Interactive effects of warming and increased nitrogen deposition on 15N tracer retention in a temperate old field: seasonal trends

The extent to which increased atmospheric nitrogen (N) deposition will drive changes in plant productivity and species composition over the next century will depend on how other influential global change factors, such as climate warming, affect the N retention of ecosystems. We examined the interactive effects of simulated climate warming and N deposition on the recoveries of ¹⁵N‐labeled ammonium and ¹⁵N‐labeled nitrate tracers added as a pulse to grass‐dominated, temperate old‐field plots at spring thaw. In addition to the year‐round warming treatment, a winter‐only warming treatment was applied to a set of plots to explore the contribution of this component of climate warming to the overall warming effect. By the end of the plant growing season, there was approximately twice as much ¹⁵N enrichment in the plant roots and bulk soil from ¹⁵NH₄⁺‐addition plots than from ¹⁵NO₃^?‐addition plots, but there were no effects of warming or N fertilization on ¹⁵N recovery. Over winter, approximately half of the excess ¹⁵N present in plant shoots was lost, which corresponded with large ¹⁵N losses from bulk soil in N fertilized plots and large ¹⁵N increases in bulk soil in nonfertilized plots. By the next spring, there was decreased ¹⁵N recovery in plants in response to N fertilization, which was largely offset by increases in plant ¹⁵N recovery in response to year‐round warming. However, ¹⁵N retention in bulk soil, where the major part of the ¹⁵N label was recovered, was approximately 40% higher in nonfertilized plots than in N fertilized plots. Overall, our results indicate that climate warming increases plant N sequestration in this system but this effect is overwhelmed by the overall effect of nitrogen deposition on ecosystem N losses.     

Bryophyte biomass and species richness on the Park Grass Experiment,Rothamsted, UK

The relationships between bryophyte biomass and species richness and soil pH, nutrient applications and vascular plant biomass and species richness were analyzed for the Park Grass Experiment (Rothamsted, UK). The study examined the abundance of bryophytes in relation to long-term fertilizer and lime application and to fertilizer treatments recently being ceased on some plots. The probability of bryophytes being present on a plot increased with increasing soil pH, and on plots at soil pH 3.3–4.5, the lowest values in this experiment, there were virtually no mosses present. Total bryophyte biomass decreased with increasing vascular plant biomass and vascular plant richness. Both bryophyte biomass and species richness showed a curvilinear response to soil pH. Bryophyte biomass was markedly increased on plots where nitrogen (N) fertilization had recently been ceased. The abundance of the common bryophyte species showed individualistic responses to treatments. N had a negative effect on the abundance of Brachythecium rutabulum. Increasing soil pH, and the application of phosphorus (P) and potassium (K) fertilizer together, had a positive effect on Eurhynchium praelongum. This species was also negatively affected by N, but tolerated larger amounts of it (100–150 kg ha^–1 N) than B. rutabulum. An ephemeral moss, Bryum subapiculatum, had a unimodal response to soil pH but showed no response to N, P, K or other explanatory variables.     

Loss of forb diversity in relation to nitrogen deposition in the UK: regional trends and potential controls

In this study we investigate the impact of nitrogen (N) deposition on the diversity of three different vegetation functional groups – forbs, grasses and mosses – using a field survey of acid grasslands across Great Britain. Our aim is to identify the vegetation types that are most vulnerable to enhanced N deposition, and to shed light on the mechanisms that may be driving N‐initiated species changes in the UK. Sixty‐eight randomly selected grasslands belonging to the UK National Vegetation Classification group U4 (Festuca ovina–Agrostis capillaris–Galium saxatile grassland) were studied along a gradient of atmospheric N deposition ranging from 6 to 36 kg N ha^?1 yr^?1. At each site, vegetation was surveyed and samples were taken from the topsoil and subsoil. Aboveground plant material was collected from three species: a forb, grass and moss. Both the species richness and cover of forbs declined strongly with increasing N deposition, from greater than eight species/20% cover per m² quadrat at low levels of N to fewer than two species/5% cover at the highest N deposition levels. Grasses showed a weak but significant decline in species richness, and a trend toward increasing cover with increasing N input. Mosses showed no trends in either species richness or cover. Most of the decline in plant species richness could be accounted for by the level of ammonium deposition. Soil KCl‐extractable ammonium concentration showed a significant positive correlation with N input, but there was no relationship between N deposition and extractable nitrate. In the soil O/A horizon, there was no relationship between N deposition and %N, and only a very weak positive relationship between the level of N deposition and the C : N ratio. Finally, in the vegetation, there was no relationship between N deposition and either shoot tissue N concentration or N : P ratio for any of the three reference species. Combining our regional survey with the results of published N‐addition experiments provides compelling evidence that there has been a significant decline in the species richness and cover of forbs across Great Britain, and that the primary cause is competition due to an increase in the cover of grasses in response to enhanced deposition of reactive N, primarily NH₄⁺.     

