Beziehung zwischen Kationen/Anionen-Aufnahme von Rotklee und Protonenabscheidung der Wurzeln

Relationship between the cation/anion uptake and the release of protons by roots of red clover Red Clover was cultivated in Mitscherlich pots on a brown podzolic soil. Besides a low N rate at the beginning of the experiment, the clover received only symbiontically fixed N. Soil pH dropped under clover from 7.2 to 4.5 during a period of 14 months. During this time seven cuts were obtained. In a parallel pot experiment with rye-grass grown on the same soil and under the same environmental conditions but supplied with NH₄NO₃ after each clipping, no drop in soil pH was observed. In the aerial plant parts of clover the cation excess was high and amounted to about 60 % of the H⁺ quantity required for the decrease in soil pH from 7.2 to 4.5. It is concluded that the cations taken up in excess were electrostatically balanced by an equivalent amount of protons secreted by the roots into the soil. The alcalinity assessed in the upper plant parts of clover was approximately equivalent to the cation excess. It is therefore supposed that the H⁺ release from roots resulted in an alkalinization in the plant cells which in return led to a synthesis of organic anions being equivalent to the amount of H⁺ released.     

Der Einfluß gesteigerter Stickstoffgaben auf die Entwicklung der Wurzelknöllchen,auf die symbiontische Stickstoffassimilation sowie auf das Wachstum und den Ertrag von Ackerbohnen (Vicia faba L.)

The effect of increasing nitrogen fertilizer rates on the development of root nodules, on the symbiotic N₂ assimilation, and on growth and yield of broad beans (Vicia faba L.) In pot experiments with broad beans (Vicia faba minor) the effect of increasing N fertilizer rates (0–800 mg N/pot) on nodulation, symbiotic N assimilation, plant growth, and yield has been studied. The plants were harvested at the beginning and at the end of flowering and at maturity. The following results were obtained: 1. No significant yield differences between treatments were found at the 1st and 2nd harvest, with exception of the 800 mg N/pot treatment which gave higher root yields at the first harvest. 2. At the 1st harvest (beginning of flowering) root nodule yield of the N-zero treatment was higher than the root nodule yield of the N treated plants. At later stages, however, no major differences in root nodule yield between the treatments N₁₀₀ and N₂₀₀ and the N-zero treatment were obtained. 3. Highest grain yields were obtained in the treatment with the lowest N-rate (100 mg N/pot) and in the treatment with the highest N rate (800 mg N/pot). The absolute highest amount of symbiotically fixed N was produced in the low N treatment (100 mg N/pot). Provided that the soil is low in available N a low fertilizer rate is required in order to exploit the full N₂ assimilation potential. 4. The total soil N remained fairly constant throughout the growing period.     

Vernachlässigte Phosphor‐ und Kaliumdüngung im ökologischen Landbau senkt die biologische Stickstofffixierung bei Rotklee und den Kornertrag bei nachfolgendem Hafer

Neglected P and K fertilization in organic farming reduces N₂ fixation and grain yield in a red clover‐oat rotation N₂ fixation is the most important N source in organic farming. An insufficient P, K, and S supply to legumes may reduce their N₂ fixation capacity. Consequently, the total yield of plant production may also be reduced. This problem was studied in a pot experiment with red clover followed by oat. Soil was taken from a field where organic farming had been practiced for more than 30 years without applying any mineral fertilizers or buying additional fodder. The soil (luvisol from loess) was characterized by: pH (CaCl₂) 5.4; lactate‐soluble (CAL) P 5 mg kg^–1 and K 110 mg kg^–1. 6 kg dry soil were mixed with 400 mg P applied as (i) triplesuperphosphate (TSP), (ii) rock phosphate (RP) or (iii) compost from organic household residues (BAK). An additional treatment (iv) with TSP received 1000 mg K as K₂SO₄ (TSP+K) and an additional treatment with RP (v) received only 200 mg P (RP/2). A control treatment received no fertilizer. P application significantly improved the P nutritional status of the plants (P content) and increased the N amount in the shoots of red clover (with 400 mg P per pot by 64 % to 139 % as compared to the control) and the dry matter (DM) yield by 60 % to 130 %. No significant differences between TSP and RP were found. The application of BAK resulted in a significantly higher N yield than the application of RP and TSP. The treatment TSP+K resulted in the highest DM yield (230 %), removal of P was 343 %, of K 228 %, and of N 239 % as compared to the control plants. This indicates a synergistic effect of P, K, and S on N₂ fixation, which was also found with BAK. Oat grown after red clover increased its grain yield by 132 % (200 mg P as RP) to 165 % (400 mg P treatments). This was mainly due to a higher P uptake (up to 172 %) and a higher N uptake (up to 172 %) as compared to the control.     

