The effects of pH and aluminium toxicity on the growth and symbiotic development of cowpeas (Vigna unguiculata (L.) Walp)

Summary The interaction of pH (4 or 6), aluminium (0 or 16 ppm at pH 4) and N source (symbiotic or combined) on the growth and nutrient status of cowpea (Vigna unguiculata) was studied in a glasshouse experiment.Low pH significantly decreased the growth of the plants dependent on symbiotic nitrogen fixation but at pH 4 the addition of 16 ppm Al further depressed growth in both nitrogen regimes. Al-ions appear to exert their effect primarily on the root system, as shown by the reduction in total length and fresh weight. The symbiotic development of the plants was affected by low pH but more markedly by the Al treatment.Shoot nitrogen concentrations were reduced from ca. 2.6% at pH 6 to 1.8% and 0.9% at pH 4 without and with aluminium respectively. Calcium concentration was decreased by low pH and further by Al in both nitrogen regimes.In all Al-treated plants, the aluminium was mainly accumulated in the roots and was associated with an increase in their phosphorus concentration.     

Aluminium speciation and phytotoxicity in alkaline soils

Aim

Highly alkaline soils (pH?>?9.0) may adversely affect agricultural crop productivity. Problems encountered include poor structure and nutrient deficiency. Research based on solution cultures suggests that aluminium (Al) phytotoxicity may occur in soils with pH?>?9.0, but little research has been undertaken on actual soils under controlled conditions. The nature of the Al species responsible and the pH regime of the soils when this occurs are unknown.

Methods

The charge and species of Al responsible for this toxicity was investigated using Zeta Potential measurement, Nuclear Magnetic Resonance (NMR) spectroscopy, Al precipitation characteristics and electrical conductivity as a function of pH. An anion exchange resin was used to evaluate Al availability to plants at alkaline pH. To verify Al phytotoxicity, a pot experiment was performed with plants grown at near neutral and high pH, with and without Al.

Results

The anionic aluminate species of aluminium was ubiquitous at highly alkaline pH, and was the dominant charged species at pH 9.2. Aluminium was phytotoxic at high pH, significantly reducing the stem and root development of field pea test plants over and above that caused by alkalinity alone. The effects of both alkalinity in general and aluminium in particular became noticeable at pH 9.0 and debilitating at pH?>?9.2.

Conclusion

As this corresponds to the pH where aluminate becomes dominant, it is probably responsible for the phytotoxicity.     

Effects of aluminium on growth and nutrient uptake of Betula pendula seedlings

The response to aluminium concentrations was evaluated for birch seedlings ( Betula pendula Roth, formerly Betula verrucosa Ehrh.) by using a growth technique that provides stable internal concentrations of nutrients in plants. Aluminium was added as aluminium nitrate and aluminium chloride and pH was kept at 3.8±0.2 by adding HCl or NaOH. The seedlings were grown in two different series of nutrient treatments, either with near-optimum conditions (relative addition rate 25% day⁻¹) or with constant nutrient stress (relative addition rate 10% day⁻¹) before the aluminium addition. Growth reduction occurred at aluminium concentrations greater than 3 m M , and lethal effects at aluminium concentrations greater than 15 m M . In plants subjected to near-optimum conditions before aluminium addition, the internal nutrient concentrations decreased with increasing aluminium concentration for all macronutrients. The concentration of the macronutrients N, K and P decreased gradually with increasing aluminium concentration, while the concentration of Ca and Mg decreased fairly abruptly when aluminium concentrations exceeded 1 m M . The same tendency was observed in nutrient stressed birch seedlings, but the pattern was more scattered. Relative growth rate of the seedlings was not affected by a low Ca/Al ratio. In all treatments, the molar Ca/Al ratio in/on the roots was below 0.2 at the end of the experiments. As decrease in growth occurs only at high aluminium concentrations, there is no reason to suggest that aluminium in acid soils is growth limiting for natural birch stands.     

Lack of effect of toxic aluminium concentrations on nitrogen fixation by nodulatedStylosanthes species

Summary Effects of three solution aluminium concentrations (0, 25, and 100M) on nitrogen fixation by well-nodulated plants ofStylosanthes hamata, Stylosanthes humilis andStylosanthes scabra are reported. Plants were inoculated with Rhizobium CB756 and grown for 21 days in an aluminium-free nutrient solution at pH 5.3 before imposition of the aluminium treatments.Nitrogen fixation was measured both by the increase in total nitrogen content of the plants and acetylene reduction in roots of plants harvested at 10 and 20 days after imposition of the aluminium treatments. Solution aluminium concentrations as high as 100M, had no detrimental effect on nitrogen fixation in any species.     

Effects of aluminium and micro-nutrients on the sorption of phosphorus byTrifolium repens L. cv. ‘Grasslands Huia’ from nutrient solutions during plant induced pH changes

Summary The sorption of phosphorus from nutrient solution and the pH change in the nutrient solution were monitored over a 24 hour period forTrifolium repens L. cv. ‘Grasslands Huia’ plants. Two different concentration levels of micro-nutrients (B, Cu, Fe, Mn and Zn) and Al formed the factors of a fractional replicate of a 2⁶ factorial design. Measurements were made at four time intervals (30 minutes after the plants were placed on the pots, 3 hours, 6 hours and 24 hours later). In addition to phosphorus, fourteen other nutrients (including nitrate and ammonium) were monitored throughout the experiment. The sorption of phosphorus was significantly influenced by both aluminium and iron. The effect of aluminium and iron on phosphorus sorption is attributed to physico-chemical sorption processes involving the root surface. However the effect on the removal of phosphorus by boron, copper, manganese and zinc was observed only as first order interaction effects —B−Zn, Cu−Zn, Mn−Zn. Thus these three elements (B, Cu and Mn) only affect phosphorus removal in conjunction with zinc. Aluminium and iron together had a separate but very significant effect on the removal of phosphorus at most periods throughout the experiment. In contrast, pH was affected only by aluminium, iron (the pH drop was enhanced) and manganese (the pH drop was decreased) as main effects independent of the other treatment elements.     

Inhibition of nitrate uptake by aluminium in maize

Experiments with two maize (Zea mays L.) hybrids were conducted to determine (a) if the inhibition of nitrate uptake by aluminium involved a restriction in the induction (synthesis/assemblage) of nitrate transporters, and (b) if the magnitude of the inhibition was affected by the concurrent presence of ambient ammonium. At pH 4.5, the rate of nitrate uptake from 240 μM NH₄NO₃ was maximally inhibited by 100 μM aluminium, but there was little measurable effect on the rate of ammonium uptake. Presence of ambient aluminium did not eliminate the characteristic induction pattern of nitrate uptake upon first exposure of nitrogen-depleted seedlings to that ion. Removal of ambient aluminium after six hours of induction resulted in recovery within 30 minutes to rates of nitrate uptake that were similar to those of plants induced in absence of aluminium. Addition of aluminium to plants that had been induced in absence of aluminium rapidly restricted the rate of nitrate uptake to the level of plants that had been induced in the presence of aluminium. The data are interpreted as indicating that aluminium inhibited the activity of nitrate transporters to a greater extent than the induction of those transporters. When aluminium was added at initiation of induction, the effect of ambient ammonium on development of the inhibition by aluminium differed between the two hybrids. The responses indicate a complex interaction between the aluminium and ammonium components of high acidity soils in their influence on nitrate uptake. ei]{gnA C}{fnBorstlap}     

Effect of ammonium- and nitrate-nitrogen nutrition on aluminium tolerance of soybean (Glycine max L.)

Ammonium-nitrogen supply increased Al tolerance (parameter root elongation rate) of soybean (Glycine max L.) plants compared to nitrate-nitrogen supply when grown at constant pH in solution culture. This protective effect of ammonium against Al could only partially be attributed to lowered activity of monomeric aluminium species in the ammonium solution. For ammonium and nitrate-grown plants the relationship between Al concentration in the root tips and total length could be described by the same regression equation. The higher Al tolerance of soybean plants grown in the presence of ammonium was due to restricted ad/absorption of Al which resulted from competition with positively charged Al species for binding sites in the apoplast. Induction of higher symplastic Al tolerance is unlikely because preculture with ammonium decreased rather than increased aluminium tolerance of the plants.     

Si amelioration of Al toxicity in barley (Hordeum vulgare L.) growing in two Andosols

The influence of silicon on aluminium toxicity in barley (Hordeum vulgare L. cv. Shunrai) was studied in two Andosols. Silicon sources were a solution of sodium metasilicate with pH adjusted to 5.0, silica gel, and an industrial waste, porous hydrated calcium silicate. The waste is produced in large amounts in the manufacturing processes of autoclaved light concrete, and has been used as a silicon source for rice plants. The addition of the waste increased the concentration of Si in the soil solution, soil pH and amelioration of aluminium toxicity was observed. The addition of silica gel and sodium metasilicate solution to both soils increased significantly (p<0.05) the Si concentration of the soil solutions, but no amelioration of aluminium toxicity was observed. An amelioration of aluminium toxicity by the waste porous hydrated calcium silicate was probably due to the increase in soil pH rather than to the increase of silicon concentration in the soil solution.     

