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.     

Influence of nitrogen source on the viability,functionality and persistence of Glomus etunicatum fungal propagules in an Andisol

This study investigated how two different N sources used as fertilizer (NO₃⁻ or NH₄⁺) interact with an inoculated arbuscular mycorrhizal (AM) fungus (Glomus etunicatum) in an Andisol from southern Chile. The effect of NO₃⁻ or NH₄⁺ on mycorrhizal and non-mycorrhizal wheat plants was measured on key root–soil interface activities: pH, acid phosphatase (P-ase) activity and P availability. Root AM colonization, extraradical mycelium length and spore number were also examined at three stages of AM symbiosis development (120, 150 and 240 days after sowing, DAS). The effect of N-source on AM propagule formation was used as an index of the quality and vigor of AM colonization. Mycorrhizal root length was greater with NO₃⁻ than with NH₄⁺ at all times. The NO₃⁻ source also improved extraradical mycelium density, which reached its maximum at 150 DAS. At each harvest the spore number in the rhizosphere soil was also greater with NO₃⁻ fertilization. This NO₃⁻ effect on spore formation ranged from 20% at a 120 DAS to 287% at a 240 DAS increase, compared with NH₄⁺. Extraradical mycelium and AM efficiency for P acquisition appeared to be related. The particular fungus/plant metabolism as affected by N sources (NO₃⁻ or NH₄⁺) applied did not result in differential plant growth or in changes in N plant acquisition, but affected AM development and activity. Differences in soil pH, available P or P-ase activity in soil seems not to be responsible for the improved physiological status of mycorrhizal development in NO₃⁻ fed plants. Mycorrhizal propagule formation in this soil and the high persistence of extraradical mycelium are important factors which may have a strong influence on the next crop, and thus, this aspects should be considered when a cropping system is designed. The influence of N sources on AM performance is of ecological and practical interest in volcanic soils when conventional management is used.     

Arbuscular mycorrhizal fungal hyphae mediating acidification can promote phytate mineralization in the hyphosphere of maize (Zea mays L.)

Mycorrhizal and non-mycorrhizal maize (Zea mays L.) plants were grown in two-compartment rhizoboxes, in order to study the effect of mycorrhizal hyphae-mediated acidification on organic P mineralization in the hyphosphere. The soil in the two compartments was supplemented with either KNO₃ or (NH₄)₂SO₄, and phytin (0 or 75 mg P kg⁻¹) was added to the hyphal compartment. P content in the shoots was significantly higher for the NH₄⁺ treatment than for the NO₃⁻ treatment, but only in the combined presence of phytin and AM fungal mycelium (Rhizophagus intraradices). NH₄⁺ treatment under these conditions also led to a decrease in hyphosphere pH, enhanced phosphatase activity in the hyphosphere and accelerated mineralization of phytin compared to the NO₃⁻ treatment. The results show that hyphosphere acidification induced by absorption of NH₄⁺ by the AM fungal mycelium leads to an increase in phosphatase activity, and consequently enhances mineralization of phytin and improves maize uptake of P from phytin-P.     

