Importance of plasmalemma ATPase in the retention and exclusion of inorganic ions

The experiments were focused on the question whether the plasmalemma ATPase activity (proton pump) has an influence on the efflux of major inorganic ion species. Efflux from roots of intact Trifolium pratense, Hordeum vulgare, Glycine max, and Zea mays was examined into a solution containing 100 μM CaCl₂ and 500 μM NH₄⁺ as sulfate in the control solution and 100 μM CaCl₂ and 500 μM NH₄⁺ as vanadate in the test solution. Vanadate being an inhibitor of the plasmalemma ATPase depressed significantly the H⁺ secretion of roots into the outer solution but had no major impact on the efflux of cation species. In the presence of vanadate significantly higher amounts of sulfate, phosphate, and nitrate were released into the outer solution by roots of soya and maize as compared with the control treatment (no vanadate). In the absence of vanadate, virtually no nitrate was released by all species examined whereas in the vanadate treatment significant amounts of NO₃^? were released. Vanadate inhibited the uptake of Cl^? in barley and maize and increased the uptake of Ca²⁺ in soya. It is concluded that the plasmalemma ATPase activity plays a major rule in the “ionic stat” of cells in providing protons to the apoplast for the reabsorption of sulfate, phosphate, and particularly nitrate which have leaked out of the cytosol.     

Role of Plasmalemma H+ ATPase in Sugar Retention by Roots of Intact Maize and Field Bean Plants

Net release and net uptake of sugars by roots of intact maize (Zea mays cv. Blizzard) and field bean (Vicia faba L. cv. Alfred) were studied at micromolar external sugar concentrations that are relevant to the rhizosphere. Besides various sugars not further characterized there was net release of glucose, fructose, sucrose, arabinose, ribose, and galactose. The net release of these sugars into the root medium (0.1 mM CaSO₄) was stimulated by the protonophore CCCP (10 μM), the sulfhydryl reagent NEM (300 μM), the specific inhibitor of plasmalemma H⁺ ATPase vanadate (0.5 mM), and by the inhibitor of the glucose carrier phlorizin (2 mM). Net uptake of glucose, fructose, and arabinose from 10 μM external concentrations was inhibited by these substances. Stimulation of net release and inhibition of net uptake was most pronounced for glucose. Sucrose added to the root medium was hydrolyzed by invertase activity leading to glucose and fructose uptake by roots. It is concluded that the retention of sugars by plant roots is not only determined by plasmalemma permeability but is also controlled by the H⁺ electrochemical gradient established by ATPase activity (retrieval mechanism). The proton gradient drives a sugar/H⁺ cotransport system that is selective for glucose but may also transport other sugars, particularly in the absence of glucose.     

Indications for passive rather than active release of natural nitrification inhibitors in Brachiaria humidicola root exudates

Plants have the ability to suppress microbial nitrification process through secondary metabolites released from their root exudates or/and leaf litter. For decades, grasses were suggested to control nitrification process, and recently, Brachiaria humidicola accession 26159 (BH) as a tropical and subtropical grass has been shown to reduce nitrification rates under laboratory and soil conditions. In this study, experiments were conducted under controlled conditions in nutrient solution culture to investigate whether the reported release of natural nitrification inhibitors from root exudates of BH is an active or passive phenomenon. So different variables such as N-form (nitrate vs. ammonium), collecting medium (distilled water vs. 1 mM NH₄Cl) and collecting period (6 vs. 24 hrs) were included to study the hypothesis. Results showed when root exudates were collected in distilled water there was no nitrification inhibition activity for all ammonium and nitrate grown plants. However, when collection was done in a medium containing 1 mM NH₄Cl, root exudates showed significant nitrification inhibition activity similar to results obtained by Subbarao et al. The observed nitrification inhibition activity had a positive correlation to ammonium treatment particularly in collection medium, probably due to root cells damage induced by low pH and membrane depolarization under ammonium nutrition. This was more supported by application of shoot homogenates of NH₄⁺, NO₃^? or NH₄NO₃ grown plants that showed significant nitrification inhibition activity compared to distilled water and DMPP controls in a bioassay test, independent of N-form. Potassium concentrations in root exudates (as a result of potassium leakage) were found to increase in root washings of plants, which were grown with ammonium, particularly when root exudates were collected in 1 mM NH₄Cl solution. In addition, higher electric conductivity of root washings after collection of root exudates in ammonium containing medium (low pH) and also in nitrate containing medium which adjusted to pH 3 by applying H₂SO₄, strongly suggest that release of natural nitrification inhibitors from root exudates of B. humidicola may not be an active process, but instead it is rather a passive phenomenon by ammonium induced root physicochemical damages.     

