Physiological responses of Pinus sylvestris to atmospheric ammonia

Summary Young saplings of Pinus sylvestris were fumigated for 3 months with ammonia in concentrations ranging from 0 to 240 g m^-3. Despite the much higher concentrations than normal in the field, no visible damage occurred. Photosynthesis, dark respiration, transpiration and biomass production were stimulated. At 240 g m^-3 with high irradiance (PAR: 950 mol m^-2 s^-1), net photosynthesis was stimulated by 24% and dark respiration by 76%. Intitial light use efficiency was not significantly affected. Transpiration increased, both in the dark and at 950 mol m^-2 s^-1 by 40% and 57%, respectively. In the presence of ammonia, stomatal control was less efficient. Though growth of roots was not affected by NH₃, that of current year needles was stimulated, resulting in an increased mass ratio of needles to roots. The nitrogen content of the needles increased, but the contents of other mineral components did not change significantly. Due to increased transpiration per unit of needle area and increased mass of needles per tree, water loss per tree was about twice as high in the treatment with 240 g m^-3 as in the control. Towards the end of fumigation, a 10-day period without water supply followed and then the water potential of the shoots was measured as an indicator of water demand. This demand was higher with higher concentrations of NH₃, suggesting a higher risk of injury from drought.     

Potassium and phosphate uptake of cucumber plants at different ammonia supply

Summary Experiments on cucumber plants grown in nutrient solution were conducted in order to study long and short time effects of ammonia on growth, nutrient element uptake and respiration of roots.Shoot yield and potassium concentration in tissue of plants treated 18 days with varied ammonia concentration were decreased. However, it was not assumed that K deficiency caused the yield reduction. The ammonia effect on K content was more pronounced in roots than in shoots.The decreased K concentration of plant tissue was linked to a diminished ability of plant roots to absorb potassium. The maximum rate of potassium uptake was lowered by ammonia during both, long- and short-time treatment. The results indicated that the NH₃ influence on potassium uptake was due to effects on metabolism and permeability of roots because changes of K uptake rate occurred immediately after starting the NH₃ treatment. Furthermore, it is shown that ammonia inhibited respiration of roots.During the short-time treatment net potassium efflux of roots was observed at higher NH₃ concentrations. The extent of K efflux depended on K concentration of both, root tissue and nutrient solution.Pretreating the plants for 12 hours with ammonia also resulted a decline in K uptake rate. However, plant roots subjected to ammonia concentrations up to 0.09 mM completely recovered during 24 hours after removing the NH₃ treatment whereas at higher NH₃ concentrations only a partial recovery occurred.Furthermore, it was shown that ammonia also influenced P uptake by plant roots.     

The influence of ammonia in nutrient solution on growth and metabolism of cucumber plants

Summary Shoot yield of cucumber plants grown 18 days in nutrient solution with 0.06 mM NH₃ was decreased. Root yield was diminished at 0.09 mM NH₃ The ammonia treatment caused heavy chlorosis increasing with age of leaves. This chlorosis was not due to any nutrient deficiency. Ammonia also influenced the morphology of roots. They were clearly shorter caused by a much smaller size of root cells.The decrease of yield was linked to a reduction of assimilation occurring not only after a long influence of ammonia lasting 14 days, but also within one hour after starting the NH₃ treatment. The decline of assimilation was probably caused by a higher resistance of stomata against CO₂ influx in leaf tissue as can be concluded from the observation that transpiration was decreased in the same way as assimilation.The effect of ammonia in nutrient solution could also be due to the occurrence of higher NH₃ concentrations in leaf tissue, because both, pH of plant press sap as well as NH₄ concentration of plant tissue, were increased.Furthermore, it is shown that the nitrate content of plant tissue was diminished by ammonia whereas ammonium and amide content were raised. Regulation of nitrate uptake of plants by means of ammonium and amide content of tissue is discussed.     