N2-fixing bacteria in the rhizosphere: Quantification and hormonal effects on root development

The paper summarizes the results of a series of experiments on enumeration of N₂-fixing bacteria (diazotrophs) and hormonal effects of Azospirillum on root development. Numbers of N₂-fixing and N-heterotrophic bacteria were determined on the root (rhizoplane plus “inner” root surface) and in the rhizosphere soil (0–3 mm from the root surface) of Arrhenatherum elatius, other forage grasses and some herbaceous plant species. Pot experiments involved freshly collected soil from an unfertilized grassland area containing its natural population of N₂-fixing bacteria. The MPN (most probable number) of diazotrophs in relation to the MPN of the total bacterial population was always lower on the root than in the rhizosphere soil, suggesting that diazotrophs were not selectively advantaged at the root surface. Supply of mineral nitrogen (NH₄NO₃) decreased the proportion of N₂-fixing bacteria at the rhizoplane as well as in the rhizosphere soil. Similar results were obtained when N was supplied via the leaves. The data suggest that N₂-fixing bacteria in the rhizosphere are poor competitors once they loose their competitive advantage of binding dinitrogen. Correspondingly, the increase in the MPN of the diazotrophs found during plant development was interpreted as a result of decreased available combined N in the rhizosphere. The proportion of N₂-fixing bacteria relative to the total number of bacteria was generally below 1%. Considering the potential amount of substrate released from the roots and the substrate requirement of the bacterial population, N₂-fixation was considered insignificant for plant growth under the given conditions. For the investigations on possible beneficial effects on plant development by bacterial hormones, Azospirillum brasilense was chosen because evidence suggests that amongst the soil bacteria releasing hormones, especially IAA, certain strains of this species are more important than other bacteria. Application of A. brasilense Cd (ATCC 29710) onto the roots of young wheat plants grown in soil increased the number of lateral roots, the total root length and the number of root hairs. Similar results were obtained after application of IAA. This suggests that IAA is an important factor responsible for the effects observed after inoculation with A. brasilense. The increase in root surface may improve acquisition of nutrients and enhance growth of plants. Another hormonal effect of A. brasilense was an increase in nodulation of Medicago sativa grown on agar. Again pure IAA resulted in a similar increase in nodule number. Increases in nodule number were only in part associated with a change in root morphology. Therefore an effect of IAA on the plant immanent regulation system for nodulation is likely.     

Acidification in the Rhizosphere of Rape Seedlings and in Bulk Soil by Nitrification and Ammonium Uptake

