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.     

Lignin signatures show the effects of changes in heather and bracken cover on the composition of organic matter in a moorland soil profile

Bracken (Pteridium aquilinum) is aggressively displacing heather (Calluna vulgaris) on many moorlands in Britain. We investigated the use of lignin derivatives to identify the distribution of soil organic matter (SOM) derived from bracken in moorland soil profiles formed under heather. Phenylpropanoids extracted from recently senesced litters, roots and SOM, using alkaline CuO oxidation, showed distinct signatures for bracken and heather, with vanillyl moieties dominating bracken litter extracts and vanillyