Immobilized EDDHA and DFOB as Iron Carriers to Cucumber Plants

Abstract

Iron chelators are the most effective Fe fertilizers known to date. However, due to their negative charge they are easily leached out of the root zone. Besides the risk of groundwater contamination with organic compounds and metals, repeated applications of expensive Fe chelates are often required. With the aim to reduce leaching, desferrioxamine B (DFOB) and ethylenediaminedihydroxyphenylacetic acid (EDDHA) were immobilized on Sepharose and tested as Fe sources to plants. Two cultivars of cucumber (Dlila and Kfir) grown in hydroponic cultures at pH 7.3, efficiently utilized Fe from immobilized FeDFOB, and immobilized FeEDDHA. In general, plant response to the immobilized fertilizers became comparable to that of soluble chelates within a period of 17 to 26 days. The kinetics of alleviating Fe induced chlorosis in plants treated with the immobilized chelates was slower than that obtained with soluble chelates. Moreover, the Fe³⁺ reduction rates obtained for immobilized FeDFOB were slower than those measured for soluble FeDFOB. Our observations suggest that immobilized FeDFOB can serve as a slow release Fe fertilizer. The slow kinetics of reduction and uptake from the immobilized as compared to the soluble chelates can be attributed to the lower accessibility to the plant's roots.     

Effects of synthetic and microbially produced chelates on the diffusion of iron and phosphorus to a simulated root in soil

Summary Hydroxamate siderophores (HS) are microbially produced, ferric-specific chelates, known to occur in soil, and to be capable of providing iron to higher plants. This study examined the potential for HS to influence the diffusion of both iron and phosphorus to plant roots in soil.The HS desferrioxamine-B (DFOB) and desferriferrichrome (ferrichrome) were compared with the synthetic chelates ethylenediamine [di(o-hydroxyphenylacetic)acid] (EDDHA) and ethylenediamine-tetraacetic acid (EDTA), and citrate, oxalate, and distilled water in their ability to increase diffusion of iron using a simulated root technique. Chelate solutions were pumped through porous fiber bundles imbedded in soil previously labeled with⁵⁵Fe. In a sandy loam of pH 7.5,⁵⁵Fe diffusion caused by 10^–4 M DFOB was twice that of water, but similar to that caused by 10^–4 M EDDHA. However, 10^–3 M EDDHA resulted in greater diffusion than 10^-3 M DFOB. The diffusions resulting from equimolar quantities of citrate, oxalate, and EDTA were similar to that with distilled water. In a clay soil of pH 5.2 previously labeled with⁵⁵Fe and³² P, the response in⁵⁵Fe diffusion to chelate treatments was: 10^–4 M EDDHA > 10^–4 M ferrichrome > 10^–3 M DFOB > 10^–4 M DFOB > water. Both ferrichrome and EDDHA caused² P diffusion to increase substantially over that of distilled water. These results suggest that hydroxamate siderophores present in the rhizosphere could effectively increase the level of soluble iron for root uptake and possibly increase phosphorus uptake by solubilization of phosphorus from iron phosphates at acid pH.     

Heavy metal sorption on soil minerals affected by the siderophore desferrioxamine B: the role of Fe(III) (hydr)oxides and dissolved Fe(III)

Phytoextraction of heavy metals from polluted soils has often been found to be limited by the bioavailability of the pollutants. Inorganic or organic ligands are occasionally used as complexing agents to enhance the mobility of the heavy metals. However, the opposite effect is also possible. We studied the influence of the hydroxamate siderophore desferrioxamine B (DFOB) on the sorption of Cu, Zn and Cd to clay minerals, with the emphasis on the role of dissolved Fe(III) and Fe(III) minerals. Depending on the surface charge of the minerals and on pH, sorption of heavy metals can be either enhanced or diminished. We show here that this effect of DFOB disappears if dissolved Fe(III) is added to suspensions of clay minerals in excess to DFOB. We found that the solid Fe(III) phases ferrihydrite and goethite did not impede the effect of DFOB on the sorption of heavy metal, however. Between pH 4 and 10, DFOB completely prevented Cu sorption on ferrihydrite. A strong mobilizing effect was also observed for Zn, but not for Cd. In presence of goethite, concentrations of dissolved Cu, Zn and Cd were enhanced only above approximately pH 5, 7 and 8, respectively. Below these pH values the binding of these metals to goethite was even stronger with than without DFOB. In the absence of heavy metals, DFOB‐promoted dissolution of ferrihydrite was much faster than that of goethite due to the larger surface area of ferrihydrite. In the alkaline pH range, where sorption of DFOB on the surfaces of the iron oxides was greater, dissolution of both minerals was reduced.     

