data on soil solution chemistry indicated a high degree of homeostatic control of xylem sap chemistry. This control likely represents a significant allocation of resources within the tree energy budget.     

The chemical form of trivalent chromium in xylem sap of maize (Zea mays L.)

Abstract

The toxicity, mobility and bioavailability of Cr, a versatile industrial metal and a contaminant, depends on its chemical form, viz: Cr(lll) and Cr(VI). It may enter humans through plants grown on contaminated soil or irrigated by contaminated water. The phytoavailability and transfer through agricultural food chains requires an understanding of mechanisms of Cr uptake and translocation by plants. Xylem sap transports both nutrient and non-nutrient ions after absorption by roots to aerial parts of the plant. lt transports cations by complexation with organic ligands. Trivalent chromium, though prone to hydrolysis, also complexes O donor ligands. The chemical form in which Cr(lll) is transported by xylem sap was investigated. ln vitro studies were performed by mixing the xylem sap of maize plants at three stages of plant growth with radiotagged Cr(III). The speciation change was investigated after 10 days and 30 days by anion and cation exchange elution chromatography. The elution curves were compared with those of pure Cr(III) and Cr(III) complexes of different synthetic acids. Complexation of Cr(III) with ligands of xylem sap especially with carboxylates was evident. Cationic Cr(III) was vitally being transported as anionic organic complex species. The major species seemed to be that of Cr(III)-citrate. Citric acid was the major complexing acid of xylem sap as determined by HPLC. These mobile and soluble complexes may get immobilized and stored in leaves and other edible plant parts. This may also be a mechanism used by plants for detoxification of toxic Cr(VI) which may become reduced and then complexed.     

Translocation and cycling through roots of recently absorbed nitrogen and sulphur in wheat (Triticum aestivum) during vegetative and generative growth

¹⁵N-Nitrate and ³⁵S-sulphate labelling experiments were performed with spring wheat ( Triticum aestivum L. cv. Timmo) 44. 64, 79, 95 and 115 days after sowing (growth stages arbitrarily denoted I to V). Label was fed to the plants via a fraction of the root system, termed "donor root", whereas the rest of the root ("receiver root") was fed non-labelled nutrient solution. Net uptake rates for both nitrate and sulphate per unit root weight changed little from growth stage I to IV, but were considerably lower at stage V. On a whole-plant weight basis, uptake declined from stage I to IV, because root contribution to total plant weight declined. Between 80 and 95% of absorbed label was translocated to the shoot at all growth stages. At stage V, up to 30% of absorbed label was recovered in the ears. Labelling of the receiver root indicated that, at all growth stages, 10 to 17% of N and 12 to 32% of S translocated to the shoot was retranslocated to the root. This corresponds to between 35 and 85% of the label actually recovered in the roots. Analysis of ¹⁵N-labelling of xylem sap collected from receiver roots at growth stages I to IV indicated that about half of the reduced N in the sap is derived from cycling through roots of recently assimilated N. Evidence of cycling was also obtained at stage V. Labelled sulphate was the only form of S cycled in the plant, but it accounted for only 1 to 7% of the sulphate in the xylem sap.     

Changes in the mineral composition of soybean xylem sap during monocarpic senescence and alterations by depodding

Evidence from earlier studies with explants (stem cutting with a leaf and a pod) indicates that a decline in the supply of mineral nutrients from the roots may prepare the leaves for induction of monocarpic senescence in soybean [ Glycine max (L.) Merrill cv. Anoka). In order to assess the changes in mineral flux from the root system, xylem sap was collected from a decapitated plant under 100 kPa pressure over 50 min. The sap volume yield declines after flowering starts, but increases during pod extension and then decreases again during podfill. The concentrations of K, Ca, Mg, P, S, Zn, Fe, Mn, Cu, Mo and Si rise and then fall during reproductive development, but the exact timing differs among the elements. In contrast, B, Al and Na concentrations show a slow rise initially with a large increase in late pod development. Depodding, which prevents the early death of the plant, inhibits the changes of some elements (K, Mg) but not others (Ca. Mg, P, S, Zn. Fe. Mn, B, Cu, Al), and it does not prevent the decrease in sap volume delivered. Inasmuch as the mineral concentration of xylem sap quantitatively reflects upward mineral flux, the supply of most minerals to the shoot declines, and this decrease seems to be an important factor in the preparatory phase of monocarpic senescence. The different minerals show different patterns of change, which indicate differences in the transport mechanisms and their regulation.     

