Plant induced alteration in the rhizosphere and the utilisation of soil heterogeneity

Plant-soil interactions result in a special rhizosphere soil chemistry, differing from that of the bulk soil found only a few mm from the root. The aim of this study was to investigate adaptation mechanisms of herbs growing in acid soils through studying their rhizosphere chemistry in a greenhouse experiment and in a field study. Ten herbs were grown in acid soil (pH 4.2 in the soil solution) in the greenhouse. The concentrations of NO₃ ^-, SO₄ ^2-, phosphates, Ca²⁺, Mg²⁺, Mn²⁺, K⁺, Na⁺, NH₄ ⁺ and pH were analysed in soil solutions obtained by centrifugation. The general pattern found was a depletion of nutrients in the rhizosphere compared with their concentrations in the bulk soil. The pH increase (up to 0.7 units) in the rhizosphere soil appeared to be caused by plant uptake of NO₃ ^- (r²=0.88). The ion concentrations in the soil solution of the rhizosphere were dependent on plant species and biomass increase. Although species with a larger biomass and higher growth rates showed a higher degree of ion depletion (except for Na⁺, SO₄ ^2-) in the rhizosphere, there were also species specific responses. A field study of five herbs at five oak forest sites in Southern Sweden (Scania) was also carried out. In addition to the soil solution concentrations, the loss on ignition (LOI) and the concentrations of 0.1 M BaCl₂ extractable K⁺, Mg²⁺, Mn²⁺, Ca²⁺, and Al ions were measured. The amount of soil solution Al was determined as free ionic (quickly reacting) Al. For all species and sites, the LOI and the concentrations of exchangeable cations were higher in the rhizosphere than in the bulk soil, apparently due to the roots preferably growing at organic-rich microsites. The concentrations of the ions as measured in the centrifuged soil solution, were either higher in the rhizosphere than in the bulk soil or were the same in both, except for NO₃ ^- and quickly reacting Al. The lower concentrations of quickly reacting Al in the rhizosphere, compared with the bulk soil could indicate the uptake of Al by the plant or the exudation of complexing substances. The pH differences were only small and mostly non-significant. Plant-soil interactions and the ability of plants to utilise heterogeneity of the soil appear to be more important for plant growth in acid soils than recognised heretofore. Rhizosphere studies provide an important means of understanding plant strategies in acid soils. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of litter incorporation and nitrogen fertilization on the contents of extractable aluminium in the rhizosphere soil of tea plant (Camallia sinensis (L.) O. Kuntze)

The effects of litter incorporation and nitrogen application on the properties of rhizosphere and bulk soils of tea plants (Camellia sinensis (L.) O. Kuntze) were examined in a pot experiment. Total of 8 treatments included four levels of tea litter additions at 0, 4.9, 9.8, and 24.5 g kg^–1 in combination with two N levels (154.6 mg kg^–1 and without). After 18 months of growth the rhizosphere soil was collected by removing the soil adhering to plant roots and other soil was referred to as bulk soil. The dry matter productions of tea plants were significantly increased by N fertilization and litter incorporation. The effect of litter was time-depending and significantly decreased the content of exchangeable Al (Al_ex, by 1 mol L^–1 KCl) and Al saturation at 9 months after litter incorporation whereas soil pH was not affected, although the litter contained high Al content. After 18 months, the contents of extractable Al by dilute CaCl₂, CuCl₂ + KCl, NH₄OAC, ammonium oxalate and sodium citrate (Al_CaCl2, Al_Cu/KCl, Al_NH4OAC, Al_Oxal, and Al_Cit respectively) and Al_ex, were not affected by litter application, except that of Al_CaCl2 in the rhizosphere soil which was decreased following litter additions. Nitrogen fertilization with NH₄ ⁺ (urea and (NH₄)₂SO₄) significantly reduced soil pH, the contents of exchangeable Ca, K, Mg and base saturation while raised extractable Al levels (Al_CaCl2, Al_Cu/KCl, Al_NH4OAC, and Al_ex). In the rhizosphere soils exchangeable K accumulated in all treatments while exchangeable Ca and Mg depleted in treatments without litter application. The depletions of Ca and Mg were no longer observed following litter incorporation. This change of distribution gradients in rhizosphere was possibly due to the increase of nutrient supplies from litter decomposition and/or preferable root growth in soil microsites rich in organic matter. Lower pH and higher extractable Al (Al_CaCl2, Al_ex, and Al_NH4OAC) in the rhizosphere soils, regardless of N and litter treatments, were distinct and consistent in all treatments. Such enrichments of extractable Al in the rhizosphere soil might be of importance for tea plants capable of taking up large amounts of Al.     