INFLUENCE OF ARBUSCULAR MYCORRHIZAL FUNGI AND AMMONIUM:NITRATE RATIOS ON GROWTH AND PUNGENCY OF SPRING ONION PLANTS

A pot experiment was carried out to study the growth and pungency of Allium fisutulosum grown in Perlite as affected by colonization by the arbuscular mycorrhizal (AM) fungi Glomus etunicatum, Glomus vesiforme, and by ammonium (NH⁺ ₄ ):nitrate (NO^? ₃ ) ratios of 5:95, 50:50, and 95:5 in 4 mM solutions. Plants were grown in a greenhouse for 20 weeks and then harvested. In general, NH⁺ ₄ :NO^? ₃ ratio of 50:50 supplied resulted in the highest shoot dry weight regardless of non-mycorrhizal and mycorrhizal plants while the effect of inoculation treatment on plant biomass was not significant. The plant sulfur (S) concentrations were usually higher in mycorrhizal plants than controls irrespective of nitrogen ratio and therefore inoculation with G. etunicatum increased the enzyme produced pyruvic acid (EPY) while inoculation with G. versiforme decreased the EPY compared with the non-mycorrhizal plants. In general, shoot pungency was lowest when NH⁺ ₄ :NO^? ₃ ratio of 95:5 supplied irrespective of mycorrhizal treatment. Colonization by both AM fungi made a substantial contribution to spring onion sulfur nutrient status but show different way on flavor characteristics of host plants.     

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.     

Soil solution chemistry in the rhizosphere of beech (Fagus silvatica L.) roots as influenced by ammonium supply

Roots can induce significant changes in the rhizosphere soil. The aim of the present study was to investigate the influence of beech (Fagus silvatica L.) roots on the chemistry of the rhizosphere soil solution. Special emphasis was given to the effect of the NH₄⁺ supply since many forest soils presently receive high NH₄⁺ inputs from atmospheric deposition. In a mature beech stand, a non‐mycorrhized long root was forced to grow into a rhizotrone filled with homogenized acidic forest soil from the Bw horizon of a Dystric Cambisol. Beside the control, a NH₄⁺ enriched treatment was installed. Thirty micro suction cups of 1 mm diameter and 0.5 cm length were placed in a systematic grid of 5 × 10 mm in each rhizotrone to enable root growth through the grid. The water potential of the soil was kept constant by supplying a synthetic soil solution. Small amounts of soil solution were sampled periodically from May to October 1999 and analyzed by capillary electrophoresis for major cations and anions. Furthermore, pH and conductivity were measured by micro electrodes. In the laboratory experiments, beech seedlings were grown in rhizotrones in a control and in a NH₄⁺ fertilized soil. The equipment for sampling soil solutions and the soil conditions in the laboratory was similar to the field experiment. In each rhizotrone a single long root grew through the lysimeter grid. The laboratory conditions induced higher rates of nitrification as compared to the field. Thus, the overall concentration range of the soil solution was not comparable between field and laboratory studies. In all treatments average soil solution concentrations of H⁺ and Al³⁺ were significantly higher in the rhizosphere than in the bulk soil. The NH₄⁺ treatment resulted, in the field and laboratory, in a strong increase of the H⁺ and Al³⁺ concentrations in the rhizosphere, accompanied by an accumulation of Ca²⁺, Mg²⁺, and NO₃^—. The observed rhizosphere gradients in soil solution chemistry were highly dynamic in time. The results demonstrate that the activity of growing beech roots results in an acidification of the soil solution in the rhizosphere. The acidification was enhanced after the addition of NH₄⁺.     

Influence of phosphorus nutrition on sulfur uptake by vesicular-arbuscular mycorrhizae of onion

Mycorrhizal and non-mycorrhizal onions were grown in pots containing soil at two P concentrations. Following ³⁵S injection into the soil, both mycorrhiza) and non-mycorrhizal plants from high P treatments had significantly higher ³⁵S concentrations in roots compared to non-mycorrhizal, low P controls. Mycorrhizal, low P plants had higher concentrations of ³⁵S in shoots than did non-mycorrhizal, low P plants. In a second experiment detached non-mycorrhizal onion roots from plants given a nutrient solution containing P for 26 days before short-term uptake experiments absorbed at greater rates from solution than roots from plants given a complete minus-P nutrient solution. This occurred at all three concentrations of S tested. 1 mM. 10μM, and 0.1 μM. Increased S uptake by mycorrhizal plants can result from increased S absorbing power of roots with enhanced P status.     

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.     