Root-induced increases in soil pH and nutrient availability to field-grown cereals and legumes on acid sandy soils of Sudano-Sahelian West Africa

A field experiment with millet (Pennisetum glaucum L.), sorghum [Sorghum bicolor (L.) Moench], cowpea (Vigna unguiculata L.) and groundnut (Arachnis hypogeae L.) was conducted on severely P-deficient acid sandy soils of Niger, Mali and Burkina Faso to measure changes in pH and nutrient availability as affected by distance from the root surface and by mineral fertiliser application. Treatments included three rates of phosphorus (P) and four levels of nitrogen (N) application. Bulk, rhizosphere and rhizoplane soils were sampled at 35, 45 and 75 DAS in 1997 and at 55 and 65 DAS in 1998. Regardless of the cropping system and level of mineral fertiliser applied, soil pH consistently increased between 0.7 and two units from the bulk soil to the rhizoplane of millet. Similar pH gradients were observed in cowpea, but pH changes were much smaller in sorghum with a difference of only 0.3 units. Shifts in pH led to large increases in nutrient availability close to the roots. Compared with the bulk soil, available P in the rhizoplane was between 190 and 270% higher for P-Bray and between 360 and 600% higher for P-water. Exchangeable calcium (Ca) and magnesium (Mg) levels were also higher in the millet rhizoplane than in the bulk soil, whereas exchangeable aluminium (Al) levels decreased with increasing pH close to the root surface. The results suggest an important role of root-induced pH increases for crops to cope with acidity-induced nutrient deficiency and Al stress of soils in the Sudano-Sahelian zone of West Africa. This revised version was published online in June 2006 with corrections to the Cover Date.     

Atmospheric ammonia and impacts of nitrogen deposition: uncertainties and challenges

Dr Willem Asman concluded that the major global sources of atmospheric NH₃ are excreta from domestic animals and fertilizers. A question raised was: how reliable are the emission estimates and extrapolations? The answer was that emission estimates are surrounded by uncertainty, which is a major handicap to sound modelling of NH₃ dry deposition and, consequently, to obtaining good estimates of critical load exceedences.
Major uncertainties in emission estimates seem to be related to the use of simple emission factors, many of which are highly empirical or have been derived from measurements carried out under conditions which deviate considerably from those following modern practices of handling and applying manure and fertilizers. An example is provided by the commonly used emission factors for synthetic fertilizers (see e.g. Bouwman et al . (1997)), which are much higher than recent micrometeorological assessments seem to suggest. Thus, emission from urea, the most widespread fertilizer used in the world (currently around 55% of world N consumption) can be completely avoided if the fertilizer is incorporated into the upper soil layers. Similarly, a growing crop can reduce losses to well below 10% of the applied amount of urea-N, i.e. to less than half of the generally used emission factors of 15% for Europe and 25% for the tropics. The emission factor for NPK-fertilizer is set at 4%, whereas that for pure calcium-ammonium-nitrate, the same N compound as is present in NPK-fertilizers, is assumed to be only 2%.     