Effects of liming on rhizosphere chemistry and growth of fine roots and of shoots of sessile oak (Quercus petraea)

Soil acidification can be detrimental to root growth and nutrient uptake, and liming may alleviate such acidification. In the following study, seedlings of sessile oak (Quercus petraea Liebl. M.) were grown in rhizotrons and subjected to liming (L) or gypsum (G) treatments and compared with the control (C). In order to study and interpret the impact of these calcium rich treatments on fine root development and tree growth, the following parameters were assessed: fine root biomass, fine root length, seedling development (height, diameter, leaves), seedling biomass, nutrient content of roots and seedlings, bulk soil and soil solution chemistry and rhizosphere soil chemistry. The results show that liming increased bulk soil pH, exchangeable Mg, Ca and the Ca/Al molar ratio, and decreased exchangeable Al, mainly in the A-horizon. Gypsum had a similar but smaller impact on exchangeable Al, Ca, H⁺ and the Ca/Al molar ratio in the A-horizon, but reacted with depth, so that exchangeable Mn, Mg and Ca were increased in the B-horizon. In the rhizosphere, the general pattern was determined by the treatment effects of the bulk soil. Most elements were more concentrated in the rhizosphere than in bulk soil, except for Ca which was less concentrated after liming or gypsum application. In the B-horizon rhizosphere pH was increased by the treatments (L > G,C) close to the root tips. Furthermore, the length of the zone with a positive root-induced pH increase was greater for the limed roots as compared with both the other treatments. Fine root growth was stimulated by liming (L > G,C) both in terms of biomass and length, whereas specific root length was not obviously affected apart from the indication of some stimulation after liming at the beginning. The live:dead ratio of fine roots was significantly higher in the limed rhizotrons as compared to the control (G not assessed), indicating lower mortality (higher longevity). Shoot growth showed greater lime-induced stimulation (L > G,C) as compared to root growth. As a result the shoot:root ratio was higher in the limed rhizotrons than in the control (L > G,C). Liming induced a higher allocation of P, S, Mg, Ca and K to the leaves, stem and twigs. Gypsum showed similar effects, but was only significant for S. Liming increased the foliar Ca/Al ratio by both increasing foliar Ca and decreasing foliar Al, whereas gypsum did not clearly improve foliar nutrition. This study suggests that a moderate application of lime can be successful in stimulating seedling growth, but that gypsum had no effect on seedling growth. It can be concluded that this lime-induced growth stimulation is directly related to the improved soil fertility status, and the alleviation of Al toxicity and acid stress, resulting in better foliar nutrition. The impact of liming on fine roots, as a consequence, was not limited to a stimulation of the total amount of fine roots, but also improved the root uptake performance. This revised version was published online in June 2006 with corrections to the Cover Date.     

The influence of soil chemistry on fine root aluminum concentrations and root dynamics in a subalpine Spodosol,Washington State,USA

The relationship between root Al concentrations and Al fractions in the soil solution was examined in a mature Abies amabilis ecosystem in the Cascade Range of Washington State. The naturally acidic soils in these ecosystems lead to high concentrations of aqueous Al in soil solutions and contribute to the biocycling of Al by the A. amabilis/T. mertensiana stand. Root concentrations of Al were very closely related to aqueous Al³⁺ activities, but poorly correlated with total aqueous Al concentrations. The solution Al/Ca molar ratios followed a seasonal cycle with low values during the fall and high values during the spring. Ratios remained <1 throughout the year in the Oa horizon while they varied between 2 and 14 in the E and Bhs horizons. The vertical distribution of roots and the mortality of fine roots may be linked to the soil solution Al/Ca ratio. Root cation exchange capacity ranged between 180 and 225 mol g^-1 and the exchangeable Al fraction represented from 12–17% of the total Al content in the root. Evidence for solid-phase co-precipitates of Al with PO₄ and oxalate was indicated from selective dissolution of the root tissue. Sufficient quantities of PO₄ and oxalate exist in the roots to tie up 20–40% of the Al present in the roots of the Oa and E horizons, but only 9% of that present in the Bhs horizon. Species differences in the distribution of Al between the above-ground and below-ground components may be dictated by these retention processes in the fine roots.     