Hyphal translocation and uptake of sulfur by vesicular-arbuscular mycorrhizae of onion

Translocation of S by external hyphae of Glomus fascieulatus, a vesicular-arbuscular (VA) mycorrhizal fungus, was demonstrated. When tracers were injected 8 cm from onion roots in soil chambers, both ³⁵S and ³²P appeared in roots of mycorrhizal plants. Neither radionuclide was present in non-mycorrhizal plants.In a second soil-chamber experiment, mycorrhizal onions took up more ³⁵S per unit dry weight than non-mycorrhizal controls when ³⁵S was injected into soil chambers in a region 3–6 cm from roots. Severing of external hyphae between the application area and the roots reduced the concentration of ³⁵S in tops of mycorrhizal plants but not in roots. Volatile ³⁵S per unit dry weight collected from all plants in each treatment was highest in the mycorrhizal-hyphae intact treatment, and higher in the mycorrhizal-hyphae severed treatment than the non-mycorrhizal treatment. Movement of ³⁵S in soil from the area of application to roots was similar for all treatments.Increased uptake of S from soil by VA mycorrhizal plants can result from hyphal translocation of S to infected roots by external mycorrhizal hyphae.     

根际pH对玉米利用磷酸单酯和双酯盐的影响

【目的】土壤有机磷在土壤全磷中占有很大比重,是植物潜在的有效磷源,但必须通过磷酸酶的水解作用释放出无机磷才能被植物利用。土壤中有机磷的主要形式为磷酸单酯和磷酸双酯。本研究中,我们探讨了无菌条件下不同形态的氮源引起的根际pH变化如何影响植物对这两种有机磷的活化利用。【方法】采用琼脂无菌培养体系种植玉米,向玉米植株供应两种形态的氮源和磷源, 氮源为硝态氮和铵态氮,磷源为植酸钙和卵磷脂,植酸钙属于磷酸单酯盐,卵磷脂属于磷酸双酯盐。不同的供氮形态会导致根际pH变化,进而研究根际pH变化对磷酸单酯盐和磷酸双酯盐的活化利用所产生的影响。【结果】当给玉米供应铵态氮时,根际pH从5.5降至4.0; 供应硝态氮时,根际pH升至6.6。测定玉米根际的琼脂中根系分泌的磷酸单酯酶和磷酸双酯酶活性发现,磷酸单酯酶活性在pH 6.0~7.0之间最高,磷酸双酯酶活性在pH 7.0~8.0之间达到最高。无论以植酸钙还是卵磷脂为有机磷源,相对于铵态氮处理,硝态氮处理能够使根际保持较高的磷酸单酯酶或磷酸双酯酶活性。有机磷的水解过程由磷酸酶活性和底物有效性两个因素控制,而植酸钙的水解受根际pH影响很大,在一定pH范围内,植酸钙的溶解度随根际pH值降低而升高,有效态磷浓度的增加,使得磷酸酶的底物有效性提高。在供应铵态氮时,根际pH值降低,玉米对植酸钙的利用效率高于硝态氮处理,尽管供硝态氮时磷酸单酯酶活性更高。同时,在供应铵态氮条件下,植株对植酸钙的利用率要显著高于卵磷脂,原因在于虽然磷酸双酯酶和磷酸单酯酶活性较低,由于植酸钙的溶解度较大,它的底物有效性更高。因此,植酸钙处理中植株的磷含量更高。相反,在供应硝态氮时,植酸钙溶解度减小而两种磷酸酶活性较高,卵磷脂处理中植株的磷含量更高。【结论】土壤中有机磷的水解过程由磷酸酶活性和有机磷底物有效性两个因素控制,酶活性与根际pH密切相关。本研究说明土壤有机磷的活化必须首先转化为溶解于水溶液中的状态,才能作为磷酸酶的底物被催化水解。我国长期施用化肥导致北方土壤大范围酸化,这种酸化无疑对土壤固有或随有机物料进入农田的有机磷的活化利用是具有重要贡献的,应该在北方土壤养分管理中应加以考虑。     

