Nitrogen fertilization alters the functioning of arbuscular mycorrhizas at two semiarid grasslands

The effects of nitrogen (N) fertilization on arbuscular mycorrhizas were studied at two semiarid grasslands with different soil properties and N-enrichment history (Shortgrass Steppe in Colorado, and Sevilleta National Wildlife Refuge in New Mexico). These sites are part of the National Science Foundation's Long-Term Ecological Research Network. The experimental plots at Shortgrass Steppe were fertilized with ammonium nitrate (NH₄NO₃) from 1971 to 1975, and have not received additional N since then. The experimental plots at Sevilleta were also fertilized with NH₄NO₃, but were established in 1995, 2 years before the soils were used for this study. Greenhouse experiments were conducted to compare the growth response of local grasses to arbuscular mycorrhizal (AM) fungi from fertilized (FERT) and unfertilized (UNFERT) field soils, at each site. Two species per site were chosen, Bouteloua gracilis and Elymus elymoides from Shortgrass Steppe, and B. gracilis and B. eriopoda from Sevilleta. Plants were grown for 3 months at HIGH N and LOW N levels, with FERT or UNFERT soil inoculum and in a non-mycorrhizal condition. Fertilization with N altered the functioning of AM fungi at both sites. Grasses inoculated with AM fungi from UNFERT soils had the most tillers, greatest biomass and highest relative growth rates. There were no significant differences in the growth response of plants inoculated with AM fungi from FERT soils and the non-mycorrhizal controls. These results were consistent across sites and species except for the plants grown at LOW N in Sevilleta soils. These plants were deficient in N and phosphorus (P) and did not show growth enhancement in response to AM inoculation with either FERT or UNFERT soils. Percent root length colonized by AM fungi was not directly related to plant performance. However, enrichment with N consistently decreased root colonization by AM fungi in the grasses grown in soils from Shortgrass Steppe with high P availability (18.4 mg kg^–1), but not in the grasses grown in Sevilleta soils with low P availability (6.6 mg kg^–1). Our study supports the hypotheses that (1) fertilization with N alters the balance between costs and benefits in mycorrhizal symbioses and (2) AM fungal communities from N fertilized soils are less beneficial mutualists than those from unfertilized soils.     

Microbial biomass and nitrogen cycling responses to fertilization and litter removal in young northern hardwood forests

The influence of site fertility on soil microbial biomass and activity is not well understood but is likely to be complex because of interactions with plant responses to nutrient availability. We examined the effects of long-term (8 yr) fertilization and litter removal on forest floor microbial biomass and N and C transformations to test the hypothesis that higher soil resource availability stimulates microbial activity. Microbial biomass and respiration decreased by 20–30 % in response to fertilization. Microbial C averaged 3.8 mg C/g soil in fertilized, 5.8 mg C/g in control, and 5.5 mg C/g in litter removal plots. Microbial respiration was 200 µg CO₂-C g^–1 d^–1 in fertilized plots, compared to 270 µg CO₂-C g^–1 d^–1 in controls. Gross N mineralization and N immobilization did not differ among treatments, despite higher litter nutrient concentrations in fertilized plots and the removal of substantial quantities of C and N in litter removal plots. Net N mineralization was significantly reduced by fertilization. Gross nitrification and NO₃ ^– immobilization both were increased by fertilization. Nitrate thus became a more important part of microbial N cycling in fertilized plots even though NH₄ ⁺ availability was not stimulated by fertilization.Soil microorganisms did not mineralize more C or N in response to fertilization and higher litter quality; instead, results suggest a difference in the physiological status of microbial biomass in fertilized plots that influenced N transformations. Respiration quotients (qCO₂, respiration per unit biomass) were higher in fertilized plots (56 µg CO₂-C mg C^–1 d^–1) than control (48 µg CO₂-C mg C^–1 d ^–1) or litter removal (45 µg CO₂-C mg C^–1 d^–1), corresponding to higher microbial growth efficiency, higher proportions of gross mineralization immobilized, and lower net N mineralization in fertilized plots. While microbial biomass is an important labile nutrient pool, patterns of microbial growth and turnover were distinct from this pool and were more important to microbial function in nitrogen cycling.     