Effects of nutrient addition on vegetation and carbon cycling in an ombrotrophic bog

We measured net ecosystem CO₂ exchange (NEE), plant biomass and growth, species composition, peat microclimate, and litter decomposition in a fertilization experiment at Mer Bleue Bog, Ottawa, Ontario. The bog is located in the zone with the highest atmospheric nitrogen deposition for Canada, estimated at 0.8–1.2 g N m⁻² yr⁻¹ (wet deposition as NH₄ and NO₃). To establish the effect of nutrient addition on this ecosystem, we fertilized the bog with six treatments involving the application of 1.6–6 g N m⁻² yr⁻¹ (as NH₄NO₃), with and without P and K, in triplicate 3 m × 3 m plots. The initial 5–6 years have shown a loss of first Sphagnum, then Polytrichum mosses, and an increase in vascular plant biomass and leaf area index. Analyses of NEE, measured in situ with climate‐controlled chambers, indicate that contrary to expectations, the treatments with the highest levels of nutrient addition showed lower rates of maximum NEE and gross photosynthesis, but little change in ecosystem respiration after 5 years. Although shrub biomass and leaf area increased in the high nutrient plots, loss of moss photosynthesis owing to nutrient toxicity, increased vascular plant shading and greater litter accumulation contributed to the lower levels of CO₂ uptake. Our study highlights the importance of long‐term experiments as we did not observe lower NEE until the fifth year of the experiment. However, this may be a transient response as the treatment plots continue to change. Higher levels of nutrients may cause changes in plant composition and productivity and decrease the ability of peatlands to sequester CO₂ from the atmosphere.     

Effects of Increased Nitrogen Deposition on the Distribution of 15N-labeled Nitrogen between Sphagnum and Vascular Plants

To elucidate the sensitivity of bog ecosystems to high levels of nitrogen (N) deposition, we investigated the fate of ¹⁵N-labeled N deposition in bog vegetation in the Netherlands, both at ambient and increased N deposition. We doubled N deposition by adding 5 g N m^?2 y^?1 as dissolved NH₄NO₃ during three growing seasons to large peat monoliths (1.1 m diameter) with intact bog vegetation kept in large outdoor containers. A small amount of ¹⁵N tracer was applied at the start of the second growing season, and its distribution among Sphagnum, vascular plant species, and peat was determined at the end of the third growing season. The ¹⁵N tracer was also applied to additional plots at the untreated field site to check for initial distribution. One week after addition, 79% of the total amount of ¹⁵N retrieved was found in the living Sphagnum layer and less than 10% had been captured by vascular plants. Fifteen months later, 63% of the total amount of ¹⁵N retrieved was still present in the living Sphagnum layer at ambient N deposition. Increased N deposition significantly reduced the proportion of ¹⁵N in Sphagnum and increased the amount of ¹⁵N in vascular plants. Deep-rooting vascular plant species were significantly more ¹⁵N enriched, suggesting that at higher atmospheric inputs N penetrates deeper into the peat. Our results provide the first direct experimental evidence for that which has often been suggested: Increased atmospheric N deposition will lead to increased N availability for vascular plants in ombrotrophic mires.     

Inertia in an ombrotrophic bog ecosystem in response to 9 years' realistic perturbation by wet deposition of nitrogen,separated by form