Ammonia in estuaries and effects on fish

This review aims to explore the biological responses of fish in estuaries to increased levels of environmental ammonia. Results from laboratory and field studies on responses of fish to varying salinity and their responses increased ammonia will be evaluated, although studies which examine responses to ammonia, in relation to varying salinity, pH and temperature together are rare. In a survey of British estuaries the continuous measurement of total ammonia showed values that ranged from background levels increasing up to c. 10 mg N l^?1 although higher values have been noted sporadically. In outer estuaries pH values tended to stabilize towards sea water values (e.g. c. pH 8). Upper reaches of estuaries are influenced by the quality of their fresh waters sources which can show a wide range of pH and water quality values depending on geological, climatic and pollution conditions. In general the ammonia toxicity (96 h LC₅₀) to marine species (e.g. 0·09–3·35 mg l^?1 NH₃) appears to be roughly similar to freshwater species (e.g. 0·068–2·0 mg l^?1 NH₃). Ammonia toxicity is related to differences between species and pH rather than to the comparatively minor influences of salinity and temperature. In the marine environment the toxicity of ionized ammonia should be considered. The water quality standard for freshwater salmonids of 21 μg l^?1 NH₃–N was considered to be protective for most marine fish and estuarine fish although the influence of cyclical changes in pH, salinity and temperature were not considered. During ammonia exposures, whether chronic or episodic, estuarine fish may be most at risk as larvae or juveniles, at elevated temperatures, if salinity is near the seawater value and if the pH value of the water is decreased. They are also likely to be at risk from ammonia intoxication in waters of low salinity, high pH and high ammonia levels. These conditions are likely to promote ammonia transfer from the environment into the fish, both as ionized and unionized ammonia, as well as promoting ammonia retention by the fish. Fish are more likely to be prone to ammonia toxicity if they are not feeding, are stressed and if they are active and swimming. Episodic or cycling exposures should also be considered in relation to the rate at which the animal is able to accumulate and excrete ammonia and the physiological processes involved in the transfer of ammonia. In the complex environment of an estuary, evaluation of ammonia as a pollutant will involve field and laboratory experiments to determine the responses of fish to ammonia as salinity and temperature vary over a period of time. It will also be necessary to evaluate the responses of a variety of species including estuarine residents and migrants.     

The time of urea treatment and its effects on the succession of ammonia fungi in two warm temperate forests in Japan

We studied the effects of the timing of urea treatment on the succession of ammonia fungi. In two evergreen Castanopsis cuspidata forests and in one deciduous Quercus serrata forest, we applied 343g urea to 25 and 15 plots of 0.5m², respectively, at three different times of the year. Ten of the early-phase (EP) species, considered to be saprotrophic, and 6 of the late-phase (LP) ones, considered ectomycorrhizal, fruited. In both phases, the commencement, peak, and cessation of fruiting took place simultaneously among all the plots treated at the same time. The fruiting occurred in summer and autumn. Quantity and size of the fruit bodies was larger in the LP than in the EP species. Fruiting of EP species was affected by the treatment time and that of LP species by interaction of the treatment time and vegetation type. EP was short and occurred as one period, whereas LP was long and occurred as two or more fruiting seasons. We found that species composition, dominant species, and degree of its dominance in fruiting of the ammonia fungi are predictable for different treatment times of the year and different vegetation types.     

Removal efficiency and mechanisms of formaldehyde by five species of plants in air-plant-water system

The foliar uptake and transport rates of formaldehyde as well as the abilities of leaf extracts to breakdown formaldehyde were investigated to discuss the formaldehyde removal efficiency and mechanism by five species of plants from air. Results showed that formaldehyde could be transported from air via leaves and roots to rhizosphere water. When exposed to 0.56 mg·m^?3 formaldehyde, the formaldehyde removal rate ranged from 18.64 to 38.47 μg·h^?1g^?1 FW (fresh weight). According to the mass balance in the air–plant–water system, the main mechanism of the formaldehyde loss was its breakdown in plant tissues caused by both enzymatic reaction and redox reaction. Higher oxidation potentials of the leaf-extracts of Wedelia chinensis and Desmodium motorium corresponded well to higher abilities to breakdown added formaldehyde than other plants. Based on the different abilities of fresh and boiled leaf-extracts to dissipate formaldehyde, the enzymatic reaction in Chenopodium album L. was the dominant mechanism while the redox reaction in Kochia scoparia (L.) Schrad. and Silene conoidea L. was the main formaldehyde breakdown mechanism when exposed to low-level formaldehyde in air. The redox mechanism suggested that the formaldehyde removal may be increased by an increasing level of reactive oxygen species (ROS) induced by the environmental stress.     

Avoidance of atmospheric ammonia by domestic fowl and the effect of early experience

An experiment was carried out to determine if broiler fowl, when given a free choice, would avoid ammoniated environments. The effect of rearing aerial environment on these preferences was tested subsequently during a similar free choice experiment.

