Response to inoculation and N fertilization for increased yield and biological nitrogen fixation of common bean (Phaseolus vulgaris L.)

Common bean (Phaseolus vulgaris L.) is able to fix 20–60 kg N ha^–1 under tropical environments in Brazil, but these amounts are inadequate to meet the N requirement for economically attractive seed yields. When the plant is supplemented with N fertilizer, N₂ fixation by Rhizobium can be suppressed even at low rates of N. Using the ¹⁵N enriched method, two field experiments were conducted to compare the effect of foliar and soil applications of N-urea on N₂ fixation traits and seed yield. All treatments received a similar fertilization including 10 kg N ha^–1 at sowing. Increasing rates of N (10, 30 and 50 kg N ha^–1) were applied for both methods. Foliar application significantly enhanced nodulation, N₂ fixation (acetylene reduction activity) and yield at low N level (10 kg N ha^–1). Foliar nitrogen was less suppressive to nodulation, even at higher N levels, than soil N treatments. In the site where established Rhizobium was in low numbers, inoculation contributed substantially to increased N₂ fixation traits and yield. Both foliar and soil methods inhibited nodulation at high N rates and did not significantly increase bean yield, when comparing low (10 kg N ha^–1) and high (50 kg N ha^–1) rates applied after emergence. In both experiments, up to 30 kg N ha^–1 of biologically fixed N₂ were obtained when low rates of N were applied onto the leaves.     

Nitrogen fixation by the woody legumeLeucaena leucocephala in Tanzania

Summary The nitrogen fixation rate in a 4-year-old stand of the woody legumeLeucaena leucocephala (Lam.) de Wit. was estimated in the field at a rather dry site in Tanzania by use of an acetylene reduction technique. The diurnal mean value during April–May was 35 nmol C₂H₄ mg^–1 (dry weight) nodules h^–1, with a variation between 22±8 and 48±12 nmol C₂H₄ mg^–1 (dry weight) nodules h^–1 in early morning and at midday, respectively. The nodule biomass was determined by auger sampling to be 51±16 kg (dry weight) ha^–1. Most of the nodules were found at the 10–30 cm soil depth level. A rough calculation of the amount of nitrogen fixed annually arrived at 110±30 kg ha^–1. The results give strong support for the use ofL. leucocephala for soil enrichment in less humid areas of tropical Africa.     

Persistence and competitiveness of threeBradyrhizobium japonicum inoculant strains in clay loam Nile valley soil

The nitrogen contribution from the shoot and root system of symbiotically grown leucaena was evaluated in a field experiment on an Alfisol at IITA in Southern Nigeria. Maize in plots that received prunings from inoculated leucaena contained more N and grain yield was increased by 1.9 t.ha.^–1. Large quantities of nitrogen were harvested with leucaena prunings (300 kg N ha^–1 in six months) but the efficiency of utilization of this nitrogen by maize was low compared to inorganic N fertilizer (ammonium sulphate) at 80 kg N ha^–1. Maize yield data indicated that nitrogen in leucaena prunigs was 34 and 45% as efficient as 80 kg N ha^–1 of (NH₄)₂SO₄ for uninoculated and inoculated plants with Rhizobium IRc 1045, respectively. In plots where the prunings were removed, the leaf litter and decaying roots and nodules contributed N equivalent of 32 kg ha^–1. Twenty-five kg ha^–1 was the inorganic N equivalent from nitrogen fixed symbiotically by leucaena when inoculated with Rhizobium strain IRc 1045. Application of prunings from inoculated leucaena resulted in higher soil ogranic C, total N, pH and available NO₃.     

Water chemistry and nutrient budgets in an undisturbed evergreen rainforest of southern Chile