Ammonium salts used as fertilizers may cause soil acidification by two different processes: nitrification in soil and net release of protons from roots. Their influence on soil pH may vary depending on the distance from root surface. The aim of this study was to distinguish between these two processes. For this purpose rape seedlings were grown 10 d in a system which separated roots from soil by a fine-meshed screen. As a function of distance from the plane root layer formed on the screen, pH, titratable and exchangeable acidity and NO₃- and NH₄-nitrogen were determined. The soil, a luvisol from loess, was supplied with no N or (NH₄)₂SO₄ either with or without a nitrification inhibitor (DCD). The bulk soil pH remained unaffected when no N or 400 mg NH₄? N kg^?1 soil plus DCD was applied but it decreased from 6.6 to 5.8 without DCD. In contrast, rhizosphere pH decreased in all cases, mainly within a distance of 1 mm from the root plane only, but with gradients extending to between 2 and 4 mm into the soil. The strongest pH decrease, from 6.6 to 4.9, occurred at the root surface of plants treated with both NH₄-N and DCD where most of the mineral N remained as ammonium. In this case Al was solubilized in the rhizosphere as indicated by exchangeable acidity. Total soil acidity produced in the NH₄ treatment without DCD was mainly derived from nitrification compared to root released protons. However, acidification of the rhizosphere was diminished by nitrification because nitrate ions taken up by the roots counteracted net proton release. It is concluded that nitrification inhibitors may reduce proton input from ammonium fertilizers but enhance acidification at the soil-root interface which may cause Al toxicity to plants.     

Nitrogen fixation in paddy soils

Several important features of the N. fixation in paddy fields which were reported previously were confirmed and some new additional results regarding the evaluation of the N₂ fixation in the rhizosphere were obtained by reinvestigation in the fields. In addition, rice plants were cultivated in the submerged soil in pots and various parts of the soil were analyzed for the N₂-fixing activity as well as several other properties. The results of the pot experiments were found to be fairly similar to those observed in the field investigations, indicating the validity of the submerged soil in a pot as a rather simulated model for the actual paddy field. By using this model system, the following facts were ascertained: (1) Water-percolation had almost no effect on the N₂-fixing activities of both the rhizosphere and the non-rhizosphere soils. (2) Suppressing effect of washing the root of rice plant on the N₂-fixing activity was slight in the seedling stage and marked in the tillering and flowering stages. (3) The N₂-fixing activity of a single rice root varied from tip to base.     

Slow-release 15N fertilizer formulations to measure N2-fixation by isotope dilution

Pot experiments were carried out to examine the effects of slow-release fertilizer formulations on estimates of N₂-fixation determined by the isotope dilution method. Soybeans were used as the N₂-fixing plants, with non-nodulated soybeans and maize as the non-fixing controls. The ¹⁵N-fertilizer formulations used were (¹⁵NH₄)SO₄, K¹⁵NO₃, gypsum-pelleted K¹⁵NO₃, (¹⁵NH₄)₂SO₄ + glucose, ground plant material enriched with ¹⁵N or ¹⁵N-oxamide. The estimate of the amount of N₂ fixed by the nodulated soybean plants depended upon both the control plant and the fertilizer formulation used. Maize took up N later than non-nodulated soybean and estimates of soil N-pool (soil “A” value + fertilizer N added) calculated from the enrichment of this control were about twice as large as those calculated from the enrichment of non-nodulated soybean receiving the same fertilizer treatment. As a consequence, estimates of N₂-fixation relative to this control were lower than those relative to non-nodulating soybean (mean 140 mg N per pot compared with 292 mg N per pot). With unstablilized ¹⁵N salts errors were sufficient to produce negative estimates of fixation relative to maize. Even with a “well-matched” control (non-nodulated soybean) estimates of fixation varied with fertilizer formulation.     

Depth of root symbiont occurrence in soil

Summary The woody legume Prosopis glandulosa (mesquite) growing in the California Sonoran Desert develops functional root symbiotic associations (N₂-fixing nodules, vesicular-arbuscular mycorrhizal fungi) at depths greater than 4 m in moist soil above a seasonally stable water table. Population densities of symbiotic microorganisms are substantially greater at depth than near the surface. Inferences of plant symbiotic dependence based upon examination of surface roots and soil may be incorrect since deep roots can support the symbioses which are critical for plants utilizing deep water.     