Does H+ pumping by plasmalemma ATPase limit leaf growth of maize (Zea mays) during the first phase of salt stress?

In the first phase of salt stress, growth of plants is impaired mainly by osmotic stress. To elucidate the effect of NaCl salinity on elongation growth of maize leaves in the first phase of salt stress, we investigated the effect of NaCl on gene expression and activity of the plasmalemma H⁺ ATPase of elongating leaves of maize (Zea mays L.). Treatment of maize plants with 125 mM NaCl for 3 d decreased leaf growth relative to control plants (1 mM NaCl). Whereas H⁺ ATPase hydrolytic activity was unaffected, the ability of the H⁺ ATPase to establish a pH gradient was strongly reduced. Total mRNA of plasmalemma H⁺ ATPase was slightly increased. However, mRNA of the ATPase isoform MHA1 was significantly reduced and ATPase isoform MHA4 was strongly increased at the mRNA level. Synthesis of total H⁺ ATPase protein was unchanged as revealed by western blot. The results indicate that reduced pumping of H⁺ ATPase in leaf plasmalemma under salt stress may be caused by a switch to gene expression of the specific isoform MHA4, which shows inferior H⁺‐pumping efficiency in comparison to isoforms expressed under control conditions. We propose that reduced H⁺ pumping of plasmalemma H⁺ ATPase is involved in the reduction of leaf growth of maize during the first phase of salt stress.     

Acidification of Red Pine Forest Soil Due to Acidic Deposition in Chunchon, Korea

The effect of acidic deposition on the soil under red pine forest in Chunchon, Korea was investigated. Precipitation, stream water, and soil solution chemistry were monitored at the watershed from 1997 to 1998. Acidity of the open-bulk precipitation was often neutralized by large amounts of ammonia (NH₃) that might have originated from livestock farming and fertilization. Estimated elemental budget at the watershed showed a positive correlation between loss of base cations and proton (H⁺) production due to nitrogen transformation in soil (ΔH⁺ _NT: ([NH₄ ⁺]_in-[NH₄ ⁺]_out)- ([NO₃ ^?]_in-[NO₃ ^?]_out)). When ΔH⁺ _NT increased, concentrations of nitrate in soil solutions also increased. Consequently, pH values of soil solutions decreased, although ion exchange with base cations contributed to buffer reaction. Since acid buffering capacity of the red pine forest soil was small, it was concluded that the input of ammonium nitrogen enhanced nitrification in soil thus causing soil acidification represented by loss of base cations from the watershed.     

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%.     

Effects of different N sources on N nutrient and microbial distribution across soybean rhizosphere

Abstract

Nitrogen is the most important nutrient for plant growth. In the present study, investigations were carried out on the effects of sodium nitrate, ammonium sulfate, urea, and two types of controlled-release coated urea (LP-40 and LP-70) fertilizers on the NO₃ ^?-N, NH₄ ⁺-N concentrations, and microbial numbers as well as pH distribution across the rhizosphere of soybean (Glycine max L. Merrill, var. Heinong 35). The study was conducted on a typical black soil using a rhizobox system. The results showed that NO₃ ^?-N was the main source of nitrogen, which was deficient in the rhizosphere in the treatments of ammonium sulfate, urea, LP-40, and LP-70, but accumulated considerably in the sodium nitrate treatment. The NH₄ ⁺-N concentration slightly increased in the rhizosphere in the ammonium sulfate treatment, and decreased in the rhizosphere when the other four kinds of N fertilizers were supplied. In an the treatments, bacterial and fungal numbers were highest in the central compartment (C.C.) of the rhizoboxes where the soybean root system was confined, but the rhizosphere width estimated from the increase in the microbial abundance differed among different N fertilizers. The experimental results also indicated that the fungal composition in the C.C. was less diverse than in other parts of the rhizobox compartments, and that the majority of fungal groups was represented by Penicillium spp., suggesting that the microbial distribution across the soybean rhizosphere differed both quantitatively and qualitatively.     