In a pristine evergreen rainforest of Nothofagus betuloides, located at the Cordillera de los Andes in southern Chile (41 °S), concentrations and fluxes of nutrients in bulk precipitation, cloud water, throughfall water, stemflow water, soil infiltration and percolation water and runoff water were measured. The main objectives of this study were to investigate canopy-soil-atmosphere interactions and to calculate input-output budgets. From May 1999 till April 2000, the experimental watershed received 8121 mm water (86% incident precipitation, 14% cloud water), of which the canopy intercepted 16%. Runoff water volume amounted 9527 mm. Bulk deposition of inorganic (DIN) and organic (DON) nitrogen amounted 3.6 kg ha^–1 year^–1 and 8.2 kg ha^–1 year^–1 respectively. Occult deposition (clouds + fog) contributes for 40% to the atmospheric nitrogen input (bulk + occult deposition) of the forest. An important part of the atmospheric ammonium deposition is retained within the canopy or converted to nitrate or organic nitrogen by epiphytic bacteria or lichens. Also the export of inorganic (0.9 kg ha^–1 year^–1) and organic (5.2 kg ha^–1 year^–1) nitrogen via runoff is lower than the input to the forest floor via throughfall and stemflow water (3.2 kg DIN ha–1 year^–1 and 5.6 kg DON ha^–1 year^–1). The low concentrations of NO ₃ ^– and NH ₄ ⁺ under the rooting depth suggest an effective biological immobilization by vegetation and soil microflora. Dry deposition and foliar leaching of base cations (K⁺, Ca²⁺, Mg²⁺) was estimated using a canopy budget model. Bulk deposition accounted for about 50% of the total atmospheric input. Calculated dry and occult deposition are both of equal value (about 25%). Foliar leaching of K⁺, Ca²⁺, and Mg²⁺ accounted for 45%, 38% and 6% of throughfall deposition respectively. On an annual basis, the experimental watershed was a net source for Na⁺, Ca²⁺ and Mg²⁺.     

Effects of planted tree fallows on soil nitrogen dynamics,above-ground and root biomass,N2-fixation and subsequent maize crop productivity in Kenya

The effects of planted fallows of Sesbania sesban (L.) Merr. and Calliandra calothyrsus (Meissner) on soil inorganic nitrogen dynamics and two subsequent maize crops were evaluated under field conditions in the highlands of eastern Kenya. Continuous unfertilised maize, maize/bean rotation and natural regrowth of vegetation (weed fallow) were used as control treatments. The proportion of symbiotic N₂-fixation was estimated by measuring both leaf ¹⁵N enrichment and whole-plant ¹⁵N enrichment by the ¹⁵N dilution technique for Sesbania and Calliandra, using Eucalyptus saligna (Sm.) and Grevillea robusta (A. Cunn) as reference species. Above- and below-ground biomass and N contents were examined in Sesbania, Calliandra, Eucalyptus and Grevillea 22 months after planting. Both the content of inorganic N in the topsoil and the quantity of N mineralised during rainy seasons were higher after the Sesbania fallows than after the other treatments. Compared to the continuous unfertilised maize treatment, both residual crop yields were significantly higher when mineral N (one application of 60 kg N ha^–1) was added. Furthermore, the second crop following the Sesbania fallow was significantly higher than the continuous maize crop. The above-ground biomass of the trees at final harvest were 31.5, 24.5, 32.5 and 43.5 Mg ha^–1 for the Sesbania, Calliandra, Grevillea and Eucalyptus, respectively. For the total below-ground biomass the values for these same tree species were 11.1, 15.5, 17.7, and 19.1 Mg ha^–1, respectively, of which coarse roots (>2 mm), including tap roots, amounted to 70–90%. About 70–90% of the N in Sesbania, and 50–70% in Calliandra, was derived from N₂-fixation. Estimates based on leaf ¹⁵N enrichment and whole-plant ¹⁵N enrichment were strongly correlated. The N added by N₂-fixation amounted to 280–360 kg N ha^–1 for Sesbania and 120–170 kg N ha^–1 for Calliandra, resulting in a positive N balance after two maize cropping seasons of 170–250 kg N ha^–1 and 90–140 kg N ha^–1, for Sesbania and Calliandra, respectively. All the other treatments gave negative N balances after two cropping seasons. We conclude that Sesbania sesban is a tree species well suited for short duration fallows due to its fast growth, high nutrient content, high litter quality and its ability to fix large amounts of N₂ from the atmosphere.     

Estimation of residual N effect of faba bean and pea on two succeeding cereals using15N methodology

A field experiment was conducted using¹⁵N methodology to study the effect of cultivation of faba bean (Vicia faba L.), pea (Pisum sativum L.) and barley (Hordeum vulgare L.) on the N status of soil and their residual N effect on two succeeding cereals (sorghum (Sorghum vulgare) followed by barley). Faba bean, pea and barley took up 29.6, 34.5 and 53.0 kg N ha^–1 from the soil, but returned to soil through roots only 11.3, 10.8 and 5.7 kg N ha^–1, respectively. Hence, removal of faba bean, pea and barley straw resulted in a N-balance of about –18, –24, and –47 kg ha^–1 respectively. A soil nitrogen conserving effect was observed following the cultivation of faba bean and pea compared to barley which was of the order of 23 and 18 kg N ha^–1, respectively. Cultivation of legumes resulted in a significantly higher A_N value of the soil compared to barley. However, the A_N of the soil following fallow was significantly higher than following legumes, implying that the cultivation of the legumes had depleted the soil less than barley but had not added to the soil N compared to the fallow. The beneficial effect of legume cropping also was reflected in the N yield and dry matter production of the succeeding crops. Cultivation of legumes led to a greater exploitation of soil N by the succeeding crops. Hence, appreciable yield increases observed in the succeeding crops following legumes compared to cereal were due to a N-conserving effect, carry-over of N from the legume residue and to greater uptake of soil N by the succeeding crops when previously cropped to legumes.     