Phosphorus uptake of maize as affected by ammonium and nitrate nitrogen - Measurements and model calculations -

Phosphorus uptake is often enhanced by ammonium compared to nitrate nitrogen nutrition of plants. A decrease of pH at the soil-root interface is generally assumed as the cause. However, an alteration of root growth and the mobilization of P by processes other than net release of protons induced by the source of nitrogen may also be considered. To study these alternatives a pot experiment was conducted with maize using a fossil Oxisol high in Fe/Al-P with low soil solution P concentration. Three levels of phosphate (0, 50, 200 mg P kg^?1) in combination with either ammonium or nitrate nitrogen (100 mg N kg^?1) were applied. Plants were harvested 7 and 21 d after sowing, P uptake measured and root and shoot growth determined. To assess the importance of factors involved in the P transfer from soil into plants, calculations were made using a model of Barber and Claassen. In the treatments with no and low P supply NH₄-N compared to NO₃-N nutrition increased the growth of the plants by 25 % and their shoot P content by 38 % while their root growth increased by 6 % only. The rhizosphere pH decreased in the NH₄-N treatments by 0.1 to 0.6 units as compared to the bulk soil while in the NO₃-N treatments it increased by 0.1 to 0.5 units. These pH changes had a minor influence on P uptake only, as was demonstrated by artificially altering the soil pH to 4.7 and 6.3 respectively. At the same rhizosphere pH, however, P influx was doubled by the application of NH₄-compared to NO₃-N. It is concluded that in this soil the enhancement of P uptake of maize plants after ammonium application cannot be attributed to the acidification of the rhizosphere but to effects mobilizing soil phosphate or increasing P uptake efficiency of roots. Model calculation showed that these effects accounted for 53 % of the P influx per unit root length in the NO₃-N and 72 % in the NH₄-N supplied plants if no P was applied. With high P application the respective figures were only 18 and 19%.     

Hydroxyl release by maize (Zea mays L.) roots under acidic conditions due to nitrate absorption and its potential to ameliorate an acidic Ultisol

Purpose

Hydroxyl ion release by maize (Zea mays L.) roots under acidic conditions was investigated with a view to develop a bioremediation method for ameliorating acid soils in tropical and subtropical regions.

Materials and methods

Two hydroponic culture experiments and one pot experiment were conducted: pH, nitrogen state, and rhizobox condition, which investigated the effects of different nitrogen forms on hydroxyl release by maize roots under acidic conditions.

Results and discussion

The pH of the culture solution increased as culture time rose. The gradient of change increased with rising NO₃ ^?/NH₄ ⁺ molar ratios. Maize roots released more hydroxyl ions at pH 4.0 than at pH 5.0. The amount of hydroxyl ions released by maize roots at a constant pH was greater than those at a nonconstant pH. Application of calcium nitrate reduced exchangeable acidity and increased the pH in an Ultisol rhizosphere, compared with bulk soil. The increasing magnitude of soil pH was greater at higher doses of N. The absorption of NO₃ ^?–N increased as the NO₃ ^?/NH₄ ⁺ molar ratios rose, which was responsible for hydroxyl ion release and pH increases in culture solutions and rhizosphere.

Conclusions

Root-induced alkalization in the rhizosphere resulting from nitrate absorption by maize plants can be used to ameliorate acidic Ultisols.     

Effects of Arbuscular Mycorrhizal Fungi and Ammonium: Nitrate Ratios on Growth and Pungency of Onion Seedlings

