The dynamics of protons,aluminium, and calcium in the rhizosphere of maize cultivated in tropical acid soils: experimental study and modelling

The goals of this work were to understand the dynamics of H⁺, Al and Ca in the rhizosphere of maize cultivated in tropical acid soils, and to evaluate the contribution of the dissolution kinetics of the Al-hydroxides to Al dynamics. The study of the dissolution kinetics was based on a comparison between experimental and simulated data, using a model of the chemical processes in the rhizosphere. Two Oxisols, pH 5.1 and 4.6, and one Ultisol, pH 5.2, were studied. An Al-tolerant maize variety (Zea mays L.) was grown for 14 days on a 3-mm thick soil layer. The composition of the soil and the soil solution, together with the concentration of Al in the roots, were determined throughout the experiment. The results showed that root growth (i) decreased the soil solution pH, up to one pH unit, (ii) increased Al concentration in the soil solution, (iii) increased exchangeable Al, and (iv) decreased exchangeable Ca. Soil solution pH, exchangeable Al, and exchangeable Ca were closely linked. Exchangeable Al increased 1.5 – 3.0 times, due to the dissolution of easily mobilised Al components. In addition, Al accumulation in roots depended mainly on Al in the soil solution. Modelling the interactions between H⁺, Al, and Ca proved that the main factor determining Al in the soil solution was the kinetic reactivity of the easily mobilised Al components. These components, probably poorly crystallised Al-hydroxides, are key players in the functioning of the rhizosphere in tropical acid soils.     

Rhizotoxicity of aluminate and polycationic aluminium at high pH

Although monomeric Al species are often toxic in acidic soils, the effects of the aluminate ion (Al(OH) ₄ ⁻ ) on roots grown in alkaline media are still unclear. Dilute, alkaline (pH 9.5) nutrient solutions were used to investigate the effects of Al(OH) ₄ ⁻ on root growth of mungbean (Vigna radiata L.). Root growth was reduced by 13% after 3 d growth in solutions with an Al(OH) ₄ ⁻ activity of 16 μM and no detectable polycationic Al (Al₁₃). This decrease in root growth was associated with the formation of lesions on the root tips (due to the rupturing of the epidermal and outer cortical cells) and a slight limitation to root hair growth (particularly on the lateral roots). When roots displaying these symptoms were transferred to fresh Al(OH) ₄ ⁻ solutions for a further 12 h, no root tip lesions were observed and root hair growth on the lateral roots improved. The symptoms were similar to those induced by Al₁₃ at concentrations as low as 0.50 μM Al which are below the detection limit of the ferron method. Thus, Al(OH) ₄ ⁻ is considered to be non-toxic, with the observed reduction in root growth in solutions containing Al(OH) ₄ ⁻ due to the gradual formation of toxic Al₁₃ in the bulk nutrient solution resulting from the acidification of the alkaline nutrient solution by the plant roots.     

Root exudates as mediators of mineral acquisition in low-nutrient environments

Plant developmental processes are controlled by internal signals that depend on the adequate supply of mineral nutrients by soil to roots. Thus, the availability of nutrient elements can be a major constraint to plant growth in many environments of the world, especially the tropics where soils are extremely low in nutrients. Plants take up most mineral nutrients through the rhizosphere where micro-organisms interact with plant products in root exudates. Plant root exudates consist of a complex mixture of organic acid anions, phytosiderophores, sugars, vitamins, amino acids, purines, nucleosides, inorganic ions (e.g. HCO₃ ^–, OH^–, H⁺), gaseous molecules (CO₂, H₂), enzymes and root border cells which have major direct or indirect effects on the acquisition of mineral nutrients required for plant growth. Phenolics and aldonic acids exuded directly by roots of N₂-fixing legumes serve as major signals to Rhizobiaceae bacteria which form root nodules where N₂ is reduced to ammonia. Some of the same compounds affect development of mycorrhizal fungi that are crucial for phosphate uptake. Plants growing in low-nutrient environments also employ root exudates in ways other than as symbiotic signals to soil microbes involved in nutrient procurement. Extracellular enzymes release P from organic compounds, and several types of molecules increase iron availability through chelation. Organic acids from root exudates can solubilize unavailable soil Ca, Fe and Al phosphates. Plants growing on nitrate generally maintain electronic neutrality by releasing an excess of anions, including hydroxyl ions. Legumes, which can grow well without nitrate through the benefits of N₂ reduction in the root nodules, must release a net excess of protons. These protons can markedly lower rhizosphere pH and decrease the availability of some mineral nutrients as well as the effective functioning of some soil bacteria, such as the rhizobial bacteria themselves. Thus, environments which are naturally very acidic can pose a challenge to nutrient acquisition by plant roots, and threaten the survival of many beneficial microbes including the roots themselves. A few plants such as Rooibos tea (Aspalathus linearis L.) actively modify their rhizosphere pH by extruding OH^– and HCO₃ ^– to facilitate growth in low pH soils (pH 3 – 5). Our current understanding of how plants use root exudates to modify rhizosphere pH and the potential benefits associated with such processes are assessed in this review.     