Effects of nitrogen fertilizer on the acidification of two typical acid soils in South China

Purpose

A laboratory incubation under constant temperature and humidity was conducted to estimate the impacts of nitrogen (N) fertilizers on the acidification of two acid soils (Plinthudult and Paleudalfs) in south China.

Materials and methods

The experiment had three treatments, i.e., control (CK), addition of urea (U), and addition of ammonium sulfate (AS). We measured soil pH, nitrate (NO₃ ^?), ammonium (NH₄ ⁺), exchangeable hydrogen ion (H⁺), and aluminum ion (Al³⁺) concentrations at various intervals during the 90 days of incubation. Soil buffering capacity (pHBC) was also measured at the end of the experiment.

Results and discussion

The application of N fertilizers resulted in soil acidification. The U treatment caused greater acidification of the Plinthudult soil than the AS treatment, while there were no differences between U and AS treatments on the acidification of the Paleudalfs. At the end of the trial, the pHBC of Plinthudult in AS treatment was greater than that in CK and U treatments, which may be due to the buffering system of NH₄ ⁺ and NH₄OH. However, the pHBC of Paleudalfs was unchanged between treatments. The dynamics of exchangeable H⁺ and Al³⁺ corresponded to that of soil pH. Correlation analysis showed that both soil exchangeable H⁺ and soil exchangeable Al³⁺ were significantly related to soil pH.

Conclusions

Application of urea and ammonium sulfate caused acidification in both soils and increased soil exchangeable Al³⁺ and H⁺ concentrations in the Paleudalfs. The application of urea increased exchangeable Al³⁺, and ammonium sulfate increased pHBC in the Plinthudult.     

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.     

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.     

盐胁迫下氮素形态对油菜和水稻幼苗离子运输和分布的影响

【目的】土壤盐碱化是制约农作物产量的主要因素之一,盐胁迫影响养分运输和分布,造成植物营养失衡,导致作物发育迟缓,植株矮小,严重威胁着我国的粮食生产。在必需营养元素中,氮素是需求量最大的元素,NO-3和NH+4是植物吸收氮素的两种离子形态。植物对盐胁迫的响应受到不同形态氮素的调控,研究不同形态氮素营养下植物的耐盐机制对提高植物耐盐性及产量具有重要的意义。【方法】本文以喜硝植物油菜(Brassica napus L.)和喜铵植物水稻(Oryza sativa L.)为试验材料,采用室内营养液培养方法,研究了NO-3和NH+4对Na Cl胁迫下油菜及水稻苗期生长状况、对Na+运输和积累的影响,以对照与盐胁迫植株生物量之差与Na+积累量之差的比值,评估Na+对植株的伤害程度。【结果】1)在非盐胁迫条件下,硝态氮营养显著促进油菜和水稻根系的生长;盐胁迫条件下,油菜和水稻生物量均显著受到抑制,Na Cl对供应铵态氮营养植株的抑制更为显著。2)盐胁迫条件下,两种供氮形态下,油菜和水稻植株Na+含量均显著增加,硝态氮营养油菜叶柄Na+显著高于铵态氮营养,叶柄Na+含量/叶片Na+含量大于铵营养油菜,硝态氮营养水稻根系Na+含量显著低于铵营养,地上部则相反。3)铵营养油菜和水稻Na+伤害度显著高于硝营养植株。4)盐胁迫条件下,硝态氮营养油菜地上部和水稻根系K+含量均显著高于铵态氮营养。5)盐胁迫条件下,硝营养油菜和水稻木质部Na+浓度,韧皮部Na+和K+浓度及水稻木质部K+浓度均高于铵营养植株。【结论】与铵营养相比,硝营养油菜和水稻具有更好的耐盐性。硝态氮处理油菜叶柄Na+显著高于铵态氮处理,能够截留Na+向叶片运输。同时,供应硝态氮营养更有利于油菜和水稻吸收K+,有助于维持植物体内离子平衡。盐胁迫下,硝营养油菜和水稻木质部Na+浓度,韧皮部Na+和K+浓度及水稻木质部K+浓度均高于铵营养植株,表明硝态氮营养油菜和水稻木质部-韧皮部对离子有较好的调控能力,是其耐盐性高于铵营养的原因之一。     

Changes in net proton release by roots of intact maize plants after local nutrient supply in a split root system

The effect of local nutrient supply to maize roots (Zea mays L. cv. Blizzard) on net proton release was studied using the split root technique (SRNS, SRCa) to compare plants that were cultivated with their roots completely in either nutrient solution (NS) or 0.1 mM CaSO₄ (Ca). Roots in NS released more protons than roots in Ca. This higher net proton release was associated with significantly higher ATP concentrations in the root tissue. Higher net proton release and ATP concentrations were also observed after a 4 h lag phase when 20 μM abscisic acid were exogenously applied to roots in 0.1 mM CaSO₄. It is suggested that higher metabolic activity in roots supplied with nutrients increased ATP concentrations and thus the substrate supply of the plasma membrane H⁺ ATPase. When only half of the root system was supplied with nutrient solution with the other half bathed in 0.1 mM CaSO₄, the roots in the SRNS compartment released significantly higher amounts of protons relative to the NS control plants. Conversely, roots in the SRCa compartment showed net proton uptake in contrast to the roots of control plants in 0.1 mM CaSO₄ which significantly acidified the root medium. These differences in proton release by roots in the split root system and control roots could not be explained in terms of differences in ATP concentrations. It is therefore suggested that an internal signal may lead to a modification of the plasma membrane H⁺ ATPase as shown earlier during plant adaptation to low pH in the root medium.     

Method for the quantification of acid production by plants in gel

Abstract

Hydrogen (H⁺) and hydroxyl ion (OH^‐) production by the tropical grass, Brachiaria humidicola, is quantified using a method in which the plants are grown in soil then transferred to agar gel for 24 h. The amount of H⁺ and OH^‐produced was calculated from the pH of the melted gel and the gels’ buffer curve. Values were obtained for plants of different ages and with nitrogen (N) supplied in the gel as nitrate (NC₃ ^‐), ammonium (NH₄ ⁺), or ammonium nitrate (NH₄NO₃) and compared with data calculated using the sum of H⁺ changes in differently colored zones of the gel. Daily H⁺ and OH^‐ production increased with plant age and total dry matter for the NH₄ ⁺‐ and NO₃ ^‐‐fed plants, respectively. By integrating the data over time, a value of 0.33 mmol H⁺ plant^‐1 was obtained for the total H⁺ production over 62 d. The proposed method was sufficiently rapid and versatile to allow the comparison between plant species or genotypes, which were grown using a variety of nutrient supplies. This procedure may indicate how acid production affects plant nutrient acquisition and aid the prediction of soil acidification by different plant species or cultivars.     