Study of aluminium concentration and speciation of surface water in four catchments in the Limousin region (France)

This study highlights the contamination of the upstream catchment of several rivers (Vienne, Gartempe, Vézère) in the Limousin (France) by aluminium in the absence of atmospheric pollution. The presence of acid soils on a granitic platform is a natural factor which contributes to the presence of protons and aluminium in water. In the Limousin, it seems that the presence of aluminium in surface water is due to a combination of natural factors: poor acid soils, numerous wet moors and peat bogs. It is currently difficult to evaluate the real impact of intensive cultivation of coniferous trees on the aluminium concentrations found in water in this area. In water, the concentration in total aluminium increases with a decrease in pH and an increase in organic matter. Despite, high concentrations of total aluminium at low pH (close to or lower than 6), the monomeric toxic forms of aluminium, computed with a speciation software, are always inferior to the toxic values for fish. Under such conditions, the concentration in aluminium recorded in some upstream catchments of the Limousin rivers may not cause damage to aquatic life.     

Aluminium tolerance in plants and the complexing role of organic acids

The aluminium cation Al(3+) is toxic to many plants at micromolar concentrations. A range of plant species has evolved mechanisms that enable them to grow on acid soils where toxic concentrations of Al(3+) can limit plant growth. Organic acids play a central role in these aluminium tolerance mechanisms. Some plants detoxify aluminium in the rhizosphere by releasing organic acids that chelate aluminium. In at least two species, wheat and maize, the transport of organic acid anions out of the root cells is mediated by aluminium-activated anion channels in the plasma membrane. Other plants, including species that accumulate aluminium in their leaves, detoxify aluminium internally by forming complexes with organic acids.     

Cinnamate toxicity expression on phenylalanine ammonia-lyase activity,germination and growth of cucumber (Cucumis sativis) seedlings

Low pH (5.2) decreased nodule number and acetylene reduction. Aluminium further depressed those parameters in theRhizobium leguminosarum-Pisum sativum associations examined. In the Al-treated plants nodule formation by strains 128C53 and 128C30 was not affected by 3 or 15 and 30 or 60 μM Al, respectively, as compared with the number of nodules on plants grown at pH 5.2 in the absence of Al. However, improved nodulation rates by those strains did not enhance plant dry weight or reduced nitrogen content. No differences in nitrogenase activity were found among strains of nodulating plants grown at the same aluminium level. These results suggest that Al-ions affected specifically nitrogenase activity and that this effect was primarily responsible for the reduction in plant growth.     

The importance of acid episodes in determining faunal distributions in Welsh streams

SUMMARY. 1. In regional studies of surface-water acidification, annual means of chemical variables are often used to describe differences and change. Outputs from hydrochemical models are often in the form of mean values, which are used in biological models, and these, in turn, are usually derived from responses to mean conditions. Thus, biological forecasts are constrained to ignore the possible effects of the short-term variations in water chemistry which characterize acid streams. This approach requires appraisal.
2. Here, regional Welsh data and daily records from a smaller number of streams were used to investigate the estimation of pH parameters. Variations in aluminium concentration in relation to pH were also assessed. Empirical relationships between invertebrate assemblages, fish populations, mean stream chemistry and measures of fluctuation in pH and aluminium concentration were explored.
3. In general, pH or Al variability and mean pH or Al in Welsh streams were closely related, so that the biological influences of episodes could not easily be separated from those of chronic conditions. Mean pH and mean aluminium concentration were the most effective pH and aluminium statistics used in multivariate models of trout density, which were not improved by including other pH or aluminium variables. For models of invertebrate assemblages based on mean pH or mean aluminium, the inclusion of variables related to episodicity (e.g. pH minimum, aluminium maximum) gave moderate increases in precision.
4. This analysis indicates that it is reasonable to use means of stream chemical variables in biological models of acidification. Consideration of chemical variability could give improvements in some cases, but at the expense of increased model complexity and effort in parameter selection. Nevertheless, we emphasize the need for accurate calibration of both biological and hydrochemical models.     