Preliminary compositional nutrient diagnosis norms for cowpea (Vigna unguiculata (L.) Walp.) grown on desert calcareous soil

This study calculated the compositional nutrient diagnosis (CND) norms of cowpea (Vigna unguiculata (L.) Walp.), as well as identified significant nutrient interactions of this crop growing in an irrigated calcareous desert soil. Three genotypes were distributed in rows in a 2-ha field. The soil showed high heterogeneity in its chemical properties. For statistical analysis, 86 foliar composite samples from healthy plants were used. Preliminary CND norms were developed using a cumulative variance ratio function and the ² distribution function. Means and standard deviations of row-centered log ratios V_X of five nutrients (N, P, K, Ca, and Mg) and a filling value R, which included all nutrients not chemically analyzed. Preliminary CND norms are: V_N^*=0.174±0.095, V_P^*=–2.172±0.234, V_K^*=–0.007±0.267, V_Ca^*=–0.022±0.146, V_Mg^*=–1.710±0.132, and V_R5^*=3.728±0.084. These CND norms are associated with dry bean yields higher than 1.88 t ha^–1, and are associated with the following foliar concentrations: 26.2 g N kg^–1, 2.5 g P kg^–1, 22.9 g K kg^–1, 21.6 g Ca kg^–1, and 4 g Mg kg^–1. Cowpea plants growing in desert calcareous soils took up lower amounts of N, P, and K than those considered as optimum in a previous report. Six interactions were strongly indicated for cowpea through principal component analyses: positive for Ca–Mg, and negative for N–Ca, N–Mg, Ca–P, Mg–P, and K–P. Furthermore, two interactions were identified using simple correlations, negative N–P and positive K–Ca.     

Intracanopy variation in nitrogen cycling through leaves is influenced by irradiance and proximity to developing fruit in mature walnut trees

Intracanopy variation in net leaf nitrogen (N) resorption and N cycling through leaves in mature walnut (Juglans regia L. cv Hartley) trees were monitored in 3 different years. Differential irradiance among the spurs sampled was inferred from differences among leaves in dry weight per unit area (LW/LA) which varied from 4.0 mg · cm^–2 to 7.0 mg · cm^–2 in shaded (S) and exposed (E) canopy positions, respectively. Our results, using ¹⁵N-depleted (NH₄)₂SO₄ validated the concept that N influx and efflux through fully expanded leaves occurred concurrently during the period of embryo growth. Additionally, it also suggested that N influx into leaves was substantially greater in exposed as compared with shaded canopy positions. Because of its well documented phloem immobility, leaf Ca accumulation was used to better estimate the relative influx of N into exposed and shaded leaves. N cycling varied locally within the tree canopy, i. e. Ca (and presumably N) influx was 100% greater in exposed than shaded tree canopy positions, but influx was not influenced significantly by the proximity of developing fruit. In contrast, both the amount and percentage N efflux was significantly greater during embryo growth in fruit-bearing than defruited spurs. Net leaf N resorption averaged 2–4 times greater (25–30%) in fruit-bearing spurs than the 5–10% decrease in the leaf N content in defruited spurs. Since about 90% of leaf N content reportedly occurs as protein, fruit N demand apparently influenced protein turnover and catalysis in associated spur leaves. The amount of leaf N resorption was greater in exposed than shaded positions in the tree canopy in 2 of the 3 years of data collection. Our data show that like leaf N content, N influx, N efflux and net leaf N resorption vary throughout mature walnut tree canopies under the combined local influences of fruiting and irradiance.     