Soil nitrogen availability under grassland and cultivated corn

Summary Adjacent corn and ryegrass plots were fertilized with rates of 0, 50, 100, 150, and 200 kg N as ammonium nitrate/ha. Corn growing on this soil did not respond to fertilizer N while ryegrass responded to rates of up to 200 kg N/ha. The differences in N availability was also reflected in the higher profile NO₃–N under corn than under ryegrass. The same general trends occurred on a second soil, where N availability for the hay crop was also less than for corn crop. Compared with corn, hay responded more to N fertilizer and had lower soil NO₃–N levels.Grasslands appear to respond to higher N fertilizer rates than cultivated crops on the same soil.Vermont Agricultural Experiment Station Journal Article No. 495.     

Fine root production and nutrient content in wet and moist arctic tundras as influenced by chronic fertilization

We used ingrowth cores to estimate fine root production in organic soils of wet sedge and moist tundra ecosystems near Toolik Lake on Alaska's North Slope. Root-free soil cores contained in nylon mesh tubes (5 cm diameter, 20–30 cm long) were placed in control and chronically fertilized (N plus P) plots in mid-August 1994 and were retrieved 1 year later. Estimated fine root production in control plots was 75 g m^–2 year^–1 in wet sedge and 56 g m^–2 year^–1 in moist tussock tundra. Fine root production in fertilized plots was 85 g m^–2 year^–1 in wet sedge and 67 g m^–2 year^–1 in moist tussock tundra. Although our estimates of fine root production were higher on fertilized than control plots, differences were not statistically significant within either tundra type. Comparisons between our estimates of fine root production and other estimates of aboveground (plus rhizome) production on the same (wet sedge tundra) or similar (moist tussock tundra) plots suggest that fine root production was about one-third of total net primary production (NPP) under non-fertilized conditions and about one-fifth of total NPP under chronic fertilization. Fine root N and P concentrations increased with fertilization in both tundra types, but P concentrations increased more than N concentrations in wet sedge tundra, whereas relative increases in N and P concentrations in moist tundra roots were similar. These data are consistent with other studies suggesting that NPP in wet sedge tundra is often P limited and that co-limitation by N and P is more important in moist tussock tundra.     

The effect of increased nutrient availability on vegetation dynamics in wet heathlands

A three year fertilization experiment was conducted in which nitrogen (N series: 20 g N m^–2 yr^–1), phosphorus (P series: 4 g P m^–2 yr^–1) and potassium (K series: 20 g K m^–2 yr^–1) were added to a mixed vegetation of Erica tetralix and Molinia caerulea. At the end of each growing season the percentage cover of each species was determined. At the end of the experiment percentage cover of each species was found to be positively correlated with the harvested biomass. In the unfertilized control series the cover of Erica and Molinia did not change significantly during the experiment. In all fertilized series however, especially in the P series, cover of Erica decreased significantly. The cover of Molinia increased significantly in the P series only.In the fertilized series the biomass of Erica and total biomass per plot did not change significantly compared with the control series. In the P series the biomass of Molinia increased significantly.It is suggested that with increasing phosphorus or nitrogen availability Molinia outcompetes Erica because the former invests more biomass in leaves which in turn permits more carbon to be allocated to the root system, which thereupon leads to a higher nutrient uptake.     

Long-term soil nutrient dynamics and lateral nutrient movement in fertilized and unfertilized red pine plantations

In this study, we use arepeated-measures analysis to test thehypothesis that soil fertility underpotassium-limited red pine (Pinusresinosa Ait.) stands at the Charles LathropPack Demonstration Forest in Warrensburg, NewYork is increasing toward a steady state thatwas artificially induced in fertilized standsby K-fertilization over 50 years ago. Wemeasured soil K by horizon and added new datato a 53-year database. We examine onemechanism that explains the higher rate of Kaccumulation in unfertilized stands comparedwith fertilized – lateral movement offertilizer K from treated plots to untreated –using the rubidium/potassium reverse tracermethod. Over the past five decades, soil Kconcentrations under both fertilized andunfertilized red pine have increasedsignificantly. The trends under fertilized andunfertilized plots demonstrate the gradualconvergence of soil K under unfertilized plotstoward concentrations in fertilized plots. Five decades after fertilization, treated soilsstill contain greater concentrations ofexchangeable K and lower bulk densities thanunfertilized plots. Analysis of Rb/K ratios inthe forest floor of fertilized and unfertilizedplots confirms the hypothesis that lateraltransport of surface broadcast fertilizer,applied over 50 years ago, extendsapproximately 11–16 m from the edges offertilized plots. The four unfertilized plotsclosest to fertilized plots have beensignificantly affected by inputs of fertilizerK, while the remaining five plots arerelatively unimpacted. Approximately 36% ofthe K in fertilized plots, and 23% of the K inunfertilized plots affected by fertilizermigration were derived directly from thefertilizer applied 5 decades ago, demonstratingthe highly conservative nature of mineralnutrient cycling in aggrading forests.     