Effects of aluminium on the peanut (Arachis hypogaea L.)/ Bradyrhizobium symbiosis

The effects of low activities of the monomeric Al species, Al³⁺, Al(OH)₂ ⁺ and Al(OH)²⁺, on the peanut/Bradyrhizobium symbiosis were examined in solution culture. In flowing solution culture, growth of the host plant was depressed at activities ≥5 μM. Neither shoot dry weight, root dry weight nor root length were inhibited by 3 μM Al, an activity which reduced nodule number by 70%. Low nodule number was compensated for, at this activity, by an increase in weight per nodule. In non-flowing solution culture of similar composition, survival of a streptomycin resistant mutant of Bradyrhizobium spp. NC92 in the bulk solution or in the rhizosphere of peanut roots was unaffected by 20 μM Al. The site of infection by Bradyrhizobium was examined by scanning electron microscopy. Lateral root axils of plants exposed to ≥2 μM Al did not display the rosette of multicellular root hairs which is characteristic in normal plants. The detrimental effects of Al on nodulation appear to be related to structural changes at the site of infection which are observed at Al activities too low to cause any depression in growth of the host plant, including root length, and at activities of Al which do not affect survival of the free-living Bradyrhizobium.     

Effects of aluminium on growth and root reactions of phosphorus stressed Betula pendula seedlings

The impact of two constant non-toxic levels of Al stress (0.2 and 0.4 mM) on growth and ³²P uptake capacity on sub-optimal (P-limited) Betula pendula seedlings grown in sand culture was examined.Seedling growth was under optimum controlled conditions in a growth chamber where nutrient additions were made at a predetermined relative addition rate (R_A) of 10% day^-1. Three treatment groups of seedlings 0, 0.2 and 0.4 mM Al were harvested at 15, 29 and 42 days. The excised roots were exposed to a ³²P-labelled solution for 15 minutes to measure their capacity for P uptake. Growth was determined by weighing the roots, stems and leaves of the seedlings.Growth data showed that relative growth rate (R_G) should equal the R_A of P the most limiting nutrient, which was supplied at P/N 3% instead of an optimal 15%. Therefore, Ingestad's theory can also be used succesfully in sand culture and this may be particularly important for future studies of root and rhizosphere exudates. Low levels of Al (< 0.2 mM) in combination with low P supply significantly lowered the R_G of the birch seedlings by further reducing P supply. However, previous studies of birch seedling growth and nutrient uptake using Ingestad's solution culture technique with optimumal P supply did not show any effect of Al on growth untill the Al was in excess of 3 mM. Aluminium was not directly toxic to the plants and therefore roots could respond to the ³²P bioassay.     

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.     