Einfluß der NO3- bzw. NH4-Ernährung auf Ertrag und Nitratgehalt von Spinat und Kopfsalat

Influence of NO₃ and NH₄-nutrition on yield and nitrate content of spinach and lettuce Field and pot experiments were conducted in order to study the effect of N-supply (N_min + N-fertilization) and partial and complete NH₄-nutrition on yield and nitrate content of spinach and lettuce. The highest yields were obtained when the N-supply of spring spinach was 250 kg N/ha, autumn spinach 200 kg N/ha and lettuce 120 kg N/ha. Nitrate content of plants could be diminished by reducing N-supply. However, if N-supply is below the optimum value yield depressions are inevitable. NH₄-nutrition reduced in both species nitrate content. But, spinach yield was reduced heavily. Very low nitrate content of plants occured, when N-nutrition of plants cultivated in nutrient solution, was changed from NO₃ to NH₄ during the last days before harvest. Herewith similar weights and head quality were obtained as with permanent NO₃-nutrition.     

Effect of dinoseb on nitrogen fixation of red clover (Trifolium pratense)

The effect of the herbicide dinoseb (2-sec-butyl-4,6-dinitrophenol) on the symbiotic nitrogen fixation of red clover (Trifolium pratense cv. Venla) was studied in field, greenhouse and laboratory experiments. In the field, the herbicide reduced the levels of nitrogenase (C₂H₂) activity of the plants up to 18 days after treatment, but increased the yields compared to hand-weeded and unweeded controls. Rhizobium strains isolated from root nodules of plants treated with dinoseb showed resistance to the herbicide when grown on laboratory media containing dinoseb. However, in a pot experiment, inoculation with a strain which grew in the presence of 200 μg dinoseb ml^?1 did not improve the performance of red clover treated with dinoseb. In the pot experiment dinoseb inhibited nitrogenase activity when sprayed on the leaves, but not when added to the growth medium. Application of starter nitrogen did not influence the effect of dinoseb. The results suggest that dinoseb did not act directly on the nodules, but affected nitrogen fixation by damaging the photosynthetic parts of the plant.     

Increased phosphorus availability from sewage sludge ashes to maize in a crop rotation with clover

A recycling of Phosphorus (P) from the human food chain is mandatory to secure the future P supply for food production. However, many available recycled P fertilizers from sewage sludge do not have an adequate P bioavailability and, thus, are not suitable for their application in soils with pH >5.5–6.0, unless being combined with efficient mobilization measures. The aim of the study was to test the P mobilization ability of red clover (Trifolium pratense L.) from two thermally recycled P fertilizers for a subsequently grown maize. Two sewage sludge ashes (SSA) were investigated in a pot experiment at soil pH 7.5 with red clover differing in its nitrogen (N) supply (added N fertilizer or biological N₂ fixation (BNF)), followed by maize (Zea maize L.). Shoot dry matter of maize was almost doubled when N supply of previous grown clover was covered by BNF, instead of receiving added N fertilizer. Similarly, shoot P removal of maize following clover with BNF was significantly increased. It is suggested that the P mobilization is related to the BNF, and a proton release of N₂ fixing clover roots led to the measured decrease in soil pH and thereby increased P availability of the tested fertilizers.     

Wurzelstudien und Phosphat-Aufnahme von Weidelgras und Rotklee unter Feldbedingungen

Root studies and phosphorus uptake of rye-grass and red clover under field conditions Root parameters (fresh weight, density, surface, length, cation exchange capacity) and phosphate uptake were studied with rye grass and red clover, grown in the field on a brown podsolic soil. In all root parameters, ry grass was superior to red clover. Also, phosphate uptake of rye grass was higher than that of red clover. The greatest difference between both species was found in root length, that of rye grass being about five times longer than that of red clover. Rye grass had longer root hairs than red clover; whereas root diameter of clover was about twice as thick the average rye grass. Significant correlations were observed between root parameters and phosphate uptake in the plants studied. The highest correlation coefficients were obtained for the relationship P-uptake versus root length (clover 0.91***, grass 0.87***) and P-uptake versus root fresh weight (clover 0.92***, grass 0.88***). The phosphate uptake per unit root parameters was significantly higher in red clover, compared with rye grass, for the parameters root fresh weight, cation exchange capacity and root length. Because of this high P-uptake rate for clover it is assumed that clover also requires a higher P-concentration in soil solution as compared with grass. Thus grass may still grow with low P concentrations in the soil solution without P deficiency at which clover cannot grow. It is for this reason that in mixed swards clover is depressed by grass, if the available P in the soil is low.     