Interaction of iron chelates with several soil materials and with a soil standard

Frequently the effectiveness of iron (Fe) chelates is low because they can be retained or destroyed by soil materials. The high cost of these Fe fertilizers makes it necessary to study soil material reaction with Fe chelates. Commercial Fe chelates with EDTA, EDDHA, and EDDHMA as ligands and their standards, prepared in the laboratory, were shaken for one hour with various soil materials [amorphous Fe(III) oxide, acid peat, calcium (Ca)‐montmorillonite and calcium carbonate (CaCO₃)] and with a soil standard made in the laboratory. After agitation, the chelate‐soil mixtures were filtered and the micronutrients and chelated Fe that remained in solution were determined. Among the soil materials used, amorphous Fe(III) oxide and acid peat had the greatest affect on the amount of chelated Fe remaining in solution. The type of chelating agent was the next major factor that affected the availability of soluble Fe following reaction with the soil materials. Another factor was the commercial formulation of the Fe chelates. The chelates comprised of EDDHA or EDDHMA maintained the highest percentages of chelated Fe in solution after interaction with the solid phases, except for the acid peat. The last soil material, acid peat, retained more chelated Fe for the Fe chelates with EDDHA or EDDHMA than with EDTA as the chelating agent. The commercial Fe‐EDDHA chelates had greater losses of chelated Fe than their standard after interaction with all the solid phases. The commercial Fe‐EDDHA chelate (Sequestrene) and the commercial Fe‐EDDHMA chelate (Hampirón) solubilized the highest amount of copper (Cu) from soil standard. This was attributed to the presence of by‐products in the commercial formulations since the Fe‐EDDHA standard did not have Cu in solution after the interaction. Therefore, the commercial Fe chelate by‐products are able to form Cu‐complexes which could affect chelated Fe and its availability to plants.     

Silicon dynamics in the rhizosphere: Connections with iron mobilization

Silicon (Si) is a beneficial element for plants as it increases their resistance to several biotic and abiotic stresses. In the rhizosphere, root exudates, especially when released by nutritionally stressed plants, promote the mineral weathering and, consequently, influence Si biogeochemistry. This study aims at evaluating the mineralogical alterations in the rhizosphere of Fe‐deficient or Fe‐sufficient barley plants grown either in a natural or in an artificial calcareous soil, focusing on the dynamics of both Fe and Si. After 6 d of soil–plant contact, X‐ray diffraction (XRD) analysis of rhizosphere soil samples of Fe‐deficient plants revealed, for both natural and artificial soil, a decrease of amorphous phases and an increase of smectite compared to the unplanted soil. Root exudates released by Fe‐deficient plants were most likely the main responsible for the weathering of the amorphous phases by a ligand controlled dissolution mechanism. When the soil–plant contact was prolonged up to 12 d, plants overcame Fe nutritional stress and their effect on soil mineralogy completely changed, as proved by the considerable increase of amorphous and decrease of smectite. Smectite decrease might evidence the effort of plant to mobilize Si and micronutrients other than Fe from the soil through the exudation of organic ligands. When the artificial soil was treated with Fe‐sufficient barley plants, the mineral weathering trend appeared reversed compared to the experiments with Fe‐deficient plants. Plant nutritional status regulates the root exudation pattern and, consequently, drives mineral weathering processes in the rhizosphere. Barley has shown to be able to mobilize Si from smectite, yet depending on its Fe supply and proving the strict connection between Si and Fe dynamics in the rhizosphere.     

Influence of pH, iron source, and Fe/ligand ratio on iron speciation in lignosulfonate complexes studied using Mössbauer spectroscopy. Implications on their fertilizer properties