Root morphological and proteomic responses to growth restriction in maize plants supplied with sufficient N

The primary objective of this study was to better understand how root morphological alteration stimulates N uptake in maize plants after root growth restriction, by investigating the changes in length and number of lateral roots, ¹⁵NO₃⁻ influx, the expression level of the low-affinity Nitrate transporter ZmNrt1.1, and proteomic composition of primary roots. Maize seedlings were hydroponically cultured with three different types of root systems: an intact root system, embryonic roots only, or primary roots only. In spite of sufficient N supply, root growth restriction stimulated compensatory growth of remaining roots, as indicated by the increased lateral root number and root density. On the other hand, there was no significant difference in ¹⁵NO₃⁻ influx between control and primary root plants; neither in ZmNrt1.1 expression levels in primary roots of different treatments. Our data suggested that increased N uptake by maize seedlings experiencing root growth restriction is attributed to root morphological adaptation, rather than explained by the variation in N uptake activity. Eight proteins were differentially accumulated in embryonic and primary root plants compared to control plants. These differentially accumulated proteins were closely related to signal transduction and increased root growth.     

Carbon distribution in grass (Festuca pratensis L.) during regrowth after cutting—utilization of stored and newly assimilated carbon

Distribution of net assimilated C in meadow fescue (Fectuca pratensi L.) was followed before and after cutting of the shoots. Plants were continuously labelled in a growth chamber with ¹⁴C-labelled CO₂ in the atmosphere from seedling to cutting and with ¹³C-labelled CO₂ in the atmosphere during regrowth after the cutting. Labelled C, both ¹⁴C and ¹³C, was determined at the end of the two growth periods in shoots, crowns, roots, soil and rhizosphere respiration. Distribution of net assimilated C followed almost the same pattern at the end of the two growth periods, i.e. at the end of the ¹⁴C- and the ¹³C-labelling periods. Shoots retained 71–73% of net assimilated C while 9% was detected in the roots and 11–14% was released from the roots, determined as labelled C in soil and as rhizosphere respiration. At the end of the 2nd growth period, after cutting and regrowth, 21% of the residual plant ¹⁴C at cutting (¹⁴C in crowns and roots) was found in the new shoot biomass. A minor part of the residual plant ¹⁴C, 12%, was lost from the plants. The decreases in ¹⁴C in crowns and roots during the regrowth period suggest that ¹⁴C in both crowns and roots was translocated to new shoot tissue. Approximately half of the total root C at the end of the regrowth period after cutting was ¹³C-labelled C and thus represents new root growth. Root death after cutting could not be determined in this experiment, since the decline in root ¹⁴C during the regrowth period may also be assigned to root respiration, root exudation and translocation to the shoots. ei]{gnH}{fnLambers} ei]{gnA C}{fnBorstlap}     

Localization and quantification of net fluxes of H+ along maize roots by combined use of pH-indicator dye videodensitometry and H+-selective microelectrodes

Two methods for measuring proton fluxes along intact maize roots grown with NH ₄ ⁺ or NO ₃ ⁻ at pH 6.5 were compared. Videodensitometric measurement of changes in a pH-indicator dye by video camera was used to map pH around roots and determine the amounts of protons released by various root regions. This method was compared with potentiometric determination of the concentration of H⁺ in the unstirred layer at the root surface using ion-selective microelectrodes. With NH ₄ ⁺ the roots released large amounts of H⁺ in preferential regions where the rate of flux can reach 1.4 or even 2.5 nmol m⁻¹ s⁻¹. Videodensitometry indicated a first region of root acidification in the subapical zone, but this was more difficult to localize with microelectrodes. With NO₃ ⁻ both methods showed that the roots released small amounts of H⁺ and that the apical region took up H⁺ in the first 10 mm then sometimes released H⁺ over the following 10 mm of root. The H⁺ flux profiles obtained by both methods were in good agreement in terms of both order of magnitude of the fluxes and spatial differences along the root. These results suggest that videodensitometry, which is easier to use than potentiometry, can be used to screen different plant species or cultivars under various experimental conditions. The microelectrode technique is indispensable, however, for studying the underlying mechanisms of net H⁺ fluxes. This revised version was published online in June 2006 with corrections to the Cover Date.     