Firstly, each of four flocks of twelve 26-day-old female Ross broiler fowl, reared in fresh air, was placed in a preference chamber with access to four compartments. After an initial 4-day acclimatisation period, where no ammonia was provided, birds were given a free choice of 4, 11, 20 and 37 ppm atmospheric ammonia over a period of 16 days. Coloured panels in each of the compartments provided the birds with a secondary cue to associate with aerial environment. Ammonia/colour cue treatment was relocated to a different compartment every 4 days. Data collection occurred during two periods that differed in light level: bright (100 lx) and dim (10 lx). The birds avoided the two higher ammonia concentrations, choosing to spend equal amounts of their time in 4 and 11 ppm during the bright phase. During the dim period, broiler fowl spent significantly more time in 4 ppm (light intensity and ammonia occupancy interaction: F_3,60 = 5.40; P = 0.002). Ammonia significantly affected compartment visit duration with back transformed mean visits of 37, 28, 20 and 15 min for 4, 11, 20 and 37 ppm, respectively (Wald statistic = 29.07; d.f. = 3; P < 0.001).

Secondly, eight flocks of 12 female broiler fowl were exposed to either 2 or 19 ppm ammonia from 1 to 26 days of age. At 26 days, each flocks, was placed in a preference chamber with access to four compartments and kept at the same atmospheric ammonia concentration (2 or 19 ppm) during an acclimatisation periods of 4 days. Following acclimatisation, a 4-day test was conducted in which the birds were given a choice of compartments containing 4, 12, 20 and 37 ppm ammonia with colour cues. Birds again avoided the two higher ammonia concentrations (F_3,15 = 8.54; P = 0.002) and there was no significant interaction between ammonia and previous exposure. Again there was an effect of ammonia on compartment visit duration (Wald statistic = 15.94; d.f. = 3; P = 0.001) but there was no significant interaction between ammonia and previous exposure. Broiler fowl avoid ammonia at concentrations commonly found on poultry units regardless of previous experience, suggesting it to be aversive at concentrations above approximately 10 ppm.     

Boron and salinity effects on grafted and non-grafted melon plants

Production of melon (Cucumis melo) may be limited by excesses of boron and salinity, and it was hypothesized that melon grafted onto Cucurbita rootstock would be more tolerant to excessive boron concentrations than non-grafted plants. The objectives of this study were (i) to determine the effects of salinity and excessive boron concentrations in irrigation water on growth and yields of grafted and non-grafted melon plants; and (ii) to study the interaction between the effects of salinity and boron on the uptake of macroelements and boron by grafted and non-grafted melon plants. The plants were grown in pots of Perlite in a greenhouse. The combined effects of boron and salinity on growth and yield were investigated for five boron concentrations, ranging from 0.2 to 10 mg L^{− 1}, and two salinity levels, electrical conductivity (EC) 1.8 and 4.6 dS m^{− 1}, in the irrigation water. With low salinity the boron concentrations in old leaves of non-grafted and grafted plants ranged from 249 to 2827 and from 171 to 1651 mg kg^{− 1} dry weight, respectively; with high salinity the corresponding concentrations ranged from 192 to 2221 and from 200 to 1263 mg kg^{− 1} dry weight, respectively. These results indicate that the grafted plants accumulated less boron than the non-grafted plants when they were exposed to similar boron concentrations, and that both plant types absorbed less boron when irrigated with the more saline irrigation water. It is suggested that: (i) the Cucurbita rootstock excluded some boron and that this, in turn, decreased the boron concentration in the grafted plants; and (ii) the low boron uptake under high-salinity irrigation was mainly a result of reduced transpiration of the plants. Significant negative linear regressions were found between fruit yield and leaf boron concentration for grafted plants, under both low and high salinity levels, and for non-grafted plants under low salinity. The fruit yield of the grafted plants was less affected by boron accumulation in the leaves than that of non-grafted plants. Increasing the water salinity decreased the sensitivity of both plant types to increases in leaf boron concentration, which indicates that the effects of boron and salinity on melon plants were not additive.     