Vergleich von Elektro-Ultrafiltration (EUF) und Extraktion mit 0,01 molarer CaCl2-Lösung zur Bestimmung des pflanzenverfügbaren Stickstoffs im Boden

Comparison of electro-ultrafiltration (EUF) and extraction by 0.01 M CaCl₂ for the determination of “available” soil nitrogen The NO₃-N, NH₄-N and N_org content of 25 different farm soils were analysed using the EUF-technique and CaCl₂ extraction in order to compare the reproducibility and reliability of the two methods. Nitrogen available to plants was measured in a micro pot experiment with Lolium multiflorum. - CaCl₂ values were better reproducibel (cv = 2.6%) than those of the EUF method (cv = 6.2%). - Close correlations were found between the soil content of CaCl₂-and EUF-extractable nitrogen (r² = 0.917). - The EUF-extractable N_org-fraction was positivly correlated with the total N-uptake by the test plants (r = 0.921). Most importently these data did not significantly deviate from the correlation of the CaCl₂ extractable N_org-fraction with total N-uptake (r = 0.932).     

Nitrat‐Austräge auf intensiv und extensiv beweidetem Grünland,erfasst mittels Saugkerzen‐ und Nmin‐Beprobung II Variabilität der NO3‐ und NH4‐Werte und Aussagegenauigkeit der Messmethoden

Nitrate leaching from intensively and extensively grazed grassland measured with suction cup samplers and sampling of soil mineral‐N II Variability of NO₃ and NH₄ values and degree of accuracy of the measurement methods Data from a grazing experiment — comparison of mean values, see Anger et al. (2002) — were used to estimate within‐field variability to asses the accuracy of two frequently used methods of estimating NO₃ leaching on pastures: (1) the ceramic suction cup sampling (with 34 cups ha^—1 minimum, calculated climatic water balance, 4 leaching periods) and (2) using the soil mineral‐N method (vertical soil NO₃ and NH₄ content in 0—0.9 m (N_min) measured at the beginning and end of two winters on a minimum of 10 different areas of 50 m² each with a minimum of 7 different sample cores). These methods were used on two permanent pastures with high mean stocking density of cattle of 4.9 LU ha^—1 on 1.3 ha with N‐fertilization of 250 kg N ha^—1 (= intensive [I]) and 2.9 LU without N fertilization on a 2.2 ha pasture (= extensive [E]). The results show that NO₃ leaching on pastures was largely due to few selectively extremely high NO₃ amounts under a few excrement spots — mainly urine spots — which would not be sampled representatively with an acceptable effort in a conventional grazing experiment. In both grazing treatments, very large spatial variation occurred. This was greater between the different suction cups than between the compound mineral N samples of each area. Therefore, a marked skewness and kurtosis demonstrated a non‐normal distribution of samples from suction cups, while mineral N values did not show this effect consistently. Sampling selected mostly spots without noticeable influence of excrement, but a few samples with very high values identified evidently urine spots from summer or autumn grazing. The differences in mean coefficient of variation (CV) between the grazing treatments and estimation methods were mainly based on the stocking rate and the density of excrement spots. CV values were 131 % [I] / 242 % [E] for NO₃ leaching measured with suction cup samplers and of 71 % [I] / 116 % [E] for soil NO₃ values and 24 % [I] / 34 % [E] for soil NH₄ values in 0—0.9 m according N_min‐method. Results of the N_min method are obviously inaccurate even with a sampling intensity much greater than 70 cores ha^—1; and so making an estimation of NO₃ leaching by this method is unsatisfactory for pastures. Compared to this, the results of suction cup sampling are more convincing; but even with a tolerated deviation of ± 20 % from the empirically estimated average and with a 95 %‐confidence interval, the calculated mean minimum number of samples in our experiment should be increased to 146 and 265 suction cups ha^—1 for the intensively and extensively grazed treatments, respectively. This requirement would be prohibitive for many field experiments.     