Mobilization of aluminium in the rhizosphere soil solution of growing tree roots in an acidic soil

Chemical conditions in the rhizosphere in many respects are different from the bulk soil. Especially in acid forest soils aluminium chemistry at the soil root interface is of particular interest because of its importance for evaluating the risk of rhizotoxicity. In the present study we have used micro suction cups to collect soil solution from the rhizosphere of oak seedlings (Quercus robur L.) in high spatial resolution and capillary electrophoresis for the determination of major ions and Al³⁺. While the concentrations of nutrient cations, especially Ca²⁺ and Mg²⁺, decreased in the vicinity of growing roots the concentrations of Al³⁺ significantly increased. Al³⁺-ions were probably released when root-exuded protons were buffered by the soil. Their occurrence indicates, that the oak roots in our experiments had only limited capabilities to detoxify Al in their rhizosphere. The restriction of this effect to the very small soil compartment close to the roots suggests, that common soil analysis which neglect rhizosphere processes might greatly underestimate the in situ concentration of Al³⁺ near tree roots. Our experiments furthermore indicate, that also suberized roots have a significant influence on rhizosphere soil solution chemistry. This revised version was published online in June 2006 with corrections to the Cover Date.     

Response of the plant root to aluminum stress: Analysis of the inhibition of the root elongation and changes in membrane function

Pea root elongation was strongly inhibited in the presence of a low concentration of Al (5 μM). In Al-treated root, the epidermis was markedly injured and characterized by an irregular layer of cells of the root surface. Approximately 30% of total absorbed Al accumulated in the root tip and Al therein was found to cause the inhibition of whole root elongation. Increasing concentrations of Ca²⁺ effectively ameliorated the inhibition of root elongation by Al and 1 mM of CaCl₂ completely repressed the inhibition of root elongation by 50 μM Al. The ameliorating effect of Ca²⁺ was due to the reduction of Al uptake. H⁺-ATPase and H⁺-PPase activity as well as ATP and PPidependent H⁺ transport activity of vacuolar membrane vesicles prepared from barley roots increased to a similar extent by the treatment with 50 μM AlCl₃. The rate of increase of the amount of H⁺-ATPase and H⁺-PPase was proportional to that of protein content measured by immunoblot analysis with antibodies against the catalytic subunit of the vacuolar H⁺-ATPase and H⁺-PPase of mung bean. The increase of both activities was discussed in relation to the physiological tolerance mechanism of barley root against Al stress.     

Bacterial inoculation of maize affects carbon allocation to roots and carbon turnover in the rhizosphere

To assess the influence of bacteria inoculation on carbon flow through maize plant and rhizosphere,¹⁴C allocation after¹⁴CO₂ application to shoots over a 5-day period was determined. Plants were grown on C- and N-free quartz sand in two-compartment pots, separating root and shoot space. While one treatment remained uninoculated, treatments two and three were inoculated withPantoea agglomerans (D5/23) andPseudomonas fluorescens (Ps I A12), respectively, five days after planting. Bacterial inoculation had profound impacts on carbon distribution within the system. Root/rhizosphere respiration was increased and more carbon was allocated to roots of plants being inoculated. After five days of¹⁴CO₂ application, more ethanol-soluble substances were found in roots of inoculated treatments and lower rhizodeposition indicated intensive C turnover in the rhizosphere. In both inoculated treatments the intensity of photosynthesis measured as net-CO₂-assimilation rates were increased when compared to the uninoculated plants. However, high C turnover in the rhizosphere reduced shoot growth of D5/23 inoculated plants, with no effect on shoot growth of Ps I A12 inoculated plants. A separation of labeled compounds in roots and rhizodeposition revealed that neutral substances (sugars) constituted the largest fraction. The relative fractions of sugars, amino acids and organic acids in roots and rhizodeposition suggest that amino acid exudation was particularly stimulated by bacterial inoculation and that turnover of this substance group is high in the rhizosphere.     

Autotoxic Ginsenosides in the Rhizosphere Contribute to the Replant Failure of Panax notoginseng

MethodsThe autotoxicities were measured using seedling emergence bioassays and root cell vigor staining. The ginsenosides in the roots, soils, and root exudates were identified with HPLC-MS.ResultsThe seedling emergence and survival rate decreased significantly with the continuous number of planting years from one to three years. The root exudates, root extracts, and extracts from consecutively cultivated soils also showed significant autotoxicity against seedling emergence and growth. Ginsenosides, including R₁, Rg₁, Re, Rb₁, Rb₃, Rg₂, and Rd, were identified in the roots and consecutively cultivated soil. The ginsenosides, Rg₁, Re, Rg₂, and Rd, were identified in the root exudates. Furthermore, the ginsenosides, R₁, Rg₁, Re, Rg₂, and Rd, caused autotoxicity against seedling emergence and growth and root cell vigor at a concentration of 1.0 µg/mL.ConclusionOur results demonstrated that autotoxicity results in replant failure of Sanqi ginseng. While Sanqi ginseng consecutively cultivated, some ginsenosides can accumulate in rhizosphere soils through root exudates or root decomposition, which impedes seedling emergence and growth.     