Lysimeter and field studies on15N in a tropical soil

Twenty four plots, each 2.0 m² in area, were established on St. Augustine loam soil series as field plots and microplots (containing lysimeters) in a completely randomised block design of four treatments (mulched fertilized, unmulched fertilized microplots; mulched fertilized and unmulched fertilized field plots), replicated three times. Labelled (¹⁵N) and unlabelled (NH₂)₂CO fertilizer were applied at rates of 400 kg N ha^–1 and CaH₂PO₄ and KCl were applied at rates of 100 and 150 g ha^–1 respectively to the field plots and microplots. Mulch (bagasse) was maintained to a depth of two cm and the plots were kept bare with regular applications of gramoxone.The maximum depth of leaching as measured by diffusion of NO ₃ ^– –¹⁵N in both the dry and wet seasons was 30 cm. The potential for downward movement of water and NO ₃ ^– –¹⁵N was low in the wet season because high intensity rainfall followed high soil moisture contents. Effects of mulching, on the mobility of applied N fertilizers were inconclusive. Infiltration rates were significantly (P=0.25) inversely correlated with soil moisture content, supporting the hypothesis that high intensity rainfall on a saturated soil surface is more likely to result in NO ₃ ^– –¹⁵N dispersion than NO ₃ ^– –¹⁵N leaching.     

An assessment of the contribution of net mineralization to N cycling in grass swards using a field incubation method

Measurements of net mineralization using a field incubation method were made over a full growing season (180 d). Soil cores, taken from cut swards which for many years had been previously grazed by cattle, were placed in jars in the field for successive incubation periods of 14 d. Acetylene was added to the incubation jars to inhibit nitrification in the soil cores and thereby prevent losses of N through denitrification. Net mineralization over 180 d amounted to 415, 321 and 310 kg N ha^–1 under grass/clover, unfertilized grass and grass receiving 420 kg N ha^–1 y^–1, respectively. At the start of the growing season, an index of potentially mineralizable N in the soil was estimated by a chemical extraction method, but this index was <50% of the estimates obtained by field incubation. The amount of N in herbage harvested regularly from the swards also under-estimated the supply of N from the soil, with apparent recoveries of 53, 82 and 74% and total yields of N of 240, 263 and 538 (kg N ha^–1) from grass/clover, unfertilized grass and fertilized grass, respectively. Mineralization rates varied significantly with seasonal soil temperature fluctuations, but the incubation method was apparently less sensitive in relation to changes in soil water content. Rates of N-turnover (as % of total soil N) were highest under grass/clover (9%), but similar under fertilized and unfertilized grass swards (approximately 5%).     

Responses of N fluxes and pools to elevated atmospheric CO2 in model forest ecosystems with acidic and calcareous soils