Iron chlorosis is a very common nutritional disorder in plants that can be treated using iron fertilizers. Synthetic chelates have been used to correct this problem, but nowadays environmental concerns have enforced the search for new, more environmentally friendly ligands, such as lignosulfonates. In this paper, Fe coordination environment and speciation in lignosulfonate (LS) complexes prepared under different experimental conditions were studied by (57)Fe M?ssbauer spectroscopy in relation to the Fe-complexing capacities, chemical characteristics of the different products, and efficiency to provide iron in agronomic conditions. It has been observed that the complex formation between iron and lignosulfonates involves different coordination sites. When Fe(2+) is used to prepare the iron-LS product, complexes form weak adducts and are sensitive to oxidation, especially at neutral or alkaline pH. However, when Fe(3+) is used to form the complexes, both Fe(2+) and Fe(3+) are found. Reductive sugars, normally present in lignosulfonates, favor a relatively high content of Fe(2+) even in those complexes prepared using Fe(3+). The formation of amorphous ferrihydrite is also possible. With respect to the agronomical relevance of the Fe(2+)/Fe(3+) speciation provided by the M?ssbauer spectra, it seems that the strong Fe(3+)-LS complexes are preferred when they are applied to the leaf, whereas root uptake in hydroponics could be more related with the presence of weak bonding sites.     

Efficacy of an artificial microbial siderophore-Fe(III) with high redox potential on correcting Fe chlorosis in rice

ABSTRACT

Microbial siderophore-chelated Fe(III) is suggested to be an important source of Fe for plants, although it is hardly reduced by plant roots. Here, we investigated the efficacy of the easily reducible artificial microbial siderophore tris[2-{(N-acetyl-N-hydroxy)glycylamino}ethyl]amine (TAGE)-Fe(III) as an alternative Fe source to correct Fe deficiency in rice plants, and compared it to that of the natural siderophore deferoxamine B (DFOB)-Fe(III). We also evaluated the absorption of Fe from TAGE-Fe(III) by the Strategy I-like system of gramineous plants using nicotianamine aminotransferase 1 (naat1) mutant rice, which does not synthesize phytosiderophores. Fe(III)-siderophores were synthesized in vitro. Nipponbare rice and its naat1 mutant were reared in soil and gel cultures to determine Fe availability. Hydroponically grown naat1 mutant seedlings were used for reducibility assays to determine the ability of rice roots to reduce Fe(III) chelated by TAGE or DFOB. The expression of a Fe-deficiency inducible gene was also determined, as well as chlorophyll and Fe concentrations. Reduci bility assays on naat1 mutant seedlings revealed that the reduction level of TAGE-Fe(III) was approximately three times higher than that of DFOB-Fe(III). Application of TAGE-Fe(III) to both culture medium and alkaline soil improved Fe chlorosis, growth, and Fe concentration in both naat1 and wild type plants, whereas application of DFOB-Fe(III) only did so in wild type plants. Easily reducible Fe(III)-chelates such as TAGE-Fe(III) can be a better source of Fe for rice plants than most natural microbial siderophores-Fe(III). Our study also demonstrated that rice plants have the ability to utilize microbial siderophores-Fe(III) as the Fe source through the Strategy I-like Fe acquisition system.     

Nature of impurities in fertilizers containing EDDHMA/Fe(3+), EDDHSA/Fe(3+), and EDDCHA/Fe(3+) chelates.

Iron chelates derived from ethylenediaminedi(o-hydroxyphenylacetic) acid (EDDHA), ethylenediaminedi(o-hydroxy-p-methylphenylacetic) acid (EDDHMA), ethylenediaminedi(2-hydroxy-5-sulfophenylacetic) acid (EDDHSA), and ethylenediaminedi(5-carboxy-2-hydroxyphenylacetic) acid (EDDCHA) are remarkably efficient in correcting iron chlorosis in plants growing in alkaline soils. This work reports the determination of impurities in commercial samples of fertilizers containing EDDHMA/Fe(3+), EDDHSA/Fe(3+), and EDDCHA/Fe(3+). The active components (EDDHMA/Fe(3+), EDDHSA/Fe(3+), and EDDCHA/Fe(3+)) were separated easily from other compounds present in the fertilizers by HPLC. Comparison of the retention times and the UV-visible spectra of the peaks obtained from commercial EDDHSA/Fe(3+) and EDDCHA/Fe(3+) samples with those of standard solutions showed that unreacted starting materials (p-hydroxybenzenesulfonic acid and p-hydroxybenzoic acid, respectively) were always present in the commercial products. 1D and 2D NMR experiments showed that commercial fertilizers based on EDDHMA/Fe(3+) contained impurities having structures tentatively assigned to iron chelates of two isomers of EDDHMA. These findings suggest that current production processes of iron chelates used in agriculture need to be improved.     