Leaching of nitrate and ammonium in heathland and forest ecosystems in Northwest Germany under the influence of enhanced nitrogen deposition

To study the impact of high atmospheric nitrogen deposition on the leaching of NO₃^– and NH₄⁺ beneath forest and heathland vegetation, investigations were carried out in adjacent forest and heathland ecosystems in Northwest Germany. The study area is subjected to high deposition of nitrogen ranging from 15.9 kg ha^–1 yr^–1 in bulk precipitation to 65.3 kg ha^–1 yr^–1 beneath a stand of Pinus sylvestris L. with NH₄–N accounting for 70–80% of the nitrogen deposited. Considerable leaching of nitrogen compounds from the upper horizons of the soil, mostly as nitrate, occurred at most of the forest sites and below a mixed stand of Calluna vulgaris (L.) Hull. and Erica tetralix, but was low in a Betula pubescens Ehrh. swamp forest as well as beneath Erica tetralix L. wet heath and heath dominated by Molinia caerulea(L.) Moench. Ground water concentrations of both NO₃–N and NH₄–N did not exceed 1 mg L^–1 at most of the sites investigated.     

Biomass and litter dynamics in a Melaleuca forest on a seasonally inundated floodplain in tropical,northern Australia

Litterfall from a Melaleuca forest was investigated as part of chemical cycling studies on the Magela Creek floodplain in tropical, northern Australia. The forest contained two species of tree, Melaleuca cajaputi and Melaleuca viridiflora, with a combined average density of 294 trees ha^–1. The M. viridiflora trees had diameter breast height measurements ranging from 11.8 to 62.0 cm, median class 25.1–30.0cm and a mean value of 29.2±1.0 cm, compared to 13.0 to 66.3 cm, 30.1–35.0cm and 33.5±1.0cm for M. cajaputi trees. A regression model between tree height, diameter breast height and fresh weight was determined and used to calculate average tree weights of 775±1.6kg for M. viridiflora and 1009±1.6kg for M. cajaputi, and a total above-ground fresh weight of 263±0.3t ha^–1. The weight of litter recorded each month on the ground beneath the tree canopy ranged from 582±103 to 2176±376 g m^–2 with a monthly mean value of 1105±51 g m^–2. The coefficient of variation of 52% on this mean indicates the large spatial and temporal variability in litter distribution over the study site. This variability was greatly affected by the pattern of water flow and litter transport during the Wet season. Litterfall from the trees was evaluated using two techniques - nets and trays. The results from these techniques were not significantly different with annual litterfall collected in the nets being 705 ± 25 g m^–2 and in the trays 716±49 g m^–2. The maximum monthly amount of litterfall, 108 ±55g m^–2, occurred during the Dry season months of June–July. Leaf material comprised 70% of the total annual weight of litter, 480±29 g m^–2 in the nets and 495 ± 21 g m^–2 in the trays. The tree density and weight of litter suggest that the Melaleuca forests are highly productive and contribute a large amount of material to the detrital/debris turnover cycle on the floodplain.     

Application of 15N and xylem ureide methods for assessing N2 fixation of three shrub legumes periodically pruned for forage

The apparently diminished capacity for N₂ fixation by the shrub legume Calliandra calothyrsus (Calliandra) relative to other woody perennial legumes was investigated in a field experiment in northern Queensland, Australia. In this trial, (i) the proportion of plant nitrogen (N) derived from symbiotic N₂ fixation (%Pfix) and the amounts of N₂ fixed were compared in Calliandra, Gliricidia sepium (Gliricidia) and Codariocalyx gyroides (Codariocalyx), (ii) variations in N₂ fixation due to season or tree age were determined, (iii) estimates of Pfix derived with the ¹⁵N natural abundance technique were compared with values obtained from ¹⁵N enrichment or xylem sap ureide procedures to determine whether the previous conclusions about Calliandra's ability to fix N had resulted from specific problems with the natural abundance methodology used in the earlier studies.Inoculated seedlings of each of the three shrub legume species were planted in dense stands (1.5 m rows, 0.5 m between trees) in two randomised blocks. The northern block was used solely for natural abundance measurements, while ¹⁵N-enriched KNO₃ (10 atom % ¹⁵N excess) was applied four times over a 52 week period to plots in the southern block. The non-nodulating tree legume Senna spectabilis (formally Cassia spectabilis) was used as a non-N₂-fixing reference for the ¹⁵N-based procedures, with Guinea grass (Panicum maximum) included as an additional non-fixing check. Growth by the trees above 75 cm was first cut and removed after 22 weeks and regrowth was subsequently pruned periodically for another 95 weeks. Sampling for dry matter production, N yield and estimates of Pfix were restricted to the central four of the 32 plants which constituted each replicate plot. Information generated during the 117 week study indicated that estimates of Pfix by ¹⁵N natural abundance were closely similar to values derived with ¹⁵N-enrichment or sap ureides. The data indicated that Calliandra had a reduced reliance upon N₂ fixation relative to Gliricidia and Codariocalyx for the first 65 weeks after establishment. This appeared to be due to more prolifc root growth by Calliandra than either of the other N₂-fixing species and an ability to extract a greater proportion of its N requirements from soil mineral N. However, after week 65 and for the remainder of the experiment, estimates of Pfix for Calliandra were similar to the other shrub legumes. Over 117 weeks, prunings from Calliandra and Gliricidia had removed 52–58 t dry matter ha^-1, and between 1471 and 1678 kg N ha^-1, of which 1026–1063 kg N ha^-1 was estimated to have been derived from N₂ fixation. At the time of final harvest, 65–73% of the fixed N was present in shoot regrowth of the N₂ fixing shrubs, 9–18% in the roots, 15% in the trunk, and 2–6% in fallen leaves.     