Benefits of mycorrhizae to soybeans grown on various regimes of nitrogen nutrition 1

Nodulating and nonnodulating isolines of soybean (Glycine max Merr ‘Clay') were grown in sand culture in a greenhouse. The plants were cultured with or without mycorrhizal (Glomus mosseae) infection, and nodulating plants were inoculated with Rhizobium iaponicum. Phosphorous was supplied as hydroxyapatite or dicalcium phosphate with N nutrition from nitrate or as combinations of nitrate and ammonium or nitrate and urea. Best growth of the nodulating isoline was with urea nutrition. Best growth with the nonnodulating isoline was with ammonium nutrition. Urea‐treated nodulating plants showed increased growth due to mycorrhizae. Urea‐treated or ammonium‐treated nonnodulating plants showed growth increases due to mycorrhizae. Nitrate‐treated plants did not show increased growth due to mycorrhizae. Mycorrhizal infection was greatest with urea nutrition, and the infection increased the tissue N content of these plants relative to nonmycorrhizal plants. Enhancement of tissue P accumulation through mycorrhizae was greater with hydroxyapatite than with dicalcium phosphate. The efficiency of the symbiotic relationship of Glvcine‐Glomus‐Rhizobium depended on a supply of reduced nitrogen, a high N:P ratio in roots, and a neutral pH in the rhizosphere. Urea nutrition met these requirements best.

Vesicular‐arbuscular mycorrhizal (VM) associations may improve the capacity of higher plants to acquire nutrients. These benefits have been studied extensively in relation to P nutrition since plant requirements for P are high relative to its availability in soils². Significant benefits may occur also in the nutrition of plants with micronutrients³. Reviews of literature suggest that the function of mycorrhizae in the acquisition of N by plants is variable⁴‘⁵‘⁶. The sources of N under various cultural or ecological conditions may account for the conflicting findings among researchers.

Hyphal transport of N is of little importance with NO3^‐nutrition because of the high mobility of this ion⁶ but may be important for the relatively immobile NH₄ ⁺ ion. Roots of NH₄ ^‐nourished plants often are restricted, and mycorrhizae may extend their absorbing surfaces⁷. Mycorrhizae may prefer NH₄‐N over NO₃‐N for their growth and development⁴. In some cases, NH₄ ⁺ ions restrict mycorrhizal infection compared to the effect of NO₃ ^‐ions⁷. The drop in ambient pH associated with NH₄ ⁺ nutrition may be a cause of this inhibition⁷‘⁸‘⁹. Plants grown on urea may not encounter the problems of acidity in the root zone and yet may have access to NH₄ ⁺ nutrition.

The N contents of mycorrhizally infected plants relative to those of uninfected plants are variable¹⁰‘^11,12,13. These diverse results could arise from differences in the levels and forms of N applied to the plants. Crops well‐infected with mycorrhizal fungi may not benefit from the association if N is limiting¹⁴‘¹⁵ although benefits may appear in soils supplemented with N¹⁵‘¹⁷ even if the level of P in the soil is high¹⁸.

An association of fungi, roots, and bacteria exists in nodulating’ legumes. Maximum benefits from this association may be achieved if N and P supplies are balanced properly. In the present study, the preference of the soybean‐Glomus mosseae‐rhizobial system for form of N was investigated to determine if the mycorrhizal benefits to the soybean could be optimized.     

Separation of tonoplast vesicles enriched in atpase and pyrophosphatase activity from maize roots

Continuous sucrose gradient (15–35%) centrifugation of maize (Zea mays L.) root microsomal membranes yielded two well‐separated fractions of tonoplast vesicles located between 19–21% (Peak I) and 25–26% sucrose (Peak II). Marker enzyme analyses indicated that both fractions were essentially free from plasma membrane, mitochondria and Golgi contaminations. The adenosine triphosphate (ATP) supported proton transport activity was found in both Peak I and Peak II with a 70 to 30% distribution. The pyrophosphatase (PP_;) supported proton transport activity was found only in Peak II. Both hydrolytic activities assumed a bell shape pH dependency with pH optimum at 6.5–7.5 and at 6.5–8.5 for ATPase and PP_; ase, respectively. The K_m of the ATPase and PP_iase, at their respective optimal pH, was found to be 1.2 mM and 0.02 mM, respectively. Both ATPase and PPjase activities were strongly inhibited by N.N'‐dicyclohexylcarbodiimide (DCCD) and diethylstilbestrol (DES) but not by molybdate. Peak I contained nitrate‐sensitive and vanadate‐insensitive ATP hydrolysis activity. In addition to catalyzing the nitrate and vanadate‐insensitive hydrolysis of PP; Peak II also contained some minor ATP hydrolysis activity that was sensitive to vanadate and nitrate. The results indicate that H⁺‐ATPases and H⁺‐PP_fase occur different populations of tonoplast vesicles from corn roots.     