The effect of phosphate rock dissolution on soil chemical properties and wheat seedling root elongation

Soils of the Appalachian region of the United States are acidic and deficient in P. North Carolina phosphate rock (PR), a highly substituted fluoroapatite, should be quite reactive in these soils, allowing it to serve both as a source of P and a potential ameliorant of soil acidity. An experiment was conducted to evaluate the influence of PR dissolution on soil chemical properties and wheat (Triticum aestivum cv. Hart) seedling root elongation. Ten treatments including nine rates of PR (0, 12.5, 25, 50, 100, 200, 400, 800, and 1600 mg P kg^-1) and a CaCO₃ (1000 mg kg^-1) control were mixed with two acidic soils, moistened to a level corresponding to 33 kPa moisture tension and incubated for 30 days. Pregerminated wheat seedlings were grown for three days in the PR treated soils and the CaCO₃ control. Root length was significantly (P<0.05) increased both by PR treatments and CaCO₃, indicating that PR dissolution was ameliorating soil acidity. The PR treatments increased soil pH, exchangeable Ca, and soil solution Ca while lowering exchangeable Al and 0.01 M CaCl₂ extractable soil Al. Root growth in PR treatments was best described by an exponential equation (P<0.01) containing 0.01 M CaCl₂ extractable Al. The PR dissolution did not reduce total soil solution Al, but did release Al complexing anions into soil solution, which along with increased pH, shifted Al speciation from toxic to nontoxic forms. These results suggest that North Carolina PR should contribute to amelioration of soil acidity in acidic, low CEC soils of the Appalachian region.     

Modelling of the dynamics of Al and protons in the rhizosphere of maize cultivated in acid substrate

The object of this study was to analyze the dynamics of Al and protons in the rhizosphere of maize cultivated in a simple acid substrate, so as to allow the use of a dynamic model of the functioning of a rhizosphere consisting of an organic phase (an agarose gel) and a mineral phase (an amorphous aluminium hydroxide). Two cultivars of maize (Zea mays L.), one Al-sensitive and the other Al-tolerant, were cultivated on this substrate in the presence of different proportions of NH ₄ ⁺ and NO ₃ ^- , which served to acidify the rhizosphere to a greater or lesser extent. The state of the agarose gel and of the cell walls of the roots were monitored using an ion exchange model which had previously been calibrated for each substrate. The experiment showed that Al and protons reduce root growth and the Ca and Mg content in the root, while relative growth varies little between pH 4.0 and pH 4.5. The model showed that competition between Al and protons for the binding sites of the cell walls might account for these results. The sensitivity of the model to the rate of Al(OH)₃ dissolution and to the cation exchange capacity of the culture substrate was tested by numerical simulation. When roots release protons and dissolve Al(OH)₃ in the rhizosphere, there is little possibility of Al desorption by protons on the cell walls at pHs compatible with good root growth of maize, plant specie sensitive to Al and H. Furthermore, the phytotoxicity of the different forms of Al hydroxides should be considered only in taking into account the dynamics of the whole system, in particular the solubilisation of Al in the rhizosphere. This revised version was published online in June 2006 with corrections to the Cover Date.     

Dynamics of phosphorus in the rhizosphere of maize and rape grown on synthetic,phosphated calcite and goethite

In calcareous soils the dynamics of phosphorus is controlled by calcite and iron oxides such as goethite which strongly retain P and consequently maintain low P concentrations in soil solution. Plants can drastically change chemical conditions in the rhizosphere, in particular by releasing H⁺ or OH⁻ or by excreting organic anions. By modifying the dissolution/precipitation and desorption/adsorption equilibria, roots can influence the mobility of soil P. The aim of this work was to test whether H⁺ or OH⁻ release can induce the mobilization of P in the rhizosphere of maize and rape supplied with NO₃-N or NH₄-N and grown on synthetic phosphated calcite or goethite as sole source of P. With P-calcite, the mobilization of P was generally related to the acidification of the rhizosphere. With P-goethite, rhizosphere acidification induced some increase of DTPA-extractable Fe and hence dissolution of goethite. Rhizosphere P was concomitantly depleted but the mechanisms involved are less clear. The difference in behavior of the two species is discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Wood-ash recycling affects forest soil and tree fine-root chemistry and reverses soil acidification