Synthetic Iron Chelates as Substrates of Root Ferric Chelate Reductase in Green Stressed Cucumber Plants

This work studied the behavior of different iron (Fe)-chelates as substrates of ferric chelate reductase (FCR) and their ability as Fe suppliers for mildly chlorotic plants. FCR activity and Fe concentration in xylem sap were determined in green stressed cucumber plants with different stress levels using different synthetic chelates as substrates. Both reduction and Fe concentration in the xylem sap were higher for the less-stable Fe chelates, except for Fe-EDTA, which presented a relatively low Fe concentration in sap. It was concluded that a high stability of the chelate in the nutrient solution reduces the Fe reduction, but other factors, such as the complexation of the Fe(II) by the chelating agents, should be considered when the complete process of Fe uptake is studied. The use of both indexes together, i.e., FCR determination and xylem sap concentration, is useful for understanding the Fe uptake from different Fe chelates.     

Stability constant of Fe2+ chelates with soluble ligands from incubated soils

Summary The stability constants (log K) of Fe²⁺ chelates were determined on the basis of the shift in peak potential during the reduction of Fe²⁺ by a consortium of soluble ligands from incubated soils. Log K values ranged from 2.6 to 4.5. On average a change in pH of 1 unit induced a change in log K of 0.92 units. Aeration of the anaerobic decomposition products increased log K. The log K for Fe²⁺ chelates was about 0.8 units larger than that for Mn²⁺ chelates. It is considered that the chelation of ferrous iron plays an important role in the mobility and availability of iron to plants.     

Nutrient status of rhizosphere and phosphorus response of radish

Radish (Raphanus sativus L.) exhibits a high efficiency in the utilization of sparingly‐soluble phosphates. A greenhouse experiment was designed to investigate the growth response of radish to different phosphorus (P) sources and the nutrient status of the rhizosphere associated with radish growth and nutrient absorption. Radish plants were grown in pots with the roots confined in rhizobags, in such a manner that the concentration of roots was very high within the rhizobag. The rhizosphere soils and non‐rhizosphere soils were analyzed separately for active silicon (Si), aluminum (Al), iron (Fe), and manganese (Mn) using Tamm's solution and for “available”; P using the Bray P1 extraction reagent. The radish growth response was mostly attributable to phosphate amount and availability, and the lime level used in the experiment. Concentrations of active Fe, Si, Al, and Mn were reduced in the rhizosphere, especially when lime and rock phosphate (Ps) were added. Available soil P was accumulated in the rhizosphere under lime and Ps addition, whereas its concentration was reduced with the zero lime treatment. Phosphorus utilization, characterized by P accumulation in shoots, was in accordance with the concentration pattern for “available”; P in the rhizosphere, but not with the growth response of radish itself. The calcium (Ca) concentration of the shoot followed the same trend as the radish growth. There was an antagonism between potassium (K) and Ca absorption as well as between Ca and magnesium (Mg) absorption. With the addition of P, shoot Mn concentration increased, while shoot Fe and Al concentrations increased with no lime addition but decreased with lime addition. The high P efficiency of radish is discussed from the view of rhizosphere chemistry. The high Mn efficiency of radish may be influenced by the same rhizosphere processes that are involved in its high P efficiency. It was concluded that rhizosphere processes and the status of nutrients determined the nutrient efficiency of radish and thus influenced its growth response and nutrient uptake.     

有机肥对小麦根际微量元素分布的影响

A rhizolbox system was used to determine the distribution of micronutrients(Fe,Mn,Cu and Zn) across the rhizosphere of wheat (Triticum aestivum).The available contents of Fe and Mn is the rhizosphere were raised by addition of manure or chemical fertilizer combined with manure,ut those of Cu and Zn were hardly affected ,which might be an important reason why manure addition could improve the Fe and Mn nutrition status of plants,Several possible mechanisms for the increase of the availbilities of Fe and Mn in the rhizosphere due to manuring are discussed as well.     