The objectives of this study were to estimate how soil type, elevated N deposition (0.7 vs. 7 g N m^–2y^–1) and tree species influence the potential effects of elevated CO₂ (370 vs. 570 mol CO₂ mol^–1) on N pools and fluxes in forest soils. Model spruce-beech forest ecosystems were established on a nutrient-rich calcareous sand and on a nutrient-poor acidic loam in large open-top chambers. In the fourth year of treatment, we measured N concentrations in the soil solution at different depths, estimated N accumulation by ion exchange resin (IER) bags, and quantified N export in drainage water, denitrification, and net N uptake by trees. Under elevated CO₂, concentrations of N in the soil solution were significantly reduced. In the nutrient-rich calcareous sand, CO₂ enrichment decreased N concentrations in the soil solution at all depths (–45 to –100%). In the nutrient-poor acidic loam, the negative CO₂ effect was restricted to the uppermost 5 cm of the soil. Increasing the N deposition stimulated the negative impact of CO₂ enrichment on soil solution N in the acidic loam at 5 cm depth from –20% at low N inputs to –70% at high N inputs. In the nutrient-rich calcareous sand, N additions did not influence the CO₂ effect on soil solution N. Accumulation of N by IER bags, which were installed under individual trees, was decreased at high CO₂ levels under spruce in both soil types. Under beech, this decrease occurred only in the calcareous sand. N accumulation by IER bags was negatively correlated with current-years foliage biomass, suggesting that the reduction of soil N availability indices was related to a CO₂-induced growth enhancement. However, the net N uptake by trees was not significantly increased by elevated CO₂. Thus, we suppose that the reduced N concentrations in the soil solution at elevated CO₂ concentrations were rather caused by an increased N immobilisation in the soil. Denitrification was not influenced by atmospheric CO₂ concentrations. CO₂ enrichment decreased nitrate leaching in drainage by 65%, which suggests that rising atmospheric CO₂ potentially increases the N retention capacity of forest ecosystems.     

Rapid whole-plant bioassay for phosphorus phytoavailability in soils

Chemical P extraction from soils is an indirect and frequently questionable index for P availability. To monitor the dynamics of P availability in soils more directly following the application of P fertilizer, manure or sludge, a rapid, whole-plant bioassay was developed using tomato (Lycopersicon esculentum Mill.), Chinese cabbage (Brassica rapa L. var.pekinensis) and wheat (Triticum aestivum L.). Plant P extracted in 0.1 M H₂SO₄ (P_i) and total P (P_t) concentration or content in stem, leaves or whole shoots were highly correlated (P < 0.01) with P fertilizer rates or water-soluble (WSP) or Olsen P in various soils, over wide ranges of soil P status. The whole-plant P_i content was found to be as informative as the more complicated indices of P_t or P_iconcentration. The assay was used to compare availability of fertilizer-P and sewage-sludge-P after incorporation into alluvial soil during 1–100 days of incubation. While both soil and plant indices had shown that fertilizer-P was more highly available than sewage-sludge-P in each period, the bioassay was much more sensitive than the Olsen-P or WSP soil indices in showing P fixation and decrease of availability during incubation time. The bioassay is sufficiently rapid (5–12 days) to allow a study of short-term changes in soil-P availability following incorporation of various P additives, and it is applicable to a very wide range of P availability values (6–535 mg Olsen-P kg^–1), extending from lower than desired for crop production to higher than permitted from an environmental standpoint.     

Phosphorus regulation of nitrogen fixation in a traditional Mexican agroecosystem

Although nitrogen is considered to be the nutrient that most commonly limits production of natural and managed terrestrial ecosystems, I propose that phosphorus may regulate productivity in many continuously cultivated agroecosystems that do not receive applications of synthetic fertilizers. One way P may limit agroecosystem productivity is by controlling nitrogen fixation of legume crops, thus affecting nitrogen availability in the overall agroecosystem. I tested this hypothesis in two studies by examining the effect of phosphorus nutrition on nitrogen fixation of alfalfa in traditional Mexican agroecosystems. All farms used in the research relied on alfalfa as the primary nitrogen source for maize cultivation and other crops, and had minimal or no reliance on synthetic fertilizers.In one study, I used the natural abundance of¹⁵N to estimate nitrogen fixation in five alfalfa plots with soils representing a wide range of P fertility. I found a correlation of r = 0.85 between foliage P concentrations and nitrogen fixation in the alfalfa plots. Mean nitrogen fixation in alfalfa plots ranged between 232–555 kg ha^–1 yr^–1 as estimated by the¹⁵N-natural abundance method.In a second study, I sampled soils from alfalfa plots on traditional farms located in 5 different physiographic regions of Mexico. Half of each soil sample was augmented with phosphorus in a greenhouse experiment. I grew alfalfa on the fertilized and unfertilized soils from each site and then determined nitrogenase activity (acetylene reduction) of the Rhizobium on the plant roots. Nitrogenase activity increased in the alfalfa grown on all soils with added phosphorus, with two of the five differences being statistically significant at P < 0.01, 0 and one at P < 0.05. Foliage P concentrations and nitrogenase activity were 0 positively correlated (r = 0.81,P < 0.01).0     