Catclaw (Mimosa buincifiera): a pest or a means to restore soil fertility in heavily eroded soil from the central highlands of Mexico?

 In the central highlands of Mexico, heavily eroded soils are often colonized by catclaw (Mimosa buincifiera): an N₂-fixing shrub. An experiment was carried out to investigate how this shrub affected characteristics of the soil and its biological functioning. Soil was sampled from outside and under the canopy of catclaw at three sites characterized by different degrees of erosion and an increase in plant density. The soil microbial biomass C, total amounts of bacteria, fungi, actinomycetes and free-living N₂-fixing micro-organisms were measured, while production of CO₂ and dynamics of nitrate (NO₃ ^–), nitrite (NO₂ ^–) and ammonium (NH₄ ⁺) were monitored in an aerobic incubation at 22±1  °C for 35 days. The C content was 1.6 times greater in the area with the largest density of plants and the least erosion (RECUP) compared with the site with the lowest density and greatest erosion (DEGR), while it was 1.2 times greater under the canopy of the catclaw than outside it (average of the three sites). The incorporation of N into the soil organic matter was greater under the canopy of the catclaw than outside it as the C:N ratio was on average 8.4 and 9. 1, respectively. The microbial biomass C, as a percentage of soil organic matter, was 1.5 times greater in the RECUP than in the DEGR site. Greatest total number of colony-forming bacteria and fungi (mean of organisms found under and outside the canopy) were found in the RECUP treatment and lowest in the DEGR treatment. Free-living N₂-fixing organisms and actinomycetes showed opposite trends. Greater total numbers of colony-forming bacteria, fungi, actinomycetes and free-living N₂-fixing organisms (mean of the three treatments) were found under the canopy of catclaw than outside of it, Production of CO₂ was 1.8 times greater in the RECUP than in the DEGR and 1.6 times greater under the canopy of catclaw than outside. Production of NO₃ ^– was 1.3 times greater in the RECUP than in the DEGR and 3.5 times greater under the canopy of catclaw than outside. There was no significant effect of location or canopy on NO₂ ^– and NH₄ ⁺ concentrations. It is concluded that the natural vegetation of catclaw increased microbial biomass and soil organic matter content under, but also outside its canopy, and preserved N better, releasing greater amounts of inorganic N upon mineralization. Catclaw can serve as a first colonizer of heavily eroded soil and be replaced by other vegetation, natural or crops, when fertility is restored. Received: 4 November 1999     

Interactions between Inorganic Nitrogen Nutrition and Root Development

Root development responds not only to the quantity of inorganic nitrogen in the rhizosphere, but to its form, NH₄⁺ or NO₃^?. Root growth of tomato showed a hyperbolic response to soil levels of inorganic nitrogen: very few roots were found in soil blocks depleted in inorganic nitrogen, roots proliferated as soils increased to 2 μg NH₄⁺-N g^?1 soil or 6 μg NO₃^?-N g^?1 soil, and root growth declined in soils with the higher levels of inorganic nitrogen. High NH₄⁺ concentrations inhibited root growth, but low concentrations promoted the development of an extensive, fine root system. Supply with NO₃^? as the sole nitrogen source led to a more compact root system. These differences in root morphology under NH₄⁺ and NO₃^? nutrition may be mediated through pH. Rice and maize roots absorbed NH₄⁺ most rapidly right at the apex and appeared to assimilate this NH₄⁺ in the zone of elongation. During NH₄⁺ assimilation, root cells must release protons, and the resulting acidification around the walls of cells in this region should stimulate root extension. By contrast, NO₃^? absorption reached a maximum in the maturation zone of rice and maize roots, and this NO₃^? was probably assimilated in more basal regions. Absorption of NO₃^? requires proton efflux, whereas NO₃^? assimilation requires proton influx. The net result under NO₃^? nutrition was only subtle shifts in rhizosphere pH that probably would not influence root elongation. The signal through which roots detect changes in rhizosphere NH₄⁺ and NO₃^? levels is still obscure. It is proposed that a product of nitrogen metabolism such as nitric oxide serves as a signal.     