Accumulation and effects of aluminium in the minnow (Phoxinus phoxinus L.) at different pH levels

The accumulation and effects of 150 pg All^-1 on minnows ( Phoxinus phoxinus L.) were studied in soft water at pH 7, 6 and 5. The fish were kept in a flow-through water system for up to 48 days. Addition of aluminium to the water resulted in poor appetite, passivity and a tendency for the fish to gather as far as possible from the aluminium inlet. The mortality was high at pH 5 and also after addition of aluminium at pH 6. Severe lesions were noted in the gills and olfactory organs after exposure to aluminium. The concentration of aluminium in the gills, liver and kidney was increased at low pH. A large proportion of the aluminium was located on the surface of the gill epithelium. The recovery capacity was pronounced when the aluminium exposure was terminated and the pH was increased from 5 to 7. No further mortality occurred and gill and olfactory organ structure returned to near normality in about 36 days.     

Changes in specific area and energy of root surface of cereal plants in Al-solution cultures. Water vapor adsorption studies

Surface areas and energetic properties of the shooting stage roots of rye (Secale L.), triticale (Triticale), barley (Hordeum L.) and four wheat (Triticum L.) varieties were estimated from experimental water vapor adsorption data. Roots stressed during 10 days at pH 4 with aluminium concentrations ranging from 0 to 40 mg dm^–3 were studied. Roots grown continuously at pH 7 were taken as controls. The surface properties of the roots grown at pH 4 without Al addition were apparently the same as those of the control roots. With the increase of the concentration of the aluminium treatment the surface area of the roots increased for all of the plants, beginning at 5 mg Al dm^–3 for barley, at 10 mg Al dm^–3for wheat and triticale, and at 40 mg Al dm^–3 for rye. The average water vapor adsorption energy of the root surface decreased in general with the increase of Al stress concentration for all plants but triticale, for which this increased. The sensitive cereal varieties seem to have greater amount of high energy adsorption centers (more polar surface) than the resistant ones (lower surface polarity), however more data is needed to justify this hypothesis. For Al-sensitive roots, fraction of high energy adsorption sites decreased and fraction of low energy sites increased under the Al stress. Smaller changes in adsorption energy sites were noted for roots of Al-resistant plants.     

Histochemical observations on the salmonids Salmo salar L. and Salmo trutta L. and the ephemeropterans Baetis rhodani (Pict.) and Ecdyonurus venosus (Fabr.) following a simulated episode of acidity in an upland stream

Salmonids (Salmo salar, Salmo trutta) and mayflies (Baetis rhodani, Ecdyonurus venosus) which had been exposed to simulated episodes of low pH, and low pH with elevated aluminium, were examined histochemically for the presence of aluminium and mucus. The control fish and mayflies and those exposed to low pH did not stain for aluminium, although increased mucus production was demonstrated in fish gills. Trout and salmon exposed to aluminium at low pH exhibited extensive aluminium and mucus coating of the secondary gill lamellae. No mucus was produced by mayflies but aluminium was apparent on all parts of the body. Mean aluminium concentrations of digested fish gills were 2950 and 3050 g g^–1 dry wt. for trout and salmon respectively, whilst for whole specimens of B. rhodani and E. venosus values were 1 200 and 3 175 g g^–1 dry wt.     

Aluminium toxicity expression nutrient uptake,growth and root morphology ofTrifolium repens L. cv. ‘Grasslands Huia’

Summary Hydroponic experiments were undertaken to examine the effect of increasing aluminium levels on the mineral nutrition and root morphology ofT. repens growing in nutrient solution. Toxicity symptoms appear between 27.8 and 47.5 M Al³⁺ activity (148 to 297 M total aluminium). The threshold level corresponding to a 10% reduction in leaf fresh weight is estimated to be approximately 20 M Al³⁺ activity.The concentration of aluminium in the leaves of white clover increases exponentially with aluminium activity in the nutrient solution. The uptake of divalent cations was inhibited but aluminium enhanced potassium and nitrogen concentrations in both leaves and roots.At high pH (pH 6.0) the speciation of aluminium is controlled by the formation of solid aluminium phosphate and aluminium hydroxide except at the lowest aluminium level (37 M) where 99.9 per cent is present as the DTPA complex. As the concentration of total aluminium increases, the percentage of Al-DTPA and soluble aluminium hydroxide decreases whilst solid Al(OH)₃ increases rapidly to reach a maximum of 91.6 percent (of the total aluminium) in the 1180 M aluminium treatment. At pH 4.5 the dominant forms of aluminium are free aluminium ion Al-DTPA, AlSO ₄ ⁺ and AlOH²⁺.The roots of aluminium stressed plants showed symptoms typical of aluminium toxicity.     