Mobilization of Cu and Zn by root exudates of dicotyledonous plants in resin-buffered solutions and in soil

It has been frequently suggested that root exudates play a role in trace metal mobilization and uptake by plants, but there is little in vivo evidence. We studied root exudation of dicotyledonous plants in relation to mobilization and uptake of Cu and Zn in nutrient solutions and in a calcareous soil at varying Cu and Zn supply. Spinach (Spinacia oleracea L.) and tomato (Lycopersicon esculentum L.) were grown on resin-buffered nutrient solutions at varying free ion activities of Cu (pCu 13.0–10.4) and Zn (pZn 10.1–6.6). The Cu and Zn concentrations in the nutrient solution increased with time, except in plant-free controls, indicating that the plant roots released organic ligands that mobilized Cu and Zn from the resin. At same pCu, soluble Cu increased more at low Zn supply, as long as Zn deficiency effects on growth were small. Zinc deficiency was observed in most treatment solutions with pZn ≥ 9.3, but not in nutrient solutions of a smaller volume/plant ratio in which higher Zn concentrations were observed at same pZn. Root exudates of Zn-deficient plants showed higher specific UV absorbance (SUVA, an indicator of aromaticity and metal affinity) than those of non-deficient plants. Measurement of the metal diffusion flux with the DGT technique showed that the Cu and Zn complexes in the nutrient solutions were highly labile. Diffusive transport (through the unstirred layer surrounding the roots) of the free ion only could not explain the observed plant uptake of Cu and of Zn at low Zn²⁺ activity. The Cu and Zn uptake by the plants was well explained if it was assumed that the complexes with root exudates contributed 0.4% (Cu) or 20% (Zn) relative to the free ion. In the soil experiment, metal concentrations and organic C concentrations were larger in the solution of planted soils than in unplanted controls. The SUVA of the soil solution after plant growth was higher for unamended soils, on which the plants were Zn-deficient, than for Zn-amended soils. In conclusion, root exudates of dicotyledonous plants are able to mobilize Cu and Zn, and plants appear to respond to Zn deficiency by exuding root exudates with higher metal affinity.     

Accounting for sap flow from different parts of the root system improves the prediction of xylem ABA concentration in plants grown with heterogeneous soil moisture

When soil moisture is heterogeneous, sap flow from, and ABA status of, different parts of the root system impact on leaf xylem ABA concentration ([X-ABA]leaf). The robustness of a model for predicting [X-ABA]leaf was assessed. 'Two root-one shoot' grafted sunflower (Helianthus annuus L.) plants received either deficit irrigation (DI, each root system received the same irrigation volumes) or partial rootzone drying (PRD, only one root system was watered and the other dried the soil). Irrespective of whether relative sap flow was assessed using sap flow sensors in vivo or by pressurization of de-topped roots, each root system contributed similarly to total sap flow during DI, while sap flow from roots in drying soil declined linearly with soil water potential (Psisoil) during PRD. Although Psisoil of the irrigated pot determined the threshold Psisoil at which sap flow from roots in drying soil decreased, the slope of this decrease was independent of the wet pot Psisoil. Irrespective of whether sap was collected from the wet or dry root system of PRD plants, or a DI plant, root xylem ABA concentration increased as Psisoil declined. The model, which weighted ABA contributions of each root system according to the sap flow from each, almost perfectly explained [X-ABA] immediately above the graft union. That the model overestimated measured [X-ABA]leaf may result from changes in [X-ABA] along the transport pathway or an artefact of collecting xylem sap from detached leaves. The implications of declining sap flow through partially dry roots during PRD for the control of stomatal behaviour and irrigation scheduling are discussed.     