Wet deposition of nitrogen (N) occurs in oxidized (nitrate) and reduced (ammonium) forms. Whether one form drives vegetation change more than the other is widely debated, as field evidence has been lacking. We are manipulating N form in wet deposition to an ombrotrophic bog, Whim (Scottish Borders), and here report nine years of results. Ammonium and nitrate were provided in rainwater spray as NH₄Cl or NaNO₃ at 8, 24 or 56 kg N ha^?1 yr^?1, plus a rainwater only control, via an automated system coupled to site meteorology. Detrimental N effects were observed in sensitive nonvascular plant species, with higher cumulative N loads leading to more damage at lower annual doses. Cover responses to N addition, both in relation to form and dose, were species specific and mostly dependent on N dose. Some species were generally indifferent to N form and dose, while others were dose sensitive. Calluna vulgaris showed a preference for higher N doses as ammonium N and Hypnum jutlandicum for nitrate N. However, after 9 years, the magnitude of change from wet deposited N on overall species cover is small, indicating only a slow decline in key species. Nitrogen treatment effects on soil N availability were likewise small and rarely correlated with species cover. Ammonium caused most N accumulation and damage to sensitive species at lower N loads, but toxic effects also occurred with nitrate. However, because different species respond differently to N form, setting of ecosystem level critical loads by N form is challenging. We recommend implementing the lowest value of the critical load range where communities include sensitive nonvascular plants and where ammonium dominates wet deposition chemistry. In the context of parallel assessment at the same site, N treatments for wet deposition showed overall much smaller effects than corresponding inputs of dry deposition as ammonia.     

Does nitrogen availability control rates of litter decomposition in forests?

The effects of increased exogenous N availability on rates of litter decomposition were assessed in several field fertilization trials. In a jack pine (Pinus banksiana Lamb.) forest, needle litter decomposed at the same rate in control plots and in plots fertilized with urea and ammonium nitrate (1350 kg N ha^-1) with or without P and K. Mixed needle litter of western hemlock (Tsuga heterophylla (Raf.) Sarg.), western red cedar (Thuja plicata Donn) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) incubated in plots recently amended with sewage sludge (500 kg N ha^-1) lost less weight during 3 years than did litter in control plots. Forest floor material also decomposed more slowly in plots amended with sewage sludge. Paper birch (Betula papyrifera Marsh.) leaf litter placed on sewage sludge (1000 kg N ha^-1), pulp sludge, or sewage-pulp sludge mixtures decomposed at the same rate as leaf litter in control plots. These experiments demonstrate little effect of exogenous N availability on rates of litter decomposition.The influence of endogenous N availability on rates of litter decomposition was examined in a microcosm experiment. Lodgepole pine (Pinus contorta var. latifolia Engelm.) needle litter collected from N-fertilized trees (525 kg N ha^-1 in ammonium nitrate) were 5 times richer in N than needles from control trees (1.56% N versus 0.33% N in control trees), but decomposed at the same rate. Green needles from fertilized trees contained twice as much N as needles from control trees (1.91% N versus 0.88% N), but decomposed at the same rate. These experiments suggest that N availability alone, either exogenous or endogenous, does not control rates of litter decomposition. Increased N availability, through fertilization or deposition, in the absence of changes in vegetation composition, will not alter rates of litter decomposition in forests.     

The high-latitude terrestrial carbon sink: a model analysis

A dynamic, global vegetation model, hybrid v4.1 ( Friend et al. 1997 ), was driven by transient climate output from the UK Hadley Centre GCM (HadCM2) with the IS92a scenario of increasing atmospheric CO₂ equivalent, sulphate aerosols and predicted patterns of atmospheric N deposition. Changes in areas of vegetation types and carbon storage in biomass and soils were predicted for areas north of 50°N from 1860 to 2100. Hybrid is a combined biogeochemical, biophysical and biogeographical model of natural, potential ecosystems. The effect of periodic boreal forest fires was assessed by adding a simple stochastic fire model. Hybrid represents plant physiological and soil processes regulating the carbon, water and N cycles and competition between individuals of parameterized generalized plant types. The latter were combined to represent tundra, temperate grassland, temperate/mixed forest and coniferous forest. The model simulated the current areas and estimated carbon stocks in the four vegetation types. It was predicted that land areas above 50°N (about 23% of the vegetated global land area) are currently accumulating about 0.4 PgC y^?1 (about 30% of the estimated global terrestrial sink) and that this sink could grow to 0.8–1.0 PgC y^?1 by the second half of the next century and persist undiminished until 2100. This sink was due mainly to an increase in forest productivity and biomass in response to increasing atmospheric CO₂, temperature and N deposition, and includes an estimate of the effect of boreal forest fire, which was estimated to diminish the sink approximately by the amount of carbon emitted to the atmosphere during fires. Averaged over the region, N deposition contributed about 18% to the sink by the 2080 s. As expected, climate change (temperature, precipitation, solar radiation and saturation pressure deficit) and N deposition without increasing atmospheric CO₂ produced a carbon source. Forest areas expanded both south and north, halving the current tundra area by 2100. This expansion contributed about 30% to the sink by the 2090 s. Tundra areas which were not invaded by forest fluctuated from sink to source. It was concluded that a high latitude carbon sink exists at present and, even assuming little effect of N deposition, no forest expansion and continued boreal forest fires, the sink is likely to persist at its current level for a century.     