Thermospectroscopic study of the adsorption mechanism of the hydroxamic siderophore ferrioxamine B by calcium montmorillonite

The behavior of iron-chelating agents in soils is highly affected by interactions with the solid phase. Still this aspect is frequently ignored. In this research the adsorption of the siderophore ferrioxamine B by Ca-montmorillonite, as a free ligand (desferrioxamine B, DFOB) and as a complex with Fe3+ (ferrioxamine B, FOB), was studied, using thermo X-ray diffraction (thermo-XRD) in the temperature range 25-360 degrees C and thermo-FTIR spectroscopy in the temperature range 25-170 degrees C. The effect of pH (4-7.5) on the adsorption was examined. Extensive use of curve-fitting analysis was required due to significant overlapping of the characteristic absorption bands of the various functional groups. Thermo-XRD analysis showed that both DFOB and FOB penetrated into the interlayer space of Ca-montmorillonite. FTIR results indicated strong interactions of DFOB within the interlayer, which involved all functional groups (NH3+, secondary amide groups, and hydroxamate groups). In contrast, the folded Fe complex of FOB retained its molecular configuration upon adsorption, and the basal spacing of the clay increased correspondingly. FOB interacted in the interlayer space of the clay, mainly through the NH of the secondary amide groups and NH3+, while the functional groups bound to the central Fe cation remained unchanged. The suspension pH had no significant effect on both DFOB and FOB adsorption at the examined range. Adsorption protected the adsorbates from thermal degradation compared to the nonadsorbed samples up to 105 degrees C. At 170 degrees C both DFOB and FOB were already partially degraded, but to a lesser extent than the nonadsorbed samples. Degradation of the molecules occurred mainly through the hydroxamic groups, which constitute the Fe-chelating center in the hydroxamic siderophore.     

Efficacy of Fe(o,o-EDDHA) and Fe(o,p-EDDHA) isomers in supplying Fe to strategy I plants differs in nutrient solution and calcareous soil

The FeEDDHA [iron(3+) ethylenediamine di(o-hydroxyphenylacetic) acid] is one of the most efficient iron chelates employed in the correction of iron clorosis in calcareous soils. FeEDDHA presents different positional isomers: the ortho-ortho (o,o), the ortho-para (o,p), and the para-para (p,p). Of these isomers, the p,p cannot chelate Fe in soil solution in a wide range of pH values, while both o,o and o,p can. The objective of this work was to compare the efficiency of both isomers (o,o and o,p) to provide Fe to two Strategy I plants (tomato and peach) in nutrient solution (pH approximately 6.0), as well as in calcareous soil (pH approximately 8.4; CALCIXEREPT). For this, chelates of both o,o-EDDHA and o,p-EDDHA with 57Fe (a nonradioactive isotope of Fe) were used, where the 57Fe acts as a tracer. The results obtained showed that the o,o isomer is capable of providing sufficient Fe to plants in both nutrient solution and calcareous soil. However, the o,p isomer is capable of providing sufficient Fe to plants in nutrient solution but not in calcareous soil.     

Role of Phosphatases,Iron Reducing,and Solubilizing Activity on the Nutrient Acquisition in Mixed Cropped Peanut and Barley

The effect of interspecific complementary and competitive root interactions and rhizosphere effects on primarily phosphorus (P) and iron (Fe) but also nitrogen (N), potassium (K), calcium (Ca), zinc (Zn), and manganese (Mn) nutrition between mixed cropped peanut (Arachis hypogaea L.) and barley (Hordeum vulgare L.). In order to provide more physiological evidence on the mechanisms of interspecific facilitation, phosphatase activities in plant and rhizosphere, root ferric reducing capacity (FR), Fe-solubilizing activity (Fe-SA), and rhizosphere pH were determined. The results of the experiment revealed that biomass yield of peanut and barley was decreased by associated plant species as compared to their monoculture. Rhizosphere chemistry was strongly and differentially modified by the roots of peanut and barley and their mixed culture. In the mixed cropping of peanut/barley, intracellular alkaline and acid phosphatases (AlPase and APase), root secreted acid phosphatases (S-APase), acid phosphatases activity in rhizosphere (RS-APase), and bulk soil (BS-APase) were higher than that of monocultured barley. Regardless of plant species and cropping system, the rhizosphere pH was acidified and concomitantly to this available P and Fe concentrations in the rhizosphere were also increased. The secretion Fe-solubilizing activity (Fe-SA) and ferric reducing (FR) capacity of the roots were generally higher in mixed culture relative to that in monoculture treatments which may improve Fe and Zn nutrition of peanut. Furthermore, mixed cropping improved N and K nutrition of peanut plants, while Ca nutrition was negatively affected by mixed cropping.     