ABSTRACT

A pot experiment was conducted to study the growth and pungency of Allium cepa L. grown in Perlite as affected by colonization by the arbuscular mycorrhizal (AM) fungi Glomus versiforme and Glomus intraradices BEG141 and by ammonium:nitrate (NH₄ ⁺:NO₃ ^?) ratios of 3:1, 1:1, and 1:3 in 4 mM solutions. Plants were harvested when bulb formation commenced. In general, mycorrhizal colonization resulted in increased shoot dry weight, shoot length, sheath diameter, root nitrogen (N) and phosphorus (P) content (except with G. intraradices and a NH₄ ⁺:NO₃ ^? ratio of 1:3), shoot N and P concentrations (except with G. versiforme and a NH₄ ⁺:NO₃= ratio of 3:1) and content. Plants inoculated with G. versiforme had higher growth parameters and N and P content than those with G. intraradices, whereas N and P concentrations showed the opposite trends. Growth parameters and N and P content of non-mycorrhizal plants were highest at a NH₄ ⁺:NO₃= ratio of 1:3, while those of plants inoculated with G. versiforme or G. intraradices were highest at a ratio of NH₄ ⁺:NO₃ ^? 3:1 or 1:1. Neither mycorrhizal colonization nor proportion of inorganic N species significantly affected bulb enzyme-produced pyruvate or total or organic sulfur (S) concentrations in plant shoots. Colonization by AM fungi made a substantial contribution to onion growth and may not have been directly related to bulb pungency at early stages of plant growth. However, the influence of AM fungi on plant N and P metabolism may have implications for onion flavor at later stages of plant growth.     

Ammonium and nitrate in the rhizosphere of spring barley, hordeum vulgare L., and take-all disease

The incidence and severity of take-all disease, due to Gaeumannomyces graminis (Sacc.) Arx & Olivier var. tritici Walker, was observed on spring barley plants growing in soil in two glasshouse experiments. Soil amendments of NH⁺₄-N significantly increased the number of diseased plants and roots during the first month after germination in comparison with controls unamended with N (P < 0.05). No significant difference in the incidence of take-all disease was detected between more mature barley plants growing in soil amended with either NH⁺₄ or NO^?₃-N and unamended controls. The least take-all disease in 3 month-old barley plants was observed when N was supplied as foliar sprays of urea at 0.5 mg N kg^?1 soil (P < 0.01). There was no significant correlation between the degree of infection and the NH⁺₄-N to NO^?₃-N ratio in the rhizosphere soil     

Mycorrhizal inoculation increases genes associated with nitrification and improved nutrient retention in soil

The effects of arbuscular mycorrhizal (AM) inoculation on prokaryote abundance within the maize rhizosphere and hyphosphere, and retention of nutrients were investigated. Maize plants were grown in pots with a membrane located at a soil depth of approximately 16 cm that allowed growth of fungal hyphae above and below the membrane, but did not allow growth of roots below the membrane. As expected, mycorrhizal inoculation significantly increased the contents of soil organic matter, Total Kjeldahl Nitrogen (primarily organic N), and Mehlich 1 phosphorus relative to the non-inoculated control. Copy numbers of 16S rRNA genes were significantly higher in the mycorrhizal compartments relative to non-mycorrhizal controls. Bacterial ammonia monooxygenase (AOB) genes, but not archaeal monooxygease genes (AOA), were significantly higher in planted treatments with and without addition of mycorrhizae, indicating that mycorrhizae stimulate prokaryotic growth and bacterial nitrification. The ecological relevance of increased NOx-N resulting from the growth of AOB in inoculated soils is not clear; however, increased mobility of NOx-N over NH₄ ⁺ could result in a competition between leaching loss and increased uptake by mycorrhizae.     

Soil Acidity Changes in Bulk Soil and Maize Rhizosphere in Response to Nitrogen Fertilization