Wood ash was applied to a forest ecosystem with the aim to recycle nutrients taken from the forest and to mitigate the negative effects of intensive harvesting. After two years, the application of 8,000 kg ha⁻¹ of wood ash increased soil exchangeable Ca and Mg. Similarly, an increase in Ca and Mg in the Norway spruce fine roots was recorded, leading to significant linear correlations between soil and root Ca and soil and root Mg. In contrast to these macronutrients, the micronutrients Fe and Zn and the toxic element Al decreased in the soil exchangeable fraction with the addition of wood ash, but not in the fine roots. Only Mn decreased in soil and in fine roots leading to a significant linear correlation between soil and root Mn. In soil, as well as in fine roots, strong positive correlations were found between the elements Ca and Mg and between Fe and Al. This indicates that the uptake of Mg resembles that of Ca and that of Al that of Fe. With the wood ash application, the pH increased from 3.2 to 4.8, the base saturation from 30% to 86%, the molar basic cations/Al ratio (BC/Al) of the soil solution from 1.5 to 5.5, and the molar Ca/Al ratio of the fine roots from 1.3 to 3.7. Overall, all below-ground indicators of soil acidification responded positively to the wood ash application within two years. Nitrate concentrations increased only slightly in the soil solution at a soil depth of 75–80 cm, and no signs of increased heavy metal concentrations in the soils or in the fine roots were apparent. This suggests that the recycling of wood ash could be an integral part of sustainable forest management because it closes the nutrient cycle and reverses soil acidification.     

Depletion of Selenium in Soil Solution due to its Enhanced Sorption in the Rhizosphere of Soybean

The effect of plant roots on selenium (Se) mobility in soil was studied by a large-scale pot experiment in order to understand the environmental behavior of Se in agricultural soils under plant growth conditions. Soybean plants (Glycine max (L.) Merrill) were grown in a greenhouse for 84 d. The concentrations of Se and major elements (K, Ca, Mg, Na, and Al) in the soil solutions and in the plants were measured at different growth periods. Concentrations of Se and major cations in soil solution decreased as the soybean plants grew, while the concentrations of Al increased. It was assumed that the soybean roots released H⁺ with the uptake of cations; consequently, due to the acidification of the rhizosphere, Al³⁺ was released starting from the soil solid phase. The decreased Se concentration in the soil solution should be due to the enhancement of Se sorption onto the soil solid phase. The increase of Se sorption level in the rhizosphere was examined in a small-scale pot experiment. The soil–soil solution distribution coefficient of Se (K _d-Se) was observed as an index of Se sorption level. K _d-Se clearly increased in the rhizosphere soil after cultivation. The effects of pH and Al³⁺ in the rhizosphere on Se sorption were assessed by K _d-Se measurements at different levels of HCl and AlCl₃. In this third experiment, a decrease in pH increased K _d-Se values, but no specific effect was observed on Se sorption due to increased Al³⁺. These results show that the Se mobility in agricultural soil could be decreased by plant roots under plant growth conditions due to enhanced Se sorption in the rhizosphere.     

Soil solution chemistry in the rhizosphere of roots of sessile oak (Quercus petraea) as influenced by lime

This study describes the soil solution chemistry in the rhizosphere of fine roots of sessile oak ( Quercus petraea (M.) Liebl.) grown in rhizotrons. A control was compared with soils treated with an equivalent CaCO₃ of 1.4 t ha^-1 CaO. Solution samples were extracted from the B-horizon using micro suction cups with a suction of _∼40 kPa. Two series of experiments were carried out: one irrigated with rain water (age of seedling 2 to 4 months) and one irrigated with demineralized water (age of seedlings 1.5 to 2 months). Half of the sampling points were choosen close to the roots and half in the bulk soil. In both experiments there was generally no rhizospheric gradient after liming. In contrast, in the control, depletion in the rhizosphere occurred for most of the ions studied (Mg, Ca, Al, K, NO₃ ^-, NH₄ ⁺, Cl^-) in the demineralized water experiment, but this was different when rainwater was used. The latter effect is probably due to the higher solution concentrations in the rainwater experiment but could also be a result of root damage due to low Ca/Al ratios in the rhizosphere solution. It was concluded, that liming improved the chemical composition in the rhizosphere soil solution by increasing overall solute concentration to levels enabling sufficient and easier nutrient uptake by roots. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of nitrogen form and phosphorus source on the growth,nutrient uptake and rhizosphere soil property of Camellia sinensis L.