Assessment of genetic variability for N2 fixation between and within provenances of Leucaena leucocephala and Acacia albida estimated by 15N labelling techniques

Nitrogen fixed in 13 provenances of Acacia albida and 11 isolines of Leucaena leucocephala inoculated with effective Rhizobium strains was measured by ¹⁵N techniques and the total N difference method. In the test soil, on the average, L. leucocephala derived about 65% of its total N from atmospheric N₂ fixation compared to about 20% by A. albida. Significant differences in the percentage of N derived from atmospheric N₂ (% Ndfa) occurred, between provenances or isolines within species. The % Ndfa ranged from 37 to 74% within L. leucocephala and from 6 to 37 within A. albida; (equivalent to 20–50 mg N plant^–1 and 4–37 mg N plant^–1 for the two species over three months, respectively) and was correlated with the nodule mass (r=0.91). The time course of N₂ fixation of three selected provenances (low, intermediate and good fixers) was followed at 12 weekly intervals over a 36 week period. The % Ndfa of all provenances and isolines increased with time; and except for one of the L. leucocephala provenances, % Ndfa was similar within species at the 36 weeks harvest. There was a significant correlation between % Ndfa and the amount of N₂ fixed (r=0.96). Significant interactions occurred between provenances and N treatments and often growth of uninoculated but N fertilized plants was less variable than for inoculated unfertilized plants.     

The effect of nitrogen fertilization on nitrogen use efficiency of irrigated and non-irrigated tobacco (Nicotiana tabacum L.).

Burley tobacco (Nicotiana tabacum L.) plants were grown in the field with or without irrigation and fertilized with 0, 120, 240 or 360 kg N ha^–1 over two growing seasons to assess nitrogen use under Mediterranean climate conditions. Kjeldahl-N and NO₃-N in leaves and stems and NO₃-N and NH₄-N in the soil at two depths (0–0.3 and 0.3–0.6 m) were determined. The effect of N fertilization on total N accumulated in the canopy biomass was markedly different between irrigated and non-irrigated plants. Under non-irrigated conditions N accumulated in the plant did not depend on the amount of N applied. In both years, the amount of N in irrigated plants increased in response to the amount of N applied, starting from 49 to 56 days after transplanting (DAT). The average amount of total N in the canopy of irrigated plants, measured across all sampling dates of both years, ranged from 30 kg ha^–1 of the unfertilized control to 88 kg ha^–1 of the 360 kg ha^–1 of N applied. The average amount of plant NO₃-N was 2.6 and 4.4 kg ha^–1 for non-irrigated and irrigated plots across all N treatments (means of 1996 and 1997). Nitrogen uptake rate (NUR) of non-irrigated plants was high between seedling establishment and the period of rapid stem elongation in 1996 (from 36 to 50 DAT) and until flowering in 1997 (from 42 to 71 DAT), but much less or negligible at later stages of plant development. Irrigation increased NUR dramatically in the second part of the growing season. Maximum NUR was estimated for plants receiving 240 or 360 kg N ha^–1 in both years. The year of study did not affect the recovery fraction (RF), physiological efficiency (PE) or agronomic efficiency (AE). Irrigation and N fertilization had significant effects on both RF and AE, but not on PE. Maximum values of RF were 45 and 22% for irrigated and non-irrigated treatments, respectively. In irrigated plots there was a negative relationship between RF and increasing N levels at all sampling dates.     

Growth response of Pinus radiata to multiple applications of nitrogen fertilizer and evaluation of the quantity of added nitrogen remaining in the forest system

Over a period of nine years, 922 kg ha^-1 of N was added in eight applications to a 16 year old P. radiata stand in a low rainfall area in north-east Tasmania, Australia. Fertilizing lifted current annual increment from 8.5 m³ha^-1 for the unfertilized plots, to 31 m³ha^-1. Increased growth was associated with improved health of the trees. Biomass measurements showed that there was a large increase in needle retention as well as needle mass on the fertilized trees. Concentration of N was also higher in fertilized trees. Fertilized plots contained 467 kg ha^-1 more N than unfertilized plots. This represented about 50 percent of the N applied. Most of this extra N was in the forest crop and in the surface 10 cm of the soil. There was no increase in organic carbon in the surface soil with the result that the C/N ratio was reduced from a very high 28 to 17.Despite the high growth rates attained in the N fertilized plots, the failure to increase soil organic matter and the loss from the site of much of the applied N indicated that long term improvement of growth of these low rainfall sites was unlikely. Growth would only be maintained with continued N fertilizer additions.     