草酸在提高大豆磷吸收利用及抗铝性中的作用

将1mmol/L草酸（pH=6.0)加入到4种难溶性含磷化合物（FePO4、CaHPO4、AlPO4和磷矿粉）的水溶液中,其溶出磷的含量均显著提高,溶磷量随着反应时间的延长首先增大,然后有所下降。水培条件下大豆利用磷矿粉中磷的能力很差,加入1mmol/L草酸（pH=6.0)能促进大豆对磷矿粉中磷的吸收利用,表明草酸可能通过螯溶磷矿粉中的磷而提高了其有效磷的含量。200μmol/L铝离子能显著抑制大豆根的生长,而60μmol/L草酸能基本解除其抑制作用。     

Excretion of ammonia and urea during development in the oyster toadfish,Opsanus tau

The oyster toadfish is one of several teleosts that has been found to produce and excrete large amounts of nitrogenous waste as urea. To clarify the role of urea in the oyster toadfish, urea and ammonia excretion rates were examined in developing fish. Ammonia and urea excretion rates were measured for groups of developing toadfish for three days a week over eight weeks of development. A distinct and significant increase in the excretion rate of urea occurred between the first three weeks (mean = 0.38 mg N/kg‐h) and the fourth through sixth and eighth week of development (mean = 4.68 mg N/kg‐h). This increase of urea excretion occurs at the time of hatching and may be important during development. Preliminary analysis (temperature, pH and salinity) was conducted on water at one toadfish nesting site to provide insight into conditions to which toadfish are exposed.     

The quantitative role of ammonia/ammonium transport and metabolism by plants in the global nitrogen cycle

Estimate of global yearly N assimilation by photolithotrophs are 417 Tmol N in the oceans and 167 Tmol on land and in freshwater, of which diazotrophy contributes 1 (sea) and 10 (land plus freshwater) Tmol N. More than half of the combined N assimilated (416 and 157 Tmol N year⁻¹ in the sea and on land plus freshwater, respectively) is due to reduced N, i. e. organic N and, mainly, NH₃/NH⁺₄. Assimilation of reduced N amounts to up to 334 Tmol N year⁻¹ in the oceans and at least 79 Tmol N year⁻¹ in freshwater and on land. Reassimilation of NH₃/NH⁺₄ within the plant which is related to photorespiration is at least as great as primary NH₃/NH⁺₄ assimilation in the sea, and 8 times greater on land. The less frequently considered reassimilation of NH₃/NH⁺₄ that is related to phenyl-propanoid (mainly lignin) synthesis in land plants is similar (111 Tmol N) to the primary assimilation of NH₃/NH⁺₄ on land each year. Shoots of terrestrial plants have higher NH₃ compensation partial pressures than most natural soils, and especially than have ocean-surface biota. However, gaseous transfer of NH₃/NH⁺₄ from land to the oceans is a negligible component of the global N cycle. Consideration of area-based N assimilation rates, diffusion distances and diffusion coefficients can rationalise why steady-state NH₃/NH⁺₄ concentrations in the sea are lower than in the soil solution. The possibility that photolithotrophs can catalyse the oxidation of NH₃/NH⁺₄, or organic N at the same redox level, to N₂, N₂O, NO, –NO₂, NO⁻, ₂, NO₂ or NO₄⁺, is critically assessed. The tentative conclusions are that such oxidation probably occurs, but is not a major component of the global conversion of reduced N to N₂ and more oxidized N species. More work is needed, especially to determine if NO generated from reduced N (conversion of arginine to citrulline plus NO) has a regulatory role in plants analogous to that established in metazoa. Relative to NO₃⁻ (or N₂) as N sources, growth using NH₃/NH⁺₄ as N source has a number of potential advantages in terms of cost of other resources. Mechanistically predicted economies for NH⁺₄ as N source are: (1) lower cost of photons used and, in transpiring plants, (2) less water lost per unit C assimilated, and (3) lower costs of catalytic Fe, Mn and Mo (unit C assimilated)⁻¹ s⁻¹, as well as (4) a higher maximum growth rate. The lower photon costs are frequently borne out by experimentation and the predicted higher maximum growth rates sometimes occur, while the predicted lower water costs are invariably contradicted. Few data are available for the cost of Fe, Mn or Mo as a function N source.     