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.     

Ethylene evolution and ammonium accumulation by nutrient‐stressed tomato plants

Environmentally stressed plants frequently have elevated rates of ethylene evolution and high accumulation of free ammonium by their foliage. The objective of this study was to investigate ethylene evolution and ammonium accumulation by nutrient‐deficient and ammonium‐stressed tomato plants (Lycopersicon esculentum Mill. ‘Heinz 1350’ and neglecta‐1) grown in a greenhouse. In soil culture, ‘Heinz 1350’ was more sensitive to ammonium toxicity and had higher ethylene evolution than neglecta‐1. High ethylene evolution corresponded with appearance of ammonium toxicity symptoms in both lines. In sand culture, ‘Heinz 1350’ and neglecta‐1 grown with K, Ca, or Mg deficiency in NO₃ ^‐‐based nutrient solutions had higher ammonium accumulation and higher ethylene evolution than plants grown with complete nutrition. P‐deficient plants had elevated ammonium accumulation but low ethylene evolution. Plants grown on NH₄ ⁺‐based nutrition with pH buffering by CaCO₃ had lower ethylene evolution and lower ammonium accumulation than plants grown in unbuffered solutions but had higher values than plants grown with NO₃ ^‐‐based nutrition. Adequate K nutrition suppressed ethylene evolution and ammonium accumulation for all plants regardless of nitrogen regimes. Ammonium accumulation and ethylene biosynthesis in plants appear to be related processes. They appear to be indicators of stress and may have roles in development of symptoms of nutritional stresses.     

Die kurzfristige pH-Pufferung von gestörten und ungestörten Waldbodenproben

Short-timed pH-buffering of disturbed and undisturbed forest soil samples The pH buffering of disturbed and undisturbed soils under spruce (podzol and podzolic cambisol derived from phyllite, eutric cambisol derived from basalt) was studied in the laboratory by adding H₂SO₄ in ecologically relevant concentrations (pH 5.6–2.0). For the cambisol with crumb structure no difference was found. 80–90% of the added protons were neutralized by release of Ca and Mg. Disturbed samples of the podzol buffer less than 70% of the applicated acid. For undisturbed samples the maximum buffering rate of 185 g H⁺/ha · h is reached with a proton load of about 500 g H⁺/ha · h (related to 4 cm soil depth). Buffering behaviour of the podzolic cambisol lies between the podzol and the cambisol. 70–90% of the proton input is buffered in the disturbed samples while the undisturbed one does not reach its maximum buffering rate, even with high proton load. In this soil Al-release is the most reactive buffer.     

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.     

Characteristics of ammonium and nitrate fluxes along the roots of Picea asperata

Nitrogen (N), ammonium (NH₄⁺) and nitrate (NO₃^?), is one of the key determinants for plant growth. The interaction of both ions displays a significant effect on their uptake in some species. In the current study, net fluxes of NH₄⁺ and NO₃^? along the roots of Picea asperata were determined using a Non-invasive Micro-test Technology (NMT). Besides, we examined the interaction of NH₄⁺ and NO₃^? on the fluxes of both ions, and the plasma membrane (PM) H⁺-ATPases and nitrate reductase (NR) were taken into account as well. The results demonstrated that the maximal net NH₄⁺ and NO₃^? influxes were detected at 13–15?mm and 8–10.5?mm from the root apex, respectively. Net NH₄⁺ influx was significantly stimulated with the presence of NO₃^?, whereas NH₄⁺ exhibited a markedly negative effect on NO₃^? uptake in the roots of P. asperata. Also, our results indicated that PM H⁺-ATPases and NR play a key role in the control of N uptake.