Primary production and nitrogen allocation of field grown sugar maples in relation to nitrogen availability

Above ground net primary production (NPP), nitrogen (N) allocation, and retranslocation from senescing leaves were measured in 7 sugar-maple dominated sites having annual net N mineralization rates ranging from 26 to 94 kg · ha^–1 · yr^–1. The following responses were observed: (1) Green sun leaves on richer sites had higher N mass per unit leaf area than sun leaves on poorer sites; (2) Total canopy N varied much less than annual net mineralization, ranging from 81 to 111 kg · ha^–1; (3) This was due to the existence of a large and relatively constant pool of N which was retranslocated from senescing leaves for use the following year (54 to 80 kg · ha^–1); (4) The percentage of canopy N retranslocated by sugar maple was also relatively constant, but was slightly higher on the richer sites. Percent N in leaf litter did not change across the gradient; (5) Above ground NPP increased linearly in relation to N allocated above ground. Therefore, N use efficiency, expressed as above ground NPP divided by N allocated above ground was constant; (6) N use efficiency expressed as (NPP above ground/total N availability) was a curvilinear function of N availability; and (7) This pattern reflected a decreasing apparent allocation of N below ground with decreasing N availability.     

Micropropagation of thirteen Malus cultivars and rootstocks,and effect of antibiotics on proliferation

Several factors that affect in vitro establishment, proliferation, and rooting of thirteen Malus cultivars and rootstocks were studied. Apple shoot tips (1.5±0.5 cm in length) were established using ascorbic and citric acids as antioxidants. Four proliferation media containing 1.0 mg 1^–1 BA and different concentrations of IBA and GA₃ were tested. Proliferation rates varied depending on the genotype and medium used. The highest proliferation rate was obtained for a rootstock that produced 11.6±2.5 shoots (1.5±0.8 cm in length) per tube per month. Rooting was induced with IBA for all the genotypes tested. The optimal IBA concentration was cultivar dependent (between 0.1 and 1.0 mg 1^–1 IBA), and lower concentrations were necessary to induce rooting in liquid rather than in solid medium.The effects on shoot-tip proliferation of cefotaxime, carbenicillin and kanamycin, three antibiotics commonly used for transformation studies, were also evaluated. Cefotaxime at 200 mg 1^–1 stimulated shoot growth and development, but at 500 mg 1^–1 caused abnormal shoot morphology. Carbenicillin at 500 mg 1^–1, alone or in combination with cefotaxime at 200 mg 1^–1, inhibited proliferation and caused excessive enlargement of the basal leaves, inducing callus formation and release of phenolic compounds in the medium. Kanamycin at 50 mg 1^–1 was phytotoxic and caused shoot chlorosis and necrosis. Consideration of the toxicity of these antibiotics is critical when designing transformation schemes for selection and recovery of transgenic apple plants.Abbreviations BA benzyladenine - cef cefotaxime - crb carbenicillin - GA₃ gibberellic acid - IBA indole-3-butyric acid - Kan kanamycin - ms Murashige and Skoog [19] macro- and micro-nutrients - NAA naphthalene-acetic acid     

Growing vetches (Vicia villosa Roth) in irrigated cotton systems: inputs of fixed N, N fertiliser savings and cotton productivity