The amount,but not the proportion,of N2 fixation and transfers to neighboring plants varies across grassland soils

ABSTRACT

Biological nitrogen fixation (BNF) is an important nitrogen source for both N₂-fixers and their neighboring plants in natural and managed ecosystems. Biological N fixation can vary considerably depending on soil conditions, yet there is a lack of knowledge on the impact of varying soils on the contribution of N from N₂-fixers in mixed swards. In this study, the amount and proportion of BNF from red clover were assessed using three grassland soils. Three soil samples, Hallsworth (HH), Crediton (CN), and Halstow (HW) series, were collected from three grassland sites in Devon, UK. A pot experiment with ¹⁵N natural abundance was conducted to estimate BNF from red clover, and the proportion of N transferred from red clover to the non-N₂ fixing grass in a grass-clover system. The results showed that BNF in red clover sourced from atmosphere in the HH soil was 2.92 mg N plant^?1, which was significantly lower than that of the CN (6.18 mg N plant^?1) and HW (8.01 mg N plant^?1) soils. Nitrogen in grass sourced from BNF via belowground was 0.46 mg N plant^?1 in the HH soil, which was significantly greater than that in CN and HW soils. However, proportionally there were no significant differences in the percentage N content of both red clover and grass sourced from BNF via belowground among soils, at 65%, 67%, 65% and 35%, 27%, 31% in HH, CN, and HW, respectively. Our observations indicate that the amount of BNF by red clover varies among grassland soils, as does the amount of N sourced from BNF that is transferred to neighboring plants, which is linked to biomass production. Proportionally there was no difference among soils in N sourced from BNF in both the red clover plants and transferred to neighboring plants.     

Nitrogen accumulation in nodulated and non‐nodulated pea plants,grown in a sandy soil at different acidities

Abstract

The growth of nitrate‐supplied and dinitrogen‐fixing pea plants was studied in a pot experiment with a sandy soil in a pH‐H_?O range from 3.4 to 5.6. Optimum growth in both treatments occurred at pH 5.0. At low pH, N₂‐plants yielded significantly less than NO₃‐plants. Planting of nodulated seedlings did not enhance yield in comparison with sowing in inoculated soil, indicating that nodulation was not the most sensitive process in restricting yield. Comparison of the nitrogen contents of shoots of planted and sown N₂‐plants allowed the suggestion that the synthesis of nitrogenous compounds was also not limiting yield. At low pH, root growth was severely reduced in dinitrogen‐fixing plants in comparison with nitrate‐supplied plants. This difference could be explained by the influence of the form of nitrogen nutrition on the cation‐anion uptake pattern of the plant and the resulting pH‐shift in the rhizosphere. It is to be expected that in an acid soil under field conditions the indirect effect of nitrate on root growth and nodulation via increase of the pH is more extensive than its direct negative effect on nodulation.     

Root-induced changes in the rhizosphere: Importance for the mineral nutrition of plants