The effects of nitrogen form and phosphorus source on the growth, nutrient uptake and rhizosphere soil property of tea (Camellia sinensis L.) were investigated in a pot experiment. The experiment was performed with a compartmental cropping device, which enables the collection of rhizosphere soil at defined distances from the root of tea plant. Nitrogen was supplied as nitrate or ammonium in combination with soluble phosphorus as Ca(H₂PO₄)₂ or insoluble P as rock phosphate. The leaf dry matter production of tea was significantly greater in the treatments with NH₄ ⁺ than NO₃ ^-, whereas dry matter production of root and stem was not significantly affected. Addition of phosphorus as either source did not influence the dry matter production. The concentrations of K in root, Mg and Ca in both the shoot and root supplied with NO₃ ^- were significantly higher than in NH₄ ⁺ and influence of P sources was minor. On the contrary, Al and Mn concentrations were significantly larger in NH₄ ^--fed plants which could be attributed to remarkably increased availability of Al and Mn caused by acidification of the rhizosphere soil (the first 1-mm soil section from the root surface) with NH₄–N nutrition. The concentration of N in shoot was also significantly higher in NH₄- than in NO₃-fed plants, indicating higher use efficiency of NH₄–N. Whatever the phosphate source, rhizosphere pH declined in ammonium compared to in nitrate treatment. The pH decrease was much larger when no P or soluble P were applied and reached 0.85–1.30 units which extended to 3–5 mm away from the root surface. Exchangeable acidity, content of exchangeable Al and Mn were also considerably higher in the rhizosphere soils of NH₄ ⁺ fed tea plants. Significant amounts of P dissolved from rock phosphate accumulated in rhizosphere of NH₄ ⁺, not NO₃ ^-, suggesting that the dissolution of rock phosphate was induced by the proton excreted by tea root fed with ammonium. With soluble P addition, shoot and root P concentrations were greater in NH₄ ⁺ than in NO₃ ^- treatment and it appeared that this difference could not be sufficiently explained by the available P content in soil which was only slightly higher in NH₄ ⁺ treatment. With rock phosphate addition, the shoot and root P concentrations were hardly affected by nitrogen form, although the available P content was much higher and accumulated in the rhizosphere soil supplied with ammonium. The reason for this was discussed with regard to the inter-relationship of Al with P uptake. This revised version was published online in June 2006 with corrections to the Cover Date.     

Relationships among foliar chemistry, foliar polyamines, and soil chemistry in red spruce trees growing across the northeastern United States

Forest trees are constantly exposed to various types of natural and anthropogenic stressors. A major long-term goal of our research is to develop a set of early physiological and biochemical markers of stress in trees before the appearance of visual symptoms. Six red spruce (t Picea rubens Sarg.) stands from the northeastern United States were selected for collection of soil and foliage samples. All of the chosen sites had soil solution pH values below 4.0 in the Oa horizon but varied in their geochemistry. Some of these sites were apparently under some form of environmental stress as indicated by a large number of dead and dying red spruce trees. Samples of soil and needles (from apparently healthy red spruce trees) were collected from these sites four times during a two-year period. The needles were analyzed for perchloric acid-soluble polyamines and exchangeable inorganic ions. Soil and soil solution samples from the Oa and B horizons were analyzed for their exchange chemistry. The data showed a strong positive correlation between Ca and Mg concentrations in the needles and in the Oa horizon of the soil. However, needles from trees growing on relatively Ca-rich soils with a low exchangeable Al concentration and a low Al:Ca soil solution ratio had significantly lower concentrations of putrescine and spermidine than those growing on Ca-poor soils with a high exchangeable Al concentration and a high Al:Ca soil solution in the Oa horizon. The magnitude of this change was several fold higher for putrescine concentrations than for spermidine concentrations. Neither putrescine nor spermidine were correlated with soil solution Ca, Mg, and Al concentrations in the B horizon. The putrescine concentrations of the needles always correlated significantly with exchangeable Al (r²=0.73, t p0.05) and soil solution Al:Ca ratios (r²=0.91, t p0.01) of the Oa horizon. This suggests that in conjunction with soil chemistry, putrescine and/or spermidine may be used as a potential early indicator of Al stress before the appearance of visual symptoms in red spruce trees.     

Mechanisms of Aluminum Tolerance in Wheat : An Investigation of Genotypic Differences in Rhizosphere pH, K, and H Transport, and Root-Cell Membrane Potentials