Using a dual-stable isotope tracer method to study the uptake, xylem transport and distribution of Fe and its chelating agent from stereoisomers of an Fe(III)-chelate used as fertilizer in Fe-deficient Strategy I plants

A dual-stable isotope tracer experiment was carried out with Fe-deficient sugar beet plants grown hydroponically and resupplied with differentially Fe labeled racemic and meso Fe(iii)-chelates of the ethylendiamine di(o-hydroxyphenylacetic) acid (o,oEDDHA). No short-term Fe isotope exchange reactions occurred in the nutrient solution and plants did not discriminate between (54)Fe and (57)Fe. After 3-6 h, stable Fe isotopes, chelating agents and chelates were analyzed in roots, xylem sap and leaves by ICP-MS and HPLC-ESI/TOFMS. Ferric chelate reductase rates, xylem transport and total uptake were 2-fold higher with the meso isomer than with the racemic one. Both chelating agent isomers were incorporated and distributed by plants at similar rates, in amounts one order of magnitude lower than those of Fe. After 6 h of Fe resupply, most of the Fe acquired was localized in roots, whereas most of the chelating agent was in leaves. In a separate experiment, Fe-deficient sugar beet and tomato plants were treated with different concentrations of Fe(iii)-o,oEDDHA (with a meso/racemic ratio of 1). The xylem sap Fe concentration at 24 h was unaffected by the chelate concentration, with xylem Fe(iii)-o,oEDDHA accounting for 1-18% of total Fe and xylem meso/racemic ratio close to 1. Although most of the Fe coming from Fe(iii)-o,oEDDHA was taken up through a reductive dissociative mechanism, a small part of the Fe may be taken up via non-dissociative mechanisms.     

Sequential response of whole plant water relations to prolonged soil drying and the involvement of xylem sap ABA in the regulation of stomatal behaviour of sunflower plants

Sunflower plants ( Helianihus animus cv. Tall Single Yellow} were grown in the greenhouse in drain pipes (100 mm inside diameter and 1 m long) rilled with John Innes No. 2 compost. When the fifth leaf had emerged, half of the plants were left unwatered for 6 days, rewatered for 2 days and then not watered for another 12 days. Measurements of water relations and abaxial stomatal conductance were made at each leaf position at regular intervals during the experimental period. Estimates were also made of soil water potentials along the soil profile and of ABA concentrations in xylem sap and leaves.
Soil drying led to some reduction in stomatal conductance alter only 3 days but leaf turgors were not reduced until day 13 (6 days after rewatering). When the water relations of leaves did change, older leases became substantially dehydrated while high turgors were recorded in younger leaves. Leaf ABA content measured on the third youngest leaf hardly changed over the first 13 days of the experiment, despite substantial soil drying, while xylem ABA concentrations changed very significantly and dynamically as soil water status varied, even when there was no effect of soil drying on leaf water relations. We argue that the highest ABA concentrations in the xylem, found as a result of substantial soil drying, arise from synthesis in both the roots and the older leaves, and act to delay the development of water deficit in younger leases.
In other experiments ABA solutions were watered on to the root systems of sunflower plants to increase ABA concentrations in xylem sap. The stomatal response to applied ABA was quantitatively very similar to that to ABA generated as a result of soil drying. There was a log-linear relationship between the reduction of leaf conductance and the increase of ABA concentration m xylem sap.     