Nitrogen cycling in a northern hardwood forest: Do species matter?

To investigate the influence of individual tree species on nitrogen (N) cycling in forests, we measured key characteristics of the N cycle in small single-species plots of five dominant tree species in the Catskill Mountains of New York State. The species studied were sugar maple (Acer saccharum), American beech (Fagus grandifolia), yellow birch (Betula alleghaniensis), eastern hemlock (Tsuga canadensis), and red oak (Quercus rubra). The five species varied markedly in N cycling characteristics. For example, hemlock plots consistently showed characteristics associated with "slow" N cycling, including low foliar and litter N, high soil C:N, low extractable N pools, low rates of potential net N mineralization and nitrification and low NO ₃ ^– amounts trapped in ion-exchange resin bags buried in the mineral soil. Sugar maple plots had the lowest soil C:N, and the highest levels of soil characteristics associated with NO ₃ ^– production and loss (nitrification, extractable NO ₃ ^– , and resin bag NO ₃ ^– ). In contrast, red oak plots had near-average net mineralization rates and soil C:N ratios, but very low values of the variables associated with NO ₃ ^– production and loss. Correlations between soil N transformations and litter concentrations of N, lignin, lignin:N ratio, or phenolic constituents were generally weak. The inverse correlation between net nitrification rate and soil C:N that has been reported in the literature was present in this data set only if red oak plots were excluded from the analysis. This study indicates that tree species can exert a strong control on N cycling in forest ecosystems that appears to be mediated through the quality of soil organic matter, but that standard measures of litter quality cannot explain the mechanism of control.     

Soil nitrogen mineralization in plantations ofJuglans nigra interplanted with actinorhizalElaeagnus umbellata orAlnus glutinosa

Nitrogen mineralization rates were estimated in 19-year-old interplantings of black walnut (Juglans nigra L.) with dinitrogen fixing autumn-olive (Elaeagnus umbellata Thunb.) or black alder (Alnus glutinosa L. Gaertn.) and in pure walnut plantings at two locations in Illinois USA. N mineralization rates were measured repeatedly over a one year period usingin situ incubations of soil cores in oxygen-permeable polyethylene bags at 0–10 and 10–20 cm soil depths, and also by burying mixed-bed ion-exchange resin in soil. Mineralization rates were highest in summer and in plots containing actinorhizal Elaeagnus and Alnus in contrast with pure walnut plots. Elaeagnus plots at one location yielded 236 kg of mineral N ha^–1 yr^–1 in the upper 20 cm of soil, a value higher than previously reported for temperate decidous forest soils in North America. The highest mean plot values for N mineralization in soil at a location were 185 kg ha^–1 yr^–1 for Alnus interplantings and 90 kg ha^–1 yr^–1 for pure walnut plots. Plots which had high N mineralization rates also had the largest walnut trees. Despite low pH (4.1 and 6.5) and low extractable P concentrations (1.4 and 0.7 mg kg^–1 dry mass) at the two locations, nitrification occurred in all plots throughout the growing season. NO ₃ ^– –N was the major form of mineralized N in soil in the actinorhizal interplantings, with NH ₄ ⁺ –N being the major form of mineral N in control plots. Walnut size was highly correlated with soil nitrogen mineralization, particularly soil NO ₃ ^– –N production in a plot.     

Soil Solution Chemistry and Element Fluxes in Three European Heathlands and Their Responses to Warming and Drought