Root-induced changes in the rhizosphere may affect mineral nutrition of plants in various ways. Examples for this are changes in rhizosphere pH in response to the source of nitrogen (NH₄-N versus NO₃-N), and iron and phosphorus deficiency. These pH changes can readily be demonstrated by infiltration of the soil with agar containing a pH indicator. The rhizosphere pH may be as much as 2 units higher or lower than the pH of the bulk soil. Also along the roots distinct differences in rhizosphere pH exist. In response to iron deficiency most plant species in their apical root zones increase the rate of H⁺ net excretion (acidification), the reducing capacity, the rate of Fe^III reduction and iron uptake. Also manganese reduction and uptake is increased several-fold, leading to high manganese concentrations in iron deficient plants. Low-molecular-weight root exudates may enhance mobilization of mineral nutrients in the rhizosphere. In response to iron deficiency, roots of grass species release non-proteinogenic amino acids (?phytosiderophores”?) which dissolve inorganic iron compounds by chelation of Fe^III and also mediate the plasma membrane transport of this chelated iron into the roots. A particular mechanism of mobilization of phosphorus in the rhizosphere exists in white lupin (Lupinus albus L.). In this species, phosphorus deficiency induces the formation of so-called proteoid roots. In these root zones sparingly soluble iron and aluminium phosphates are mobilized by the exudation of chelating substances (probably citrate), net excretion of H⁺ and increase in the reducing capacity. In mixed culture with white lupin, phosphorus uptake per unit root length of wheat (Triticum aestivum L.) plants from a soil low in available P is increased, indicating that wheat can take up phosphorus mobilized in the proteoid root zones of lupin. At the rhizoplane and in the root (root homogenates) of several plant species grown in different soils, of the total number of bacteria less than 1 % are N₂-fixing (diazotrophe) bacteria, mainly Enterobacter and Klebsiella. The proportion of the diazotroph bacteria is higher in the rhizosphere soil. This discrimination of diazotroph bacteria in the rhizosphere is increased with foliar application of combined nitrogen. Inoculation with the diazotroph bacteria Azospirillum increases root length and enhances formation of lateral roots and root hairs similarly as does application of auxin (IAA). Thus rhizosphere bacteria such as Azospirillum may affect mineral nutrition and plant growth indirectly rather than by supply of nitrogen.     

Nitrat‐Austräge auf intensiv und extensiv beweidetem Grünland,erfasst mittels Saugkerzen‐ und Nmin‐Beprobung I Einfluss der Beweidungsintensität

Nitrate leaching from intensively and extensively grazed grassland measured with suction cup samplers and sampling of soil mineral‐N I Influence of pasture management Leaching of nitrate (NO₃^—) from two differently managed cattle pastures was determined over four winters between 1993 and 1997 using ceramic suction cup samplers (with min. 34 cups ha^—1); additionally, vertical soil mineral‐N content in 0—0.9 m (N_min) was measured at the beginning and end of two winters (with min. 70 different sample cores ha^—1). The experimental site in the highlands north‐east of Cologne, Germany, is characterized by high annual precipitation (av. 1,362 mm between 1993 and 1996). An intensive continuous grazing management (1.3 ha, fertilized with 250 kg N ha^—1 yr^—1, average stocking density 4.9 LU ha^—1, = [I]) was tested against an extensive continuous grazing system (2.2 ha, av. 2.9 LU ha^—1; no N‐fertilizer but an estimated proportion of Trifolium repens up to 15 % of total dry matter in the final year, = [E]). The results can be summarized as follows: (1) Mean leaching losses of NO₃‐N, estimated from suction cup sampling and balance of drainage volume, were 85 kg NO₃‐N ha^—1 [I] and 15 kg NO₃‐N ha^—1 [E] during three wet winters with drainage volumes between 399 and 890 mm; in a dry winter with 105 mm calculated percolation, nitrate leaching decreased by a factor of 5 for both grazing treatments. (2) Although the amount of mineral N in soil (N_min) sampled in late autumn showed differences between intensive and extensive grazing, the N_min method permits no certain indication of the risk of NO₃ leaching. For example, during the winter period 1994/95 a reduction of mineral N in the soil (0—0.9 m) in both grazing treatments was found (—33 [I] / —8 [E] kg NO₃‐N ha^—1 and —26 [I] / —21 [E] kg NH₄‐N ha^—1) whereas during the winter 1996/97 an increase in almost all mean mineral N values occurred (+10 [I] / +2 [E] kg NO₃‐N ha^—1 and +10 [I] / —10 [E] kg NH₄‐N ha^—1). (3) In spite of the differences between both methods, the experiment shows that NO₃‐N leaching under extensive grazing could be reduced almost to levels close to those under mown grassland.