We compared symbiotic N₂ fixation by winter forage legumes (clovers, medics and vetches) using the ¹⁵N natural abundance technique in three experiments. Vetches (Vicia spp.) were the most productive legumes, and woollypod vetch fixed (shoot+root) up to 265 kg N ha^–1 (mean 227 kg N ha^–1) during a 4–5 months period over winter and early spring. Balansa and Berseem clovers, and Gama medic were highly productive in the first experiment, but fixed significantly less N than woollypod vetch in the second experiment. A 6-year study (1997–2003) compared cotton (Gossypium hirsutum L.) systems with and without vetch, or with faba beans (Vicia faba L.) to assess the effects of these crops on cotton production. Woollypod vetch was grown either between annual cotton crops, or between wheat (Triticum aestivumL.) and cotton crops. Vetch added 230 kg N ha^–1 (174 kg fixed N ha^–1) to the soil when incorporated as a green manure. Faba bean shoot residues and nodulated roots contributed 108 kg fixed N ha^–1 to the soil, following the removal of 80 kg N ha^–1 in the harvested seed (meaned over three crops). Lablab (Lablab purpureus L. – summer-growing and irrigated) added 277 kg N ha^–1 (244 kg fixed N ha^–1) before incorporation as a green manure in the first year of the experiment. The economic optimum N fertiliser rate for each cropping system was determined every second year when all systems were sown to cotton. Cotton following cotton required 105 kg fertiliser N ha^–1, but only 40 kg N ha^–1 when vetch was grown between each cotton crop. Cotton following wheat required 83 kg fertiliser N ha^–1 but no N fertiliser was needed when vetch was grown after wheat (the highest yielding system). Cotton following faba beans also required no N fertiliser. The vetch-based systems became more N fertile over the course of the experiment and produced greater lint yields than the comparative non-legume systems, and required less N fertiliser. While no cash flow was derived from growing vetch, economic benefits accrued from enhanced cotton yields, reduced N fertiliser requirements and improved soil fertility. These findings help explain the rotational benefits of vetches observed in other regions of the world.     

Estimation of biological nitrogen fixation associated withBrachiaria andPaspalum grasses using15N labelled organic matter and fertilizer

Summary Six pasture grasses,Paspalum notatum cv batatais,P. notatum cv pensacola,Brachiaria radicans, B. ruziziensis, B. decumbens andB. humidicola, were grown in concrete cylinders (60 cm diameter) in the field for 31 months. The soil was amended with either a single addition of¹⁵N labelled organic matter or frequent small (2 kg N. ha^–1) additions of¹⁵N enriched (NH₄)₂SO₄. In the labelled fertilizer treatment soil analysis revealed that there was a very drastic change in¹⁵N enrichment in plant-available nitrogen (NO ₃ ^– +NH ₄ ⁺ ) with depth. The different grass cultivars recovered different quantities of applied labelled N, and evidence was obtained to suggest that the roots exploited the soil to different depths thus obtaining different¹⁵N enrichments in soil derived N. This invalidated the application of the isotope dilution technique to estimate the contribution of nitrogen fixation to the grass cultivars in this treatment. In the labelled organic matter treatment the¹⁵N label in the plant-available N declined at a decreasing rate during the experiment until in the last 12 months the decrease was only from 0.274 to 0.222 atom % excess. There was little change in¹⁵N enrichment of available N with depth, hence it was concluded that although the grasses recovered different quantities of labelled N, they all obtained virtually the same¹⁵N enrichment in soil derived N. Data from the final harvests of this treatment indicated thatB. humidicola andB. decumbens obtained 30 and 40% respectively of their nitrogen from N₂ fixation amounting to an input of 30 and 45 kg N.ha^–1 year^–1 respectively.     

Deposition of ammonia to the forest floor under spruce and beech at the Höglwald site

Laboratory and field measurements of the flux of ammonia to forest floor canopies of spruce and beech stands at the Höglwald site in southern Bavaria are reported. Measurements were performed with an open chamber method. A linearity between ammonia concentration and ammonia flux from the atmosphere to the ground floor canopy was detected. Deposition of ammonia showed no saturation even at air concentrations up to 50 g NH₃ m^–3 air. Temperature, water content and the moss layer of the ground floor canopy had a minor influence on the deposition velocity in laboratory experiments. Deposition velocity of ammonia was higher to the spruce (1.3 cm s^–1), and limed spruce ground floor canopy (1.17 cm s^–1) compared to the beech stand (0.79 cm s^–1). In field studies, a diurnal course of the deposition velocity was detected with highest velocities in midday and minor during night times, but not in the climatic chamber. The flux of ammonia to the ground floor canopy was estimated of app. 10 kg N ha^–1 yr^–1 for the soil under spruce, 9 kg N ha^–1 yr^–1 for the limed spruce and 6 kg N ha^–1yr^–1 for the soil under beech. The fluxes are interpreted as fluxes from the atmosphere to the ground canopies of the stands.     