Latent soil effects of grazing and ammonium deposition on Deschampsia flexuosa tillers inserted and grown in heather moorland soil

Upland heather moorlands support a range of increasingly rare and threatened biota, making them a priority habitat for conservation and restoration. Over-grazing and nitrogen deposition are two of the most important threats to maintaining these heather moorlands, yet there remains a paucity of research into their combined long-term effects. During the summer of 2008, we established an experiment within an existing research site at Pwllpeiran, mid-Wales. The original site, established in 1996, investigated long-term grazing and N deposition treatments (ammonium and nitrate) on heather moorland. Previous findings from the site, following a decade of treatment, suggested no significant, detectable impacts of treatments on soil C:N ratio and the foliar nitrogen of vegetation.The aim of our study was to investigate short- (2008) and long-term (1996–2008) N deposition treatment impacts, in combination with past grazing (1990–2007), on soil nutrient bioavailability. Soil cores were harvested and aboveground vegetation removed. Tillers of the grass Deschampsia flexuosa were planted into these cores which were then reciprocally transplanted back into the field experiment. The N deposition treatment was continued but grazing was excluded. D. flexuosa biomass changes were next assessed and leaf chemistry investigated using the metabolic fingerprinting method Fourier-transform infrared spectroscopy (FTIR) following three months of growth in the field (May–August 2008).Grazing treatment (on its own) had significant negative impacts on aboveground biomass and significant changes in plant chemistry were also revealed through the metabolic fingerprinting method Fourier transform infrared spectroscopy (FTIR). Short-term N deposition treatments during 2008 had no impacts on D. flexuosa growth or chemistry. There were also no detectable latent effects of long-term nitrate treatments on either growth or chemistry of D. flexuosa. However, plants grown in plots that had received long-term treatments of ammonium (NH₄⁺) had significantly lower poly-phenolic contents (revealed by FTIR) than plants grown in either nitrate (NO₃⁻) or control plots, suggesting detectable latent effects of N application in its reduced form. Further work needs to be undertaken to assess the relevance of residual soil nitrogen pools post N deposition and grazing.     

Long-term effects of PO4 fertilization on the distribution of bryophytes in an arctic river

1. Each year since 1983, H₃PO₄ has been added continuously during the ice-free season to a P-limited tundra river (Kuparuk River, North Slope, Alaska). Effects on epilithic metabolism, invertebrate community structure and fish production developed quickly. 2. In 1990, 7 years after fertilization began, we noted extensive coverage by bryophytes within the fertilized reach of the river, where very little had been noted before. Bryophyte biomass from a limited set of quadrats taken in 1990 and 1991 yielded 17 ± 9 (SE) g dry mass m^?2 in control reaches and 322 ± 96 g dry mass m^?2 in fertilized reaches. 3. An initial survey of macroalgal and bryophyte cover in 1991 suggested that the moss Schistidium (Grimmia) agassizii was distributed in both control and fertilized reaches of the river. No clear difference in coverage by this species was found in either reach. 4. In contrast, two species of Hygrohypnum (H. alpestre and H. ochraceum) were found almost exclusively in the fertilized reach. An extensive point transect survey done in 1992, above, within and below the fertilized reach, indicated that increased cover and biomass of Hygrohypnum spp. were confined to the fertilized reach of the river. Detrended correspondence analysis clearly separated the macrophyte and macroalgal communities in the fertilized reach from those in the control and downstream reaches. 5. A fourth bryophyte species (Fontinalis neomexicana) also occurred almost exclusively in the fertilized reach, but was much less abundant than the Hygrohypnum species. 6. Analysis of total N and P in the tissues of the Hygrohypnum spp., and estimates of average coverage (～15%) and biomass (～150g dry weight m^?2) over an 8km fertilized reach, suggest that these species alone may have removed two-thirds of the P added in the fertilizer experiment. The bryophyte community in this river is likely to be the dominant sink for P in the fertilized reach.     