Effect of humic substances in nutrient film technique on nutrient uptake

An experiment was conducted to find out how humic substances affected nutrient uptake of plants. The test plants, oregano, thyme, and basil, were grown in nutrient film technique at two pH levels (4.5 and 6.5), in two substrates (peat and perlite), and at three levels of humic substance that was a peat extract (control, low, and high concentration). Nutrient uptake of potassium (K), calcium (Ca), magnesium (Mg), iron (Fe), manganese (Mn), zinc (Zn), and copper (Cu) were determined by elemental contents in aerial parts of the plant and its weight. Humic substance had no effect on K, Ca, and Mg uptake but lowered the uptake of Fe, Mn, Zn, and Cu for the three test plants, more pronounced with perlite than peat and more at low pH than at high pH. The lowering of the uptake might be caused by complexation of Fe, Mn, Zn, and Cu by the humic substance and the lower availability of these metals in complexed form than as a cation or as EDTA‐chelate in the case of Fe. It is not clear why the effect of the humic substance on micro‐element uptake is larger at low than at high pH. The complexation is expected to be stronger at pH 6.5 than at pH 4.5. At low pH, the high concentration of humic substance caused a low fresh weight of the shoots, perhaps caused by a toxicity of the humic substance at low pH. This was less pronounced at high pH.     

Efficacy of commercial Fe(III)‐EDDHA and Fe(III)‐EDDHMA chelates to supply iron to sunflower and corn seedlings

Use of synthetic iron (Fe) chelates is the most common and effective way to treat Fe chlorosis in plants. Most commercial products contain Fe‐EDDHA or Fe‐EDDHMA but their efficacy can be quite different. Commercial products with EDDHA or EDDHMA as active components were chosen based on the data obtained by Lucena et al. (1992) in their chemical test. The chelates present extreme differences in behavior in the mentioned chemical tests. The analysis of the products revealed that the total Fe concentration is greater than the one indicated by the manufacturer in spite of a lesser amount of FeY^‐ present. The plant response to these commercial products was tested using short‐term greenhouse hydroponic cultures. Sunflower and corn were chosen because of their different behavior under Fe‐stress conditions. No significant difference between plants treated with Fe‐EDDHA or Fe‐EDDHMA chelates were observed. Since the purity index indicates there are too many differences between commercial formulations of the same type of chelate, the differentiation between groups cannot be determined with commercial products. Index I3, described by Lucena et al. (1992), does not correlate with the plant response because it did not consider the purity percentage of the products.     

Comparison of iron chelates and complexes supplied as foliar sprays and in nutrient solution to correct iron chlorosis of soybean

The application of synthetic chelates is the most efficient remedy for correcting iron (Fe) chlorosis. However, chelates are usually expensive and nondegradable products. Recently, new degradable chelates have been proposed for their use as Fe fertilizers. Also, Fe complexes cheaper than synthetic chelates and derived from natural products are also used to correct Fe deficiencies. Fifteen products, including five different synthetic chelates (Fe‐EDDS, Fe‐IDHA, and three Fe‐EDTA formulations) and ten natural complexes (humates, lignosulfonates, amino acids, glycoproteins, polyamines, citrate, and gluconate), have been compared when applied at low concentration to soybean (Glycine max L.) chlorotic plants grown in hydroponics under controlled conditions. In the first experiment, Fe compounds were applied to the nutrient solution, while in the second trial, Fe was foliar‐supplied. Dry matter, Fe concentration in shoots and roots, and SPAD values were used to evaluate the effectiveness of the Fe in the different products. In the nutrient‐solution experiment, synthetic chelates provided better plant growth, Fe concentration, and SPAD values than complexes. Among the Fe complexes, transferrin generally provided good plant responses, similar to those obtained with synthetic chelates. After foliar application, the highest regreening was observed for plants treated with synthetic chelates and amino acid complexes, but the translocation to roots only occurred for Fe lignosulfonate. Fe‐EDDS and Fe‐EDTA performed in a similar way when applied in nutrient solution or as foliar sprays.     

Plant nutritional and soil factors in relation to microbial activity in the rhizosphere,with particular emphasis on denitrification

Examples of the effect of mineral nutrition of plants (N, P, K, Mg, Fe) on microbial activity in the rhizosphere are presented with emphasis on our own studies related to root exudation, bacterial numbers, oxygen consumption and denitrification. Direct effects concern changes of pH, e.g. by liming. As the microbial community in the rhizosphere depends on decomposable organic substances released from roots, plant nutrition also indirectly affects microbial activity via its influence on plant metabolism and growth. Important factors affecting denitrification in the rhizosphere are, besides stimulation of denitrifiers by root exudation, air-filled porosity and readily decomposable organic matter content of the soil.