Nitrogen fixation by white clover in pastures grazed by dairy cows: Temporal variation and effects of nitrogen fertilization

Effects of rate of nitrogen (N) fertilizer and stocking rate on production and N₂ fixation by white clover (Trifolium repens L.) grown with perennial ryegrass (Lolium perenne L.) were determined over 5 years in farmlets near Hamilton, New Zealand. Three farmlets carried 3.3 dairy cows ha^–1 and received urea at 0, 200 or 400 kg N ha^–1 yr^–1 in 8–10 split applications. A fourth farmlet received 400 kg N ha^–1 yr^–1 and had 4.4 cows ha^–1.There was large variation in annual clover production and total N₂ fixation, which in the 0 N treatment ranged from 9 to 20% clover content in pasture and from 79 to 212 kg N fixed ha^–1 yr^–1. Despite this variation, total pasture production in the 0 N treatment remained at 75–85% of that in the 400 N treatments in all years, due in part to the moderating effect of carry-over of fixed N between years.Fertilizer N application decreased the average proportion of clover N derived from N₂ fixation (P_N; estimated by ¹⁵N dilution) from 77% in the 0 N treatment to 43–48% in the 400 N treatments. The corresponding average total N₂ fixation decreased from 154 kg N ha^–1 yr^–1 to 39–53 kg N ha^–1 yr^–1. This includes N₂ fixation in clover tissue below grazing height estimated at 70% of N₂ fixation in above grazing height tissue, based on associated measurements, and confirmed by field N balance calculations. Effects of N fertilizer on clover growth and N₂ fixation were greatest in spring and summer. In autumn, the 200 N treatment grew more clover than the 0 N treatment and N₂ fixation was the same. This was attributed to more severe grazing during summer in the 0 N treatment, resulting in higher surface soil temperatures and a deleterious effect on clover stolons.In the 400 N treatments, a 33% increase in cow stocking rate tended to decrease P_N from 48 to 43% due to more N cycling in excreta, but resulted in up to 2-fold more clover dry matter and N₂ fixation because lower pasture mass reduced grass competition, particularly during spring.     

Nitrogen incorporation and remobilization in different shoot components of field-grown winter oilseed rape (Brassica napus L.) as affected by rate of nitrogen application and irrigation

The seasonal course of nitrogen uptake, incorporation and remobilization in different shoot components of winter oilseed rape (Brassica napus L.) was studied under field conditions including three rates of ¹⁵N labelled nitrogen application (0, 100 or 200 kg N ha^-1) and two irrigation treatments (rainfed or watered at a deficit of 20 mm). The total amount of irrigation water applied was 260 mm, split over 13 occasions in a 7-week-period ranging from 1 week before onset of flowering until 4 weeks after flowering.Nitrogen application and irrigation increased plant growth and nitrogen accumulation. Irrespective of N and irrigation treatment more than 50% of total shoot N was present in the stem when flowering started. At the end of flowering, pod walls were the main N store containing about 30–40% of shoot N. The quantities of N remobilized from stems and pod walls amounted in all treatments to about 70% of the N present in these organs at mid-flowering. At harvest, stem and pod walls each contained about 10% of total shoot N, the remaining 80% being incorporated into seeds. The main component contributing to the response of seed N accumulation to nitrogen application and irrigation was pods in axillary racemes. Up to 20 kg N ha^-1, corresponding to about 10% of final shoot N content, was lost from the plants by leaf drop.Irrigation increased the recovery at harvest of applied N from 30% to about 50%, while the level of N application did not affect the N recovery. ¹⁵N labelled (fertilizer derived) nitrogen constituted a greater proportion of the N content in old leaves than in young leaves and increased with age in the former, but not in the latter. Relative to soil N, fertilizer derived N also contributed more to the N content of vegetative than to that of reproductive shoot components. Small net changes in shoot N content after flowering reflected a balance between N import and export, leading to continuous dilution of ¹⁵N labelled N with unlabelled N.     