The capacity of nitrogen (N) fertilizers to acidify the soil is regulated principally by the rate and N source. Nitrogen fertilizers undergo hydrolysis and nitrification in soil, resulting in the release of free hydrogen (H⁺) ions. Simultaneously, ammonium (NH₄ ⁺) absorption by roots strongly acidifies the rhizosphere, whereas absorption of nitrate (NO₃ ^?) slightly alkalinizes it. The rhizosphere effects on soil acidity and plant growth in conjunction with N rate are not clearly known. To assess the impact of these multiple factors, changes in the acidity of a Typic Argiudol soil, fertilized with two N sources (urea and UAN) at two rates (equivalent to 100 and 200 kg N ha^?1), were studied in a greenhouse experiment using maize as the experimental plant. Soil pH (measured in a soil–water slurry), total acidity, exchangeable acidity, and exchangeable aluminum (Al) were measured in rhizospheric and bulk soil. Plant biomass and foliar area (FA) were also measured at the V₆ stage. Nitrogen fertilization significantly reduce the pH in the bulk soil by 0.3 and 0.5 units for low and high rates respectively. Changes in the rhizosphere (the “rhizospheric effect”) resulted in a significant increase in soil pH, from 5.9 to 6.2. The rhizospheric effect × N source interaction significantly increased exchangeable acidity in the rhizosphere relative to bulk soil, particularly when UAN was added at a low rate. Only total acidity was significantly increased by the fertilizer application rate. In spite of the bulk soil acidification, no significant differences in exchangeable aluminum were detected. Aerial biomass and FA were significantly increased by the higher N rate, but N source had no effect on them. Although changes in acidity were observed, root biomass was not significantly affected.     

Effects of thiamine and nitrogen fertilizer form on the number of N2−fixing and total bacteria in the rhizosphere of maize plants

The effects of thiamine (vitamin B1) application as seed dressing and of N form supplied (NH₄⁺ versus NO₃^?) on rhizosphere pH and on rhizosphere microorganisms were evaluated in two different soils. Imbibition of maize (Zea mays L.) seeds with thiamine (1 g kg^?1) increased seed thiamine content by a factor of 370. Maize plants from untreated and treated seeds were cultivated in a growth chamber under controlled conditions for 10 d in a sandy loam soil, pH 7.1 (Mascherode soil) or in a sandy soil, pH 4.8 (Niger soil) fertilized with two different N sources (NO₃?N or NH₄?N with dicyandiamide, 100 and 250 mg N kg^?1 soil). The rhizosphere pH was not affected by thiamine, only slightly affected by N source in the Mascherode soil, but markedly affected in the Niger soil. Thiamine application and N source affected the most probable number (MPN) of diazotrophs and total bacteria isolated from the rhizosphere soil of 10 d old maize plants. In the Mascherode soil, thiamine application increased MPN of diazotrophs 4-fold and total bacteria 2-fold when the soil was fertilized with 100 mg NO₃?N compared to untreated seedlings. Compared to Mascherode soil, in the Niger soil, MPN of diazotrophs was extremely low, especially after NH₄?N treatment which significantly decreased pH of the rhizosphere. Thiamine application had only marginal effects on the MPN of diazotrophs and total bacteria. Total bacteria isolated from Niger soil fertilized with NH₄?N were about 10-fold lower compared to the soil from Mascherode. However, in the other two treatments, total bacteria were higher in the Niger soil compared to the Mascherode soil. In the Niger soil, apparently some of the heterotrophs (the Actinomycetes dominated in this soil) might have suppressed the diazotrophs. The results of the present study demonstrate that in many cases seed treatment with thiamine enhances MPN of diazotrophs and total bacteria in the rhizosphere of maize seedlings.     

Effect of ectomycorrhizae on the growth and uptake and transport of 15N-labeled compounds by Pinus tabulaeformis seedlings under water-stressed conditions