Control of rhizosphere pH and exclusion of Al by the plasma membrane have been hypothesized as possible mechanisms for Al tolerance. To test primarily the rhizosphere pH hypothesis, wheat cultivars (Triticum aestivum L. `Atlas 66' and `Scout'), which differ in Al tolerance, were grown in either complete nutrient solution, or 0.6 millimolar CaSO₄, with and without Al at pH 4.50. A microelectrode system was used to simultaneously measure rhizosphere pH, K⁺, and H⁺ fluxes, and membrane potentials (E_m) along the root at various distances from the root apex. In complete nutrient solution, the rhizosphere pH associated with mature root cells (measured 10-40 millimeters from the root apex) of Al-tolerant `Atlas 66' was slightly higher than that of the bulk solution, whereas roots of Al-sensitive `Scout' caused a very small decrease in the rhizosphere pH. In CaSO₄ solution, no significant differences in rhizosphere pH were found between wheat cultivars, while differential Al tolerance was still observed, indicating that the rhizosphere pH associated with mature root tissue is not directly involved in the mechanism(s) of differential Al tolerance. In Al-tolerant `Atlas 66', growth in a CaSO<sub>4</sub> solution with 5 micromolar Al (pH 4.50) had little effect on net K<sup>+</sup> influx, H<sup>+</sup> efflux, and root-cell membrane potential measured in cells of mature root tissue (from 10-40 mm back from apex). However, in Al-sensitive `Scout', Al treatment caused a dramatic inhibition of K⁺ influx and both a moderate reduction of H⁺ efflux and depolarization of the membrane potential. These results demonstrate that increased Al tolerance in wheat is associated with the increased ability of the tolerant plant to maintain normal ion fluxes and membrane potentials across the plasmalemma of root cells in the presence of Al.     

Responses of maize grain yield to changes in acid soil characteristics after soil amendments

An experiment was conducted from 1997 to 2000 on an acid soil in Cameroon to assess the effectiveness of cultivating acid tolerant maize (Zea mays L.) cultivar and the use of organic and inorganic fertilizers as options for the management of soil acidity. The factors investigated were: phosphorus (0 and 60 kg ha^?1), dolomitic lime (0 and 2 t ha^?1), organic manure (no manure, 4 t ha^?1 poultry manure, and 4 t ha^?1 of leaves of Senna spectabilis), and maize cultivars (ATP-SR-Y – an acid soil-tolerant, and Tuxpeño sequia – an acid susceptible). On acid soil, maize grain yield of ATP-SR-Y was 61% higher than the grain yield of Tuxpeño sequia. Continuous maize cultivation on acid soil further increased soil acidity, which was manifested by a decrease in pH (0.23 unit), exchangeable Ca (31%) and Mg (36%) and by an increase in exchangeable Al (20%). Yearly application of 60 kg ha^?1 of P for 3 years increased soil acidity through increases in exchangeable Al (8%) and H (16%) and a decrease in exchangeable Ca (30%), Mg (11%) and pH (0.07 unit). Lime application increased grain yield of the tolerant (82%) and susceptible (208%) cultivars. The grain yield increases were associated with a mean decrease of 43% in exchangeable Al, and 51% in H, a mean increase of 0.27 unit in pH, 5% in CEC, 154% in exchangeable Ca, and 481% in Mg contents of the soil. Poultry manure was more efficient than leaves of Senna producing 38% higher grain yield. This yield was associated with increases in pH, Ca, Mg and P, and a decrease in Al. The highest mean grain yields were obtained with lime added to poultry manure (4.70 t ha^?1) or leaves of Senna (4.72 t ha^?1). Grain yield increase was more related to the decrease in exchangeable Al (r = ?0.86 to ?0.95, P<0.01) and increase in Ca (r = 0.78–0.94, P<0.01), than to pH (r = ?0.57 (non-significant) to ?0.58 (P<0.05)). Exchangeable Al was the main factor determining pH (r = ?0.88 to ?0.92, P<0.01). The yield advantage of the acid tolerant cultivar was evident even after correcting for soil acidity. Acid soil-tolerant cultivars are capable of bringing unproductive acid soils into cultivation on the short run. The integration of soil amendments together with acid soil-tolerant cultivar offers a sustainable and comprehensive strategy for the management of acid soils in the tropics.     

Origins of root-mediated pH changes in the rhizosphere and their responses to environmental constraints: A review