Mechanisms of ammonia and ammonium ion toxicity in animal cells: Transport across cell membranes

A model for transport of ammonia and ammonium ions across cell membranes is presented. The model suggests that ammonium ions compete with potassium ions for inward transport, over the cytoplasmic membrane, via potassium transport proteins like the Na⁺/K⁺-ATPase and the Na⁺K⁺2Cl⁻-cotransporter. It also explains the difference between the ammonia/ammonium that is added to the cells and which is formed by the cells during metabolism of amino acids, especially glutamine and glutamate. The ammonium transport and subsequent events lead to predictable intracellular and extracellular pH (pH_e) changes. Experiments which verified the model and the predicted consequences were performed by measurements of the pH_e in concentrated cell suspensions. Addition of ammonium ions caused a time-dependent pH_e increase which was inhibited by potassium ions. The test system is not per se specific for transport measurements but the effect of potassium ions on the pH_e strongly favors our suggested model. Simple diffusion of ammonium ions would not be counteracted by potassium ions. The results show that ammonium ion transport in the murine myeloma cell line (Sp2/0-Ag14) used is inhibited by an excess of potassium ions. Results from experiments with specific inhibitors of suggested transport proteins were not conclusive. It is postulated that one important toxic effect of ammonia/ammonium is an increased demand for maintenance energy, caused by the need to maintain ion gradients over the cytoplasmic membrane. The results also suggest that potassium ions can be used to detoxify ammonia/ammonium in animal cell cultivations.     

Manganese toxicity: The significance of magnesium for the sensitivity of wheat plants

The tolerance of wheat to manganese was investigated in soil and solution culture. Although no critical toxicity concentration could be identified, growth was reduced when the ratio of magnesium to manganese in the shoots (R_p) fell below 20:1 (mgg^–1/mgg^–1). In soil, plant growth relative to unstressed plants (Y) could be described by the empirical equation: % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaaeywaiabg2% da9iaaicdacaGGUaGaaGyoaiaaiwdacqGHsislcaaIWaGaaiOlaiaa% iMdacaaI1aGaaeyzaiaabIhacaqGWbGaaiikaiabgkHiTiaaicdaca% GGUaGaaGymaiaaiodacaaI5aGaaeOuamaaBaaaleaacaqGWbaabeaa% kiaacMcaaaa!4959!\[{\text{Y}} = 0.95 - 0.95{\text{exp}}( - 0.139{\text{R}}_{\text{p}} )\]In solution culture the value of R_p was related to the ratio of the two ions in the nutrient solution (R_s) according to the expression: % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaaeysaiaab6% gacaqGGaGaaeOuamaaBaaaleaacaqGWbaabeaakiabg2da9iaaicda% caGGUaGaaGinaiaaikdacqGHRaWkcaaIWaGaaiOlaiaaisdacaaI4a% GaaeiiaiaabMeacaqGUbGaaeiiaiaabkfadaWgaaWcbaGaae4Caaqa% baGccaGGPaaaaa!47B6!\[{\text{In R}}_{\text{p}} = 0.42 + 0.48{\text{ In R}}_{\text{s}}\]The magnesium concentration in the nutrient solution for optimum growth at a given concentration of manganese was given by: % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGqiVu0Je9sqqrpepC0xbbL8F4rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaGaaeysaiaab6% gacaqGGaGaae4waiaab2eacaqGNbGaaeyxaiabg2da9iaaikdacaGG% UaGaaGioaiaaiMdacqGHRaWkcaaIWaGaaiOlaiaaiwdacaaI0aGaae% iiaiaabMeacaqGUbGaaeiiaiaabUfacaqGnbGaaeOBaiaab2faaaa!4A0B!\[{\text{In [Mg]}} = 2.89 + 0.54{\text{ In [Mn]}}\]Magnesium increased the tolerance of plants to high concentrations of manganese in shoot tissue and also increased the ability of the plant to discriminate against manganese ions in translocation of nutrients from roots to shoots.