Soil water chemistry and element budgets were studied at three northwestern European Calluna vulgaris heathland sites in Denmark (DK), The Netherlands (NL), and Wales (UK). Responses to experimental nighttime warming and early summer drought were followed during a two-year period. Soil solution chemistry measured below the organic soil layer and below the rooting zone and water fluxes estimated with hydrological models were combined to calculate element budgets. Remarkably high N leaching was observed at the NL heath with 18 and 6.4 kg N ha^–1 year^–1 of NO₃–N and NH₄–N leached from the control plots, respectively, indicating that this site is nitrogen saturated. Increased soil temperature of +0.5°C in the heated plots almost doubled the concentrations and losses of NO₃–N and DON at this site. Temperature also increased mobilization of N in the O horizon at the UK and DK heaths in the first year, but, because of high retention of N in the vegetation or mineral soil, there were no significant effects of warming on seepage water NO₃–N and NH₄–N. Retention of P was high at all three sites. In several cases, drought increased concentrations of elements momentarily, but element fluxes decreased because of a lower flux of water. Seepage water DOC and DON was highly significantly correlated at the UK site where losses of N were low, whereas losses of C and N were uncoupled at the NL site where atmospheric N input was greatest. Based on N budgets, calculations of the net change in the C sink or source strength in response to warming suggest no change or an increase in the C sink strength during these early years.     

Indices of plant N availability in an alpine grassland under elevated atmospheric CO2

The objective of this study was to estimate whether elevated atmospheric [CO₂] alters plant N availability in a native high-elevation grassland in the Swiss Alps using two integrative, relatively non-disruptive methods. Estimates based on seasonal net plant N uptake, and those based on the amounts of NH ₄ ⁺ -N plus NO ₃ ^– -N captured by ion exchange resin (IER) bags, did not differ in plots treated with ambient (355 L L^–1) and elevated (680 L L^–1) [CO₂] in either the second (1993) or third (1994) growing season under treatment with elevated [CO₂]. The results of this study suggest that the effects of rising atmospheric [CO₂] on plant N availability may be negligible in this grassland. The results also contrast the relatively large effects of elevated atmospheric [CO₂] (increases and decreases) reported for highly disturbed artificial systems.     

Segregation of habitat and prey in two sympatric carnivorous plant species,Drosera rotundifolia and Drosera intermedia

Summary Labelled nitrogen was used to evaluate the effects of intensive forest management on soil nitrogen transformations. The total release of N into inorganic forms (ammonium plus nitrate) was much greater than net N mineralization in all treatments. Immobilization of N by microbes was greatest in minimally-treated harvested plots, while the turnover of N within soil microbes was greatest in intensively-treated plots. Ammonium was immobilized 2.4–3.2 times more rapidly than nitrate in havested plots; nitrification in disturbed sites could thus increase the availability of N to regrowing vegetation.     

Utilisation of soil organic P by agroforestry and crop species in the field,western Kenya

A field experiment in western Kenya assessed whether the agroforestry species Tithonia diversifolia (Hemsley) A. Gray, Tephrosia vogelii Hook f., Crotalaria grahamiana Wight & Arn. and Sesbania sesban (L) Merill. had access to forms of soil P unavailable to maize, and the consequences of this for sustainable management of biomass transfer. The species were grown in rows at high planting density to ensure the soil under rows was thoroughly permeated by roots. Soil samples taken from beneath rows were compared to controls, which included a bulk soil monolith enclosed by iron sheets within the tithonia plot, continuous maize, and bare fallow plots. Three separate plant biomass samples and soil samples were taken at 6-month intervals, over a period of 18 months. The agroforestry species produced mainly leaf biomass in the first 6 months but stem growth dominated thereafter. Consequently, litterfall was greatest early in the experiment (0–6 months) and declined with continued growth. Soil pH increased by up to 1 unit (from pH 4.85) and available P increased by up to 38% (1 g P g^–1) in agroforestry plots where biomass was conserved on the field. In contrast, in plots where biomass was removed, P availability decreased by up to 15%. Coincident with the declines in litterfall, pH decreased by up to 0.26 pH units, plant available P decreased by between 0.27 and 0.72 g g^–1 and P_o concentration decreased by between 8 and 35 g g^–1 in the agroforestry plots. Declines in P_o were related to phosphatase activity (R²=0.65, P<0.05), which was greater under agroforestry species (0.40–0.50 nmol MUB s^–1 g^–1) than maize (0.28 nmol MUB s^–1 g^–1) or the bare fallow (0.25 nmol MUB s^–1 g^–1). Management of tithonia for biomass transfer, decreased available soil P by 0.70 g g^–1 and P_o by 22.82 g g^–1. In this study, tithonia acquired P_o that was unavailable to maize. However, it is apparent that continuous cutting and removal of biomass would lead to rapid depletion of P stored in organic forms.