The aim of the present review is to define the various origins of root-mediated changes of pH in the rhizosphere, i.e., the volume of soil around roots that is influenced by root activities. Root-mediated pH changes are of major relevance in an ecological perspective as soil pH is a critical parameter that influences the bioavailability of many nutrients and toxic elements and the physiology of the roots and rhizosphere microorganisms. A major process that contributes root-induced pH changes in the rhizosphere is the release of charges carried by H⁺ or OH^– to compensate for an unbalanced cation–anion uptake at the soil–root interface. In addition to the ions taken up by the plant, all the ions crossing the plasma membrane of root cells (e.g., organic anions exuded by plant roots) should be taken into account, since they all need to be balanced by an exchange of charges, i.e., by a release of either H⁺ or OH^–. Although poorly documented, root exudation and respiration can contribute some proportion of rhizosphere pH decrease as a result of a build-up of the CO₂ concentration. This will form carbonic acid in the rhizosphere that may dissociate in neutral to alkaline soils, and result in some pH decrease. Ultimately, plant roots and associated microorganisms can also alter rhizosphere pH via redox-coupled reactions. These various processes involved in root-mediated pH changes in the rhizosphere also depend on environmental constraints, especially nutritional constraints to which plants can respond. This is briefly addressed, with a special emphasis on the response of plant roots to deficiencies of P and Fe and to Al toxicity. Finally, soil pH itself and pH buffering capacity also have a dramatic influence on root-mediated pH changes.     

Effects of liming and mycorrhizal colonization on soil phosphate depletion and phosphate uptake by maize (Zea mays L.) and soybean (Glycine max L.) grown in two tropical acid soils

The effects of liming and inoculation with the arbuscular mycorrhizal fungus, Glomus intraradices Schenck and Smith on the uptake of phosphate (P) by maize (Zea mays L.) and soybean (Glycine max [L.] Merr.) and on depletion of inorganic phosphate fractions in rhizosphere soil (Al-P, Fe-P, and Ca-P) were studied in flat plastic containers using two acid soils, an Oxisol and an Ultisol, from Indonesia. The bulk soil pH was adjusted in both soils to 4.7, 5.6, and 6.4 by liming with different amounts of CaCO₃.In both soils, liming increased shoot dry weight, total root length, and mycorrhizal colonization of roots in the two plant species. Mycorrhizal inoculation significantly increased root dry weight in some cases, but much more markedly increased shoot dry weight and P concentration in shoot and roots, and also the calculated P uptake per unit root length. In the rhizosphere soil of mycorrhizal and non-mycorrhizal plants, the depletion of Al-P, Fe-P, and Ca-P depended in some cases on the soil pH. At all pH levels, the extent of P depletion in the rhizosphere soil was greater in mycorrhizal than in non-mycorrhizal plants. Despite these quantitative differences in exploitation of soil P, mycorrhizal roots used the same inorganic P sources as non-mycorrhizal roots. These results do not suggest that mycorrhizal roots have specific properties for P solubilization. Rather, the efficient P uptake from soil solution by the roots determines the effectiveness of the use of the different soil P sources. The results indicate also that both liming and mycorrhizal colonization are important for enhancing P uptake and plant growth in tropical acid soils.     

Aluminium accumulation in root cell walls coincides with inhibition of root growth but not with inhibition of magnesium uptake in Norway spruce

High levels of aluminium in the soil solution of forest soils cause stress to forest trees. Within the soil profile, pH and aluminium concentration in the soil solution vary considerably with soil depth. pH strongly influences the speciation of A1 in solution, and is a factor when considering toxicity of A1 to roots. Norway spruce ( Picea abies [L.] Karst.) seedlings were grown for 7 weeks in nutrient solutions at pH 3.2, 4.0 or 5.0 containing 0, 100 or 400 µ M A1. At the end of this period, seedling growth, the cation exchange capacity of the roots and the amount of exchangeable Ca and Mg in roots were determined. A1 concentrations in whole roots, root segments, and in needles were measured. Using X‐ray microanalysis, the concentrations of Al, Ca, Mg and P were determined in cortical cell walls. We wanted to test the hypotheses that (1) the amount of Al bound to cation exchange sites can be used as a marker for Al toxicity and (2) the Mg concentration of needles is controlled by the amount of Mg bound to cation exchange sites. Low pH reduced the inhibition of Al on root growth and shoot length. Both low pH and Al lowered the concentration of Ca and Mg in needles. Al concentrations in the roots decreased as the pH decreased. In the roots, Al displaced Mg and Ca from binding sites at the root cortical cell walls. A comparison of the effects of Al at the different pH values on root growth and Mg concentration in the needles, suggests that, at pH 5.0, an Al fraction in the apoplast inhibits root growth, but does not affect Mg uptake. This fraction of Al is not available for transport to the shoots. In contrast, Mg uptake is strongly affected by Al at pH 3.2, although only very low levels of Al were detected in the roots. Thus, Al accumulation in the apoplast is a positive marker for Al effects on root growth, but not Mg uptake. The Mg concentration of needles is not controlled by the amount of Mg bound to cation exchange sites.     