Localization and fate of aluminium in the digestive gland of the freshwater snail Lymnaea stagnalis

The digestive gland of the freshwater snail Lymnaea stagnalis, exposed to water containing an elevated concentration of aluminium at neutral pH for up to 30 days, followed by a 20 day recovery period, was examined by light and electron microscopy and X-ray microanalysis. Aluminium was localized in the yellow granules present in the digestive and excretory cells and in the green and small granules present in the digestive cells. More aluminium, silicon, phosphorus and sulphur were present in all three granule types from aluminium exposed snails. The number of yellow and green granules from the digestive gland of aluminium exposed snails showed a progressive increase over the experimental period compared to controls. The number and aluminium content of the granules is likely to reflect the role of the digestive gland as a 'sink' for accumulated aluminium. We propose that intracellular monomeric silica is involved in the detoxification of aqueous aluminium which at neutral pH is largely in the form of an insoluble polyhydroxide. The increased amounts of sulphur and phosphorus in the granules are likely to be part of a broad response to metal loading but probably do not play a significant role in the storage and detoxification of aluminium.     

Geogenically Acidified Mining Lakes — Living Conditions and Possibilities of Restoration

Pyrite and marcasite oxidation in the consequence of lignite surface mining creates lakes with pH as low as 2 to 3, buffered by high contents of iron and aluminium. Living conditions in this extreme habitat for plants and animals are described as well as the characteristics of the pioneer settlement. The utilization of these fish-free lakes is very limited. As possibilities for a water quality improvement special recultivation methods of overburden, chemical neutralization and biological ecotechnologies are recommended.     

The effects of pH, aluminium concentration and temperature on the embryonic development of the European common frog, Rana temporaria

The individual and combined effects of pH, aluminium concentration and temperature, on the development of common frog ( Rana temporaria ) embryos from an upland area of northern England, were investigated in a controlled laboratory study. There was strong evidence to suggest that embryonic survival was lower at pH 4.5 compared with pH 6.0. At pH 4.5, embryonic survival was reduced at the highest aluminium concentrations. There was no strong evidence for a reduction in embryonic survival at lower aluminium concentrations.
Gastrulation and hatching appeared to be the most sensitive stages to both pH and aluminium concentration.
There was strong evidence to suggest that temperature-shocked embryos had reduced survival when compared with those kept at a constant temperature, particularly at pH 4.5.
There was substantial variation in the survival of embryos from different clutches in the same pond with respect to low pH, aluminium concentration and temperatures. Thus R. temporaria has a wide genetic base from which to tolerate environmental changes. It is suggested, however, that the lethal and sublethal effects of low pH, high aluminium concentration and low temperature, could lead to a decrease in recruitment to the adult populations of R. temporaria in upland, northern England.     

Acid-induced acute toxicity of aluminium to three species of filter feeding caddis larvae (Trichoptera, Arctopsychidae and Hydropsychidae)

1. Arctopsyche ladogensis , Hydropsyche angustipennis and Hydropsyche siltalai larvae were exposed to nominal aluminium concentrations of 0, 625, 1250, 2500 and 5000 μg Al l^–1 at pH 5.0 for 96 h. Larvae reared at pH 6.4 and without any aluminium treatment were used as controls. Morphological abnormalities in the anal papillae of the larvae were used as the response variable in estimating the median effective concentrations (EC₅₀) of aluminium.
2. No morphological abnormalities were observed in the control larvae. Only a few individuals of A. ladogensis had darkened anal papillae at pH 5.0 without additional aluminium treatment, whereas increasing aluminium concentrations significantly increased the number of individuals in all species displaying darkening and reduction of the papillae.
3. A. ladogensis appeared to be the most sensitive species to increasing aluminium concentrations, as reflected by the significantly lower mean EC₅₀ value for this species compared with those of H. siltalai and H. angustipennis . H. angustipennis larvae were the most tolerant to aluminium.
4. The results indicate that interspecific differences in sensitivity to aluminium may be a key factor influencing the guild structure of filter feeding caddis flies in acidified streams. The results also imply that toxic effects of aluminium on filter feeding caddis larvae occur due to the impairment of normal osmoregulation processes via damage to the ion-regulatory organs.