Potassium limits potential growth of bog vegetation under elevated atmospheric CO2 and N deposition

The free air carbon dioxide enrichment (FACE) and N deposition experiments on four ombrotrophic bogs in Finland, Sweden, the Netherlands and Switzerland, revealed that after three years of treatment: (1) elevated atmospheric CO₂ concentration had no significant effect on the biomass growth of Sphagnum and vascular species; and (2) increased N deposition reduced Sphagnum growth, because it increased the cover of vascular plants and the tall moss Polytrichum strictum, while vascular plant biomass growth was not affected. This paper focuses on water chemistry, plant nutrient content, and litter decomposition rates. Potassium limitation, or low supply of K and P, may have prevented a significant increase of Sphagnum growth under elevated CO₂ and N deposition. Vascular plant growth under elevated CO₂ and N deposition was also limited by K, or by K in combination with P or N (N in CO₂ experiment). Elevated CO₂ and N deposition had no effect on decomposition rates of Sphagnum and vascular plant litter. Aside from a possible effect of N deposition on light competition between species, we expect that elevated atmospheric CO₂ and N deposition concentrations will not affect Sphagnum and vascular plant growth in bogs of north‐west Europe due to K‐, or K in combination with N‐ or P‐, limited growth. For the same reason we expect no effect of elevated CO₂ and N deposition on litter decomposition. Net primary production of raised ombrotrophic bogs that are at or close to steady state, is regulated by input of nutrients through atmospheric deposition. Therefore, we hypothesize that the expected increase of plant growth under elevated CO₂ and N deposition is diminished by current levels of K (and to some extent P and N) in atmospheric deposition.     

NH4 : NO3 nutrition influence on biomass productivity and root respiration of poplar and willow clones

Nitrogen fertilization often improves the yield of intensively managed, short‐rotation coppices. However, information of N nutrition form on the growth of common species and clones used for biomass production is limited. Thus, this study aims at evaluating N form effects on the growth of two Salicaceae clones. Cuttings of the poplar clone Max 4 (Populus maximovizcii × P. nigra) and the willow clone Inger (Salix triandra × S. viminialis) were fertilized in a pot experiment with four ratios of nitrate (NO₃^?) to ammonium (50%, 62.5%, 75% and 87.5% NO₃^? balanced with ammonium (NH₄⁺) to constant total N) for one growing season and under stable soil pH. Plants were harvested for analysis of biomass and morphology of leaves, stem and roots. Respiration of fine and coarse roots (RR) was determined and related to biomass growth. Salix cv. Inger accumulated more total dry matter than Populus cv. Max 4. In both Salicaceae clones, the total biomass was significantly influenced by the nitrate ratio and greatest in plants fertilized with 50% NO₃^? of the total N supply. Both clones possess a different leaf and root morphology, but no significant influence of the NO₃^? ratio on the morphology was found. Fine RR rates differed significantly between clones, with significantly greater fine RR in Max 4; 87.5% NO₃^? fertilization increased the fine RR. Fine RR and total accumulated plant biomass were closely related. Our study is the first to show the tremendous influence of fine root respiration, especially including the carbon‐intensive reduction of NO₃^? to NH₄⁺, on the aboveground growth of Salicaceae clones. Ways to improve yield in SRC are thus to lower the assimilate consumption by fine roots and to match fertilization regimes to the used clones or vice versa.     

Changes in forest understorey vegetation in Norway related to long‐term soil acidification and climatic change

Question: Does the understorey vegetation of Norwegian boreal forests change in relation to broad‐scale, long‐term changes? Location: Norway. Methods: Permanently marked 1‐m² vegetation plots from 17 monitoring reference areas in forests dominated by Picea abies (11 areas, 620 plots) and Betula spp. (six areas, 300 plots) were analysed twice, at the start in 1988–1997 and 5 yr later (1993–2002). Species subplot frequency data were analysed separately for each area by univariate and multivariate statistical methods; 5‐yr changes in single species abundances, species number per plot and species composition were tested. Results: Two distinct patterns of change were found: 1. Abundance of several vascular plant species decreased in SE Norwegian Picea forests, most noticeably of species with a preference for richer soils, such as Oxalis acetosella. 2. Abundance of many bryophyte species as well as bryophyte species number per plot increased in forests of both types over most of Norway. Conclusions: The pattern of vascular plant changes is probably a time‐delayed response of long‐lived, mainly clonal, populations to acidified soils resulting from deposition of long‐distance airborne pollutants. The pattern bryophyte changes, with reference to the close link between climatic conditions for growth and abundance changes for Hylocomium splendens established in previous demographic studies, is related to climatic conditions favourable for bryophyte growth. We conclude that many forest understorey plants are sensitive indicators of environmental change, and that the concept used for intensive monitoring of Norwegian forests enables early detection of changes in vegetation brought about by broad‐scale, regional, impact factors.     