Regulation of Sulfur Nutrition in Wild-Type and Transgenic Poplar Over-Expressing γ-Glutamylcysteine Synthetase in the Cytosol as Affected by Atmospheric H2S

This study with poplar (Populus tremula × Populus alba) cuttings was aimed to test the hypothesis that sulfate uptake is regulated by demand-driven control and that this regulation is mediated by phloem-transported glutathione as a shoot-to-root signal. Therefore, sulfur nutrition was investigated at (a) enhanced sulfate demand in transgenic poplar over-expressing γ-glutamylcysteine (γ-EC) synthetase in the cytosol and (b) reduced sulfate demand during short-term exposure to H₂S. H₂S taken up by the leaves increased cysteine, γ-EC, and glutathione concentrations in leaves, xylem sap, phloem exudate, and roots, both in wild-type and transgenic poplar. The observed reduced xylem loading of sulfate after H₂S exposure of wild-type poplar could well be explained by a higher glutathione concentration in the phloem. In transgenic poplar increased concentrations of glutathione and γ-EC were found not only in leaves, xylem sap, and roots but also in phloem exudate irrespective of H₂S exposure. Despite enhanced phloem allocation of glutathione and its accumulation in the roots, sulfate uptake was strongly enhanced. This finding is contradictory to the hypothesis that glutathione allocated in the phloem reduces sulfate uptake and its transport to the shoot. Correlation analysis provided circumstantial evidence that the sulfate to glutathione ratio in the phloem may control sulfate uptake and loading into the xylem, both when the sulfate demand of the shoot is increased and when it is reduced.     

The role of mineral nutrients in the increased growth response of tomato plants in solarized soil

Soil solarization is a non-chemical disinfestation technique that frequently promotes plant growth in the absence of known major pathogens, a phenomenon termed increased growth response (IGR). The effect of solarization on plant nutrients and their role in the IGR was studied with tomato plants grown in solarized or non-solarized (control) sandy soil, under controlled conditions. Solarization considerably increased the soil concentrations of water extractable N, K, Ca, Mg and Na at most sites, whereas Cl and DTPA extractable Mn, Zn, Fe and Cu were decreased by the treatment. Plant growth and specific leaf area were enhanced in solarized as well as in N-supplemented control soil. In tomato plants grown in solarized soil, concentrations of most nutrients in the xylem sap, including N, were increased compared to the control, whereas Cl and SO4 levels decreased. The most significant increase in leaf nutrient concentration caused by soil solarization was recorded for N. Furthermore, leaf N concentration was highly and positively correlated with shoot growth. The concentration of Cu increased in leaves from the solarization vs. the control treatment, whereas that of SO4 and Cl decreased, the latter presumably below the critical toxicity level. The correlation between shoot growth and leaf concentration was positive for Cu and inverse for Cl and SO4. In conclusion, we found that soil solarization significantly affects nutrient composition in tomato plants, and provided strong evidence that N, and eventually also Cl, play a major role in IGR.     

Litter N retention over winter for a low and a high phenolic species in the alpine tundra

Mycorrhizal fungus colonization of roots may modify plant metal acquisition and tolerance. In the present study, the contribution of the extraradical mycelium of an arbuscular mycorrhizal (AM) fungus, Glomus mosseae (BEG 107), to the uptake of metal cations (Cu, Zn, Cd and Ni) by cucumber (Cucumis sativus) plants was determined. The influence of the amount of P supplied to the hyphae on the acquisition and partitioning of metal cations in the mycorrhizal plants was also investigated. Pots with three compartments were used to separate root and root-free hyphal growing zones. The shoot concentration of Cd and Ni was decreased in mycorrhizal plants compared to non-mycorrhizal plants. In contrast, shoot Zn and Cu concentrations were increased in mycorrhizal plants. High P supply to hyphae resulted in decreased root Cu concentrations and shoot Cd and Ni concentrations in mycorrhizal plants. These results confirm that some elements required for plant growth (P, Zn, Cu) are taken up by mycorrhizal hyphae and are then transported to the plants. Conversely, Cd and Ni were transported in much smaller amounts by hyphae to the plant, so that arbuscular mycorrhizal fungus colonization could partly protect plants from toxic effects of these elements. Selective uptake and transport of plant essential elements over non-essential elements by AM hyphae, increased growth of mycorrhizal plants, and metal accumulation in the root may all contribute to the successful growth of mycorrhizal plants on metal-rich substrates. These effects are stimulated when hyphae can access sufficient P in soil.