Excessive aluminium accumulation in the pea mutant E107 (brz)

E107 is a pleiotropic mutant of peaPisum sativum cv. ‘Sparkle’, characterized by forming few nodules and developing bronze necrotic spots on older leaves. The mutant accumulates Al and has symptoms typical of Al toxicity. The lateral roots of E107 are fewer (40%) and shorter (50%) than those of its parent. High concentrations of Al accumulate in E107 shoots (1000 mg kg^-1) and roots (3000 mg kg^-1), and three-week old E107 plants extrude 2.5 times more protons than ‘Sparkle’ plants of similar age. Al concentrations of the roots of the mutant and of its parent ‘Sparkle’ are similar for the first two weeks of growth. Thereafter they differ. In 2 week old plants Al continues to accumulate in excessive amounts in E107 primary and lateral roots whereas in ‘Sparkle’ roots, it reaches a plateau. In E107, Al is erratically distributed in the walls of root hairs and epidermal cells in both primary and lateral roots. Some of these cells have also Al in their nucleus.     

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.     

Agarose as a suitable substrate for use in the study of Al dynamics in the rhizosphere

Because of experimental difficulties, few authors have studied the dynamics of aluminium in the rhizosphere. The aim of this paper is to present a suitable method for studying rhizosphere Al dynamics. It is based on the use of agarose as a substrate for plant growth. Agar and agarose gels are often used in rhizosphere studies, but most are poorly characterized and occasionally give rise to experimental artefacts, especially with low mobility elements like Al. The results reported here show that agarose is a relatively pure substrate, nearly devoid of phosphorus and other Al-complexing substances. Aqueous extracts of agarose also exhibit Al phytotoxicity equivalent to that of a nutrient solution. Since this substrate has the properties of a variable charge exchange complex, it can be considered as a physico-chemical model for organic matter. Finally, its Al adsorption capacity is high enough for the Al reserve in the substrate not to exert a limiting effect on plants and low enough to allow accurate measurement of Al depletion in the rhizophere.     

Ammonium versus Nitrate Nutrition of Plants Stimulates Microbial Activity in the Rhizosphere

Using an alkaline calcareous soil, pot experiments were conducted to elucidate the effects of NH ₄ ⁺ vs. NO ₃ ⁻ nutrition (50 or 100 mg kg⁻¹ soil) of wheat and maize on microbial activity in the rhizosphere and bulk soils. Dicyandiamide was used as nitrification inhibitor to maintain NH ₄ ⁺ as the predominant N source for plants grown in NH ₄ ⁺ -treated soil. While maize grew equally well on both N sources, root and shoot growth of wheat was higher under NH ₄ ⁺ than under NO ₃ ⁻ nutrition. Bacterial population density on roots, but not in the rhizosphere soil, was higher under NH ₄ ⁺ than under NO ₃ ^– supplied at 150 mg N kg⁻¹ soil; whereas at both N levels applied, NH ₄ ⁺ compared to NO ₃ ⁻ nutrition of wheat and maize significantly increased microbial biomass in the rhizosphere soil. Under both plant species, NH ₄ ⁺ vs. NO ₃ ⁻ nutrition also increased aerobic and anaerobic respiration, and dehydrogenase activity in the rhizosphere. As microbial activity in the planted bulk and unplanted soils was hardly affected by the N-source, we hypothesize that the stimulation by NH ₄ ⁺ of the rhizosphere microbial activity was probably due to higher availability of root exudates under NH ₄ ⁺ than under NO ₃ ⁻ nutrition.     