Relative contributions of soil chemistry, plant physiology and rhizosphere induced changes in speciation on Ni accumulation in plant shoots

The aim of this study is to rank the relative importance of soil properties, root uptake and root-to-shoot redistribution on the transfer of the trace element nickel from soil to the shoots of non hyperaccumulatings plants. Two contrasting soils and seven plant species have been studied using the radioactive isotope, ⁶³Ni. Shoots and roots were analysed separately and the specific activity of each plant has been measured. The isotopic exchange properties of rhizosphere soil where compared with control non rhizosphere soil. Possible changes in Ni speciation in the rhizosphere have been assessed by comparing the isotopic exchange properties of the rhizosphere and control soil and by comparing the specific activities of Ni in each plant. The capacity of soil to immobilise added radiotracer largely determines root uptake, leading to between a 4- and 40-fold difference between soils for a given species. The redistribution of nickel from roots to shoots was fairly constant for plants grown on the rendzina, but varied strongly between species for the acid soil. This variation enhanced the contrast between species of the soil-to-shoot transfer factor. Root action significantly enhanced immobilisation of added nickel in an acid soil due to the modification of speciation of initially non exchangeable soil nickel, but had little effect on a neutral rendzina. Changes in rhizosphere pH were similar on the two soils. In the acid soil, these pH changes were accompanied by changes in Ni speciation but a causative link has not been established. In the neutral soil pH changes may have modified root uptake properties.     

Effects of deforestation and cultivation on soil CEC and contents of exchangeable bases: A case study in Simlipal National Park,India

Deforestation in the tropics seems to be a serious problem probably because of the reduction in soil CEC and the consequent losses of nutrients from the soils. Here, changes in these parameters as influenced by deforestation as well as vegetative cover were studied; statistical methods were applied to interpret the results. Cultivation causes a significant reduction in CEC, total content of the exchangeable bases and exchangeable Ca2+ and Mg2+ levels compared to the adjoining unmanaged forest land. Levels of exchangeable K⁺ and Na⁺, however, do not change significantly. Evergreen forest soils have the highest levels of CEC, total exchangeable bases, exchangeable Ca²⁺ and K⁺. Deciduous forest, grassland and cultivated soils have statistically similar contents of exchangeable Ca²⁺, Mg²⁺, K⁺ and Na⁺. Exchangeable Mg²⁺, however, is not affected by vegetative cover. Soil CEC shows fairly good correlation with the organic carbon content only in evergreen forest soils. In others, organic carbon apparently does not influence CEC significantly. All soils show excellent correlation between their CEC and total exchangeable bases. It is concluded that for regeneration of weathered tropical soils, an evergreen cover provides the most effective means; deciduous vegetation or grass cover do not seem promising.     

Changes in nutrient distribution in forests and soils of Walker Branch watershed,Tennessee, over an eleven-year period

Changes in vegetation, litter, and soil nutrient content were measured in selected plots on Walker Branch watershed, Tennessee, from 1972–73 to 1982. The watershed has been allowed to revert to forest since 1942, before which it consisted of small subsistence farms and woodland pastures. Changes in Ca status were of particular interest because initial nutrient cycling characterizations indicated that net Ca accumulation in vegetation could have caused large decreases in soil exchangeable Ca²⁺ within 20 years.Decreases in forest floor and subsoil (45–60 cm) N, exchangeable Ca²⁺, and Mg²⁺ content were noted in several plots from 1972 to 1982. Surface soils (0–15 cm) showed either no change or, in some cases (e.g., N and exchangeable K⁺ in certain plots), increases over the 11-year period. Reductions in forest floor and subsoil exchangeable Ca²⁺ and exchangeable Mg²⁺ on cherty, upper slope oak-hickory and chestnut oak forests were most striking. The changes in Ca²⁺ are thought to be due primarily to high rates of Ca²⁺ incorporation into woody tissues of oak and hickory species. Reductions in forest floor and subsoil exchangeable Mg²⁺ could not be accounted for by woody increment; leaching may have played a major role in causing these decreases. Changes in P and exchangeable K⁺ were variable, with both increases and decreases.There were significant increases in exchangeable Al³⁺ in both subsoils and surface soils of certain plots, but these were not accompanied by decreases in exchangeable base cations or consistent decreases in pH. Dissolution of interlayer Al from 2:1 clays may be the cause of the exchangeable Al³⁺ increases.These results suggest a general decline in fertility, especially with regard to Ca and Mg in those forests with low soil Ca and Mg supplies. Monitoring of further changes (if any) in these ecosystems will continue as the currently aggrading forests approach steady state.