 Seedlings of Chinese pine (Pinus tabulaeformis Carr.) were grown in a growth chamber using a sterilized soil/sand/vermiculite mixture, and were inoculated with the ectomycorrhizal fungus Cenococcum graniforme (Sow.) Ferd. and Winge. The function of external mycelium of ectomycorrhiza for uptake and transport of N from ¹⁵N-labeled NH₄ ⁺ and NO₃ ^– for plant nutrition was evaluated under well-watered conditions and water stress. The pots comprised two-compartments, whereby the penetration of roots was prevented by a nylon mesh bag which the mycorrhizal hyphae were allowed to pass in order to colonize the rest of the pot. ¹⁵N-labeled NH₄ ⁺ or NO₃ ^– was applied to the area of the pot to which the root had no access. At harvest, the ¹⁵NH₄ ⁺ concentration in plant tissues was significantly promoted by the formation of mycorrhizae both under well-watered conditions and water stress. The ¹⁵NO₃ ^– concentration was reduced by water stress and increased by mycorrhizal formation. The enhancement of ¹⁵NO₃ ^– uptake caused by mycorrhizal formation was more evident under water stress than under well-watered conditions. The external mycelia of the ectomycorrhizae took up and transported NH₄ ⁺ and NO₃ ^– from the soil to the plant, thereby improving plant nutrition and growth, in addition to helping the plants to avoid the effects of water stress. Received: 31 October 1997     

Specific activity of phosphorus in mycorrhizal and non-mycorrhizal plants in relation to the availability of phosphorus to plants

Phosphate was allowed to react with a soil to which iron hydroxide had been added. The P was then labelled by a subsequent addition of ³²P. Soil P was extracted by 10 mm CaCl₂, 0.5 m NaHCO₃, and acid NH₄F solutions and the specific activity of P in the extracts was measured. Subterranean clover plants were grown both with and without a mycorrhizal fungus. Phosphorus contents and the specific activities of P in the plant shoots were determined.For mycorrhizal plants, adding iron hydroxide had no effect on the amount of P taken up, but for non-mycorrhizal plants it decreased the uptake. However there was no effect of iron hydroxide or of mycorrhizal infection on the specific activity of P in the plants. Adding iron hydroxide had no effect on the amount of P extracted by acid NH₄F, but decreased the P extracted by 10mm CaCl₂ and by 0.5 m NaHCO₃. The specific activity of P in the extracts was not affected by the addition of iron hydroxide and was the same for the three extractants. Further, the specific activity of P in all extractants was similar to that of P in both mycorrhizal and non-mycorrhizal plants. Thus differences in the availability of soil P to mycorrhizal and non-mycorrhizal plants and to the extractants were not reflected by differences in labelling. It therefore follows that lack of difference in specific activity between mycorrhizal and non-mycorrhizal plants does not eliminate the possibility that mycorrhizal plants can obtain P that was unavailable to non-mycorrhizal plants.     

Nitrate and ammonium nutrition of plants: Effects on acid/base balance and adaptation of root cell plasmalemma H+ ATPase

The increase of rhizosphere pH in the course of nitrate nutrition results from H⁺ consumption in the external medium during uptake of NO₃^? in a H⁺ co-transport and from internal OH^? production during nitrate reduction. Synthesis of organic acids for NH₄⁺ assimilation as well as strong partial depolarization of membrane potential with NH₄⁺ uptake are the important reasons for rhizosphere acidification during ammonium nutrition. Despite differences in proton balance depending on N form, cytoplasmic pH changes are small due to physico-chemical buffering, biochemical pH regulation, H⁺ inclusion in vacuoles, and H⁺ release into the rhizosphere. Because of the large capacity for proton excretion the plasmalemma H⁺ ATPase of root cells plays an essential role during ammonium nutrition. An increase of the kinetic parameter V_max after ammonium nutrition relative to nitrate nutrition suggests that the capacity of H⁺ release may be adjusted to the particular requirements of ammonium nutrition. Moreover, H⁺ ATPase is adjusted not only quantitatively but also qualitatively. The increase of the kinetic parameter k_m as well as the capability of the plasmalemma vesicles in vitro to establish a steeper pH gradient favours the supposition that H⁺ ATPase isoforms are formed which allow H⁺ release into the rhizosphere under conditions of low pH or poor H⁺ buffering of the soil. In this respect species differences exist, e.g. between maize (efficient adaptation) and faba bean (poor adaptation).