Photosynthesis controls of CO2 efflux from maize rhizosphere

The effects of different shading periods of maize plants on rhizosphere respiration and soil organic matter decomposition were investigated by using a ¹³C natural abundance and ¹⁴C pulse labeling simultaneously. ¹³C was a tracer for total C assimilated by maize during the whole growth period, and ¹⁴C was a tracer for recently assimilated C. CO₂ efflux from bare soil was 4 times less than the total CO₂ efflux from planted soil under normal lighting. Comparing to the normal lighting control (12/12 h day/night), eight days with reduced photosynthesis (12/36 h day/night period) and strongly reduced photosynthesis (12/84 h day/night period) resulted in 39% and 68% decrease of the total CO₂ efflux from soil, respectively. The analysis of ¹³C natural abundance showed that root-derived CO₂ efflux accounted for 82%, 68% and 56% of total CO₂ efflux from the planted soil with normal, prolonged and strongly prolonged night periods, respectively. Clear diurnal dynamics of the total CO₂ efflux from soil with normal day-night period as well as its strong reduction by prolonged night period indicated tight coupling with plant photosynthetic activity. The light-on events after prolonged dark periods led to increases of root-derived and therefore of total CO₂ efflux from soil. Any factor affecting photosynthesis, or substrate supply to roots and rhizosphere microorganisms, is an important determinant of root-derived CO₂ efflux, and thereby, total CO₂ efflux from soils. ¹⁴C labeling of plants before the first light treatment did not show any significant differences in the ¹⁴CO₂ respired in the rhizosphere between different dark periods because the assimilate level in the plants was high. Second labeling, conducted after prolonged night phases, showed higher contribution of recently assimilated C (¹⁴C) to the root-derived CO₂ efflux by shaded plants. Results from ¹³C natural abundance showed that the cultivation of maize on Chromic Luvisol decreased soil organic matter (SOM) mineralization compared to unplanted soil (negative priming effect). A more important finding is the observed tight coupling of the negative rhizosphere effect on SOM decomposition with photosynthesis.     

Nitrogen retranslocation in plants of maize,lupin and cocklebur

Summary In a split root experiment translocation of N from shoot to root was studied using¹⁵NO ₃ ^– . The three plant species selected for this experiment differed significantly with respect to root NRA. For lupin, maize and cocklebur about 80, 50 and 6% of all absorbed NO ₃ ^– was assmilated in the roots, respectively.Although NO ₃ ^– was reduced in the roots of lupin and maize plants to a greater extent than required for the roots' demand for organic N, a significant phloem flow of N from shoot to roots was found in these plants. Unexpectedly, for cocklebur, the plant with the very low root NRA, the fraction of total N present in the root that has been imported from the shoot was only half that as found for lupin and maize.     

Altered cell wall properties are responsible for ammonium‐reduced aluminium accumulation in rice roots

The phytotoxicity of aluminium (Al) ions can be alleviated by ammonium (NH₄⁺) in rice and this effect has been attributed to the decreased Al accumulation in the roots. Here, the effects of different nitrogen forms on cell wall properties were compared in two rice cultivars differing in Al tolerance. An in vitro Al‐binding assay revealed that neither NH₄⁺ nor NO₃^? altered the Al‐binding capacity of cell walls, which were extracted from plants not previously exposed to N sources. However, cell walls extracted from NH₄⁺‐supplied roots displayed lower Al‐binding capacity than those from NO₃^?‐supplied roots when grown in non‐buffered solutions. Fourier‐transform infrared microspectroscopy analysis revealed that, compared with NO₃^?‐supplied roots, NH₄⁺‐supplied roots possessed fewer Al‐binding groups (‐OH and COO‐) and lower contents of pectin and hemicellulose. However, when grown in pH‐buffered solutions, these differences in the cell wall properties were not observed. Further analysis showed that the Al‐binding capacity and properties of cell walls were also altered by pHs alone. Taken together, our results indicate that the NH₄⁺‐reduced Al accumulation was attributed to the altered cell wall properties triggered by pH decrease due to NH₄⁺ uptake rather than direct competition for the cell wall binding sites between Al³⁺ and NH₄⁺.