桧叶白发藓对不同氮源胁迫的形态和生理响应

桧叶白发藓(Leucobryum juniperoideum)在我国东南部常见,被认为是一种理想的、适用于庭院栽培的苔藓植物,而氮是植物必需的矿质元素,但过量摄入会对其造成伤害,近年来氮沉降水平的提高对苔藓植物的多样性造成了严重影响。该研究为揭示氮沉降加剧对桧叶白发藓的影响,以经6个月断茎培养的桧叶白发藓配子体为材料,用Ca(NO3)2、NH4HCO3和NH4NO3代表三种氮源,设置2、4、8、16gN·m-2 4个水平,以喷洒去离子水为对照,进行不同氮源的胁迫试验。结果表明:氮处理浓度的增加引起组织氮含量的显著提高,增加幅度分别为69.1%、25.7%和43.1%;同时引起植株坏死率显著上升,增加幅度分别为16.5%、12.5%和13.9%。三种氮源处理对株高和净重的影响有显著差异,低浓度的铵态氮(4gN·m-2)引起株高和净重的显著增加,而硝态氮和混合态氮处理差异不显著;加氮浓度的进一步提高,引起株高和净重的减低,硝态氮处理的减低幅度最大,铵态氮的降低幅度最小。三种氮源处理均引起叶绿素含量先上升后下降,但同一水平铵态氮处理的叶绿素含量要高于其它两种氮处理,而且引起叶绿素含量下降的处理浓度要高于其它两种氮源;三种氮源均引起SOD活性显著增加、可溶性蛋白和脯氨酸含量先升后降,但不同氮源间生理指标的变化不同步。这说明桧叶白发藓对硝态氮胁迫的响应比铵态氮敏感,硝态氮的增加对桧叶白发藓造成严重危害,而少量的铵态氮(4gN·m-2)则能促进桧叶白发藓的生长。研究结果可作为桧叶白发藓繁殖与生产的氮源。     

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.     

Impacts of Soil Nitrogen and Carbon Additions on Forest Understory Communities with a Long Nitrogen Deposition History

Rates of nitrogen (N) deposition have been historically high throughout much of the northeastern United States; thus, understanding the legacy of these high N loads is important for maintaining forest productivity and resilience. Though many studies have documented plant invasions due to N deposition and associated impacts on ecosystems, less is known about whether invasive plants will continue to increase in dominance with further shifting nutrient regimes. Using soil N and carbon additions, we examined the impact of both increasing and decreasing soil N on native and invasive understory plant dynamics over 4 years in a northeastern deciduous forest with a long history of N deposition. Despite applying large quantities of N, we found no difference in soil nitrate (NO₃) or ammonium (NH₄ ⁺) pools in N addition plots over the course of the study. Indicative of the potential N saturation in these forest soils, resin-available NO₃ ^? and NH₄ ⁺ showed evidence that the added N was rapidly moving out of the soil in N addition plots. Accordingly, we also found that adding N to soil altered neither invasive nor native plant abundance, though adding N temporally increased invasive plant richness. Carbon additions decreased soil N availability seasonally, but did not alter the total percent cover of invasive or native plants. Rather than being suppressed by excess N availability, native plant species in this ecosystem are primarily inhibited by the invasive species, which now dominate this site. In conclusion, understory plant communities in this potentially N-saturated ecosystem may be buffered to future alterations in N availability.     