Seasonal changes in apparent hydraulic conductance and their implications for water use of European beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) and sessile oak [<Emphasis Type="Italic">Quercus petraea</Emphasis> (Matt.) Liebl] in South Europe

The water status of Fagus sylvatica L. and Quercus petraea (Matt) Liebl. was analysed during a cycle of progressive natural drought in southern Europe. Predawn (Ψ_pd) and midday water potential were measured in transpiring (Ψ_leaf) and non-transpiring leaves (Ψ_xyl). Furthermore, photosynthesis (A), stomatal conductance to water vapour (g_s) and sap flow (F_d) were recorded on the same dates. Apparent leaf specific hydraulic conductance in the soil–plant–air continuum (K_h) and whole tree hydraulic conductance (K_hsf) were calculated by using the simple analogy of the Ohm’s law. K_h was estimated at different points in the pathway as the ratio between transpiration (E) in the uppermost canopy leaves at midday and the gradient of water potential in the different compartments of the continuum soil–roots–stem–branches–leaves. There was a progressive decrease in water potential measured on non-transpiring leaves at the base of tree crown in both species (Ψ^l_xyl) from the beginning of the growing season to the end of summer. A similar decrease was shown in shoot water potential (Ψ^u_xyl) at the uppermost canopy. Predawn water potential (Ψ_pd) was high in both species until late July (28 July); afterwards, a significant decrease was registered in F. sylvatica and Q. petraea with minimum values of −0.81±0.03 and −0.75±0.06 MPa, respectively, by 15 September. In both species, leaf specific hydraulic conductance in the overall continuum soil–plant–air (K_h) decreased progressively as water stress increases. Minimum values of K_h and K_hsf were recorded when Ψ_pd was lower. However, Q. petraea showed higher K_h than F. sylvatica for the same Ψ_pd. The decrease in K_h with water stress was mainly linked to its fall from the soil to the lowermost canopy (K_srs). Nevertheless, a significant resistance in the petiole–leaf lamina (K_pl) was also recorded because significant differences in all dates were found on Ψ between transpiring and non-transpiring leaves from the same shoot. The decline in K_h was followed by an increase in stomatal control of daily water losses through the decrease of stomatal conductance to water vapour (g_s) during the day. It promoted a seasonal increase in the stomatal limitation to carbon dioxide uptake for photosynthesis (A). These facts were more relevant in F. sylvatica, which had concurrently a higher decline in water use at the tree level than Q. petraea. The results showed a strong coupling in F. sylvatica and Q. petraea between processes at leaf and tree level. It may be hypothesised a role of specific hydraulic conductance not only in the regulation of water losses by transpiration but also of carbon uptake.     

Exchange of N-gases at the Höglwald Forest – A summary

During 4 years continuous measurements of N-trace gas exchange were carried out at the forest floor-atmosphere interface at the Höglwald Forest that is highly affected by atmospheric N-deposition. The measurements included spruce control, spruce limed and beech sites. Based on these field measurements and on intensive laboratory measurements of N₂-emissions from the soils of the beech and spruce control sites, a total balance of N-gas emissions was calculated. NO₂-deposition was in a range of –1.6 –2.9 kg N ha^–1 yr^–1 and no huge differences between the different sites could be demonstrated. In contrast to NO₂-deposition, NO- and N₂O-emissions showed a huge variability among the different sites. NO emissions were highest at the spruce control site (6.4–9.1 kg N ha^–1 yr^–1), lowest at the beech site (2.3–3.5 kg N ha^–1 yr^–1) and intermediate at the limed spruce site (3.4–5.4 kg N ha^–1 yr^–1). With regard to N₂O-emissions, the following ranking between the sites was found: beech (1.6–6.6 kg N ha^–1 yr^–1) >> spruce limed (0.7–4.0 kg N ha^–1 yr^–1) > spruce control (0.4–3.1 kg N ha^–1 yr^–1). Average N-trace gas emissions (NO, NO₂, N₂O) for the years 1994–1997 were 6.8 kg N ha^–1 yr^–1 at the spruce control site, 3.6 kg N ha^–1 yr^–1 at the limed spruce site and 4.5 kg N ha^–1 yr^–1 at the beech site. Considering N₂-losses, which were significantly higher at the beech (12.4 kg N ha^–1 yr^–1) than at the spruce control site (7.2 kg N ha^–1 yr^–1), the magnitude of total gaseous N losses, i.e. N₂-N + NO-N + NO₂-N + N₂O-N, could be calculated for the first time for a forest ecosystem. Total gaseous N-losses were 14.0 kg N ha^–1 yr^–1 at the spruce control site and 15.5 kg N ha^–1 yr^–1 at the beech site, respectively. In view of the huge interannual variability of N-trace gas fluxes and the pronounced site differences in N-gas emissions it is concluded that more research is needed in order to fully understand patterns of microbial N-cycling and N-gas production/emission in forest ecosystems and mechanisms of reactions of forest ecosystems to the ecological stress factor of atmospheric N-input.