Changes in species composition of European acid grasslands observed along a gradient of nitrogen deposition

Question: Which environmental variables affect floristic species composition of acid grasslands in the Atlantic biogeographic region of Europe along a gradient of atmospheric N deposition? Location: Transect across the Atlantic biogeographic region of Europe including Ireland, Great Britain, Isle of Man, France, Belgium, The Netherlands, Germany, Norway, Denmark and Sweden. Materials and Methods: In 153 acid grasslands we assessed plant and bryophyte species composition, soil chemistry (pH, base cations, metals, nitrate and ammonium concentrations, total C and N, and Olsen plant available phosphorus), climatic variables, N deposition and S deposition. Ordination and variation partitioning were used to determine the relative importance of different drivers on the species composition of the studied grasslands. Results: Climate, soil and deposition variables explained 24% of the total variation in species composition. Variance partitioning showed that soil variables explained the most variation in the data set and that climate and geographic variables accounted for slightly less variation. Deposition variables (N and S deposition) explained 9.8% of the variation in the ordination. Species positively associated with N deposition included Holcus mollis and Leontodon hispidus. Species negatively associated with N deposition included Agrostis curtisii, Leontodon autumnalis, Campanula rotundifolia and Hylocomium splendens. Conclusion: Although secondary to climate gradients and soil biogeochemistry, and not as strong as for species richness, the impact of N and S deposition on species composition can be detected in acid grasslands, influencing community composition both directly and indirectly, presumably through soil‐mediated effects.     

Plant responses to nutrient addition and predictive ability of vegetation N:P ratio in an austral fen

1. Previous studies of the N:P ratio in wetland plants have been carried out in northern hemisphere wetlands where atmospheric nitrogen deposition is higher. There is little research on foliar N:P ratio as a potential indicator of nutrient limitation in vegetation communities in southern hemisphere wetlands. This study aimed to redress this knowledge gap and answer the following questions: how well does the plant tissue nitrogen to phosphorus (N:P) ratio predict wetland plant community nutrient limitation, as indicated by vegetation standing stocks and below-ground biomass, in southern hemisphere fens? Secondly, what are the impacts of realistic upper levels of farm nutrient run-off on natural montane fen vegetation?
2. Low (35 kg ha⁻¹ year⁻¹) and high (70 kg ha⁻¹ year⁻¹) levels of nitrate-N or ammonium-N with and without P (20 kg ha⁻¹ year⁻¹) were added to 81 vegetation plots over a period of 2.75 years. Species composition, plant nutrient status, and above-ground live vegetation standing stocks were assessed after 3 years, and below-ground biomass after 2 years.
3. Plant tissue analysis suggested the community was N limited or N and P co-limited; we found greater standing stocks of vegetation in plots treated with 70 kg ha⁻¹ year⁻¹ ammonium-N, indicating N limitation. No difference between other treatments was found in above-ground standing stocks or below-ground biomass. Plant species cover increased in both high N treatments, consistent with N limitation. These changes in plant species cover were accompanied by significant decreases in species richness in both high N treatments. Native species dominated the vegetation and this was unaffected by nutrient addition (90% cover).
4. This is one of the first studies to test and find support for the N:P ratio in southern hemisphere wetlands. Observed declines in species richness after N fertilisation in an N-limited fen suggests increased N may pose risks to austral wetlands. Responses by plant communities (changes in composition, biomass) to lower levels of nutrient addition may require longer periods of fertilisation to be apparent in slow growing ecosystems.

     

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.     

氮素和水分添加对贝加尔针茅草原植物多样性及生物量的影响

为了解草原植物群落物种多样性和植物地上生物量对氮沉降增加和降水变化的响应,在内蒙古贝加尔针茅(Stipa baicalensis)草原,分别设置对照(N0)、1.5 g/m2(N15)、3.0 g/m2(N30)、5.0 g/m2(N50)、10.0 g/m2(N100)、15.0 g/m2(N150)、20.0g/m2(N200)和30.0 g/m2(N300)(不包括大气沉降的氮量)8个氮素(NH4NO3)添加梯度和模拟夏季增加降水100mm的水分添加交互试验,研究氮素和水分添加对草原群落植物物种多样性和几种常见植物地上生物量的影响。结果表明:(1)氮素和水分的添加降低了草原群落植物物种多样性,且氮素和水分有显著的互作效应。在水分添加的条件下,随着施氮水平的增加,群落植物物种多样性减小;在无水分添加的条件下,随着施氮水平的增加,群落植物物种多样性呈先增加后减小的"单峰"变化趋势。(2)不同植物对氮素和水分添加的响应不同,随着施氮水平的增加,羊草地上生物量显著增加;贝加尔针茅、羽茅、糙隐子草、寸草苔和冷蒿先增加后减少,呈单峰曲线;星毛委陵菜、牧马豆、扁蓄豆和线叶菊地上生物量则逐渐减少。而且氮素和水分对贝加尔针茅、羽茅、扁蓄豆地上生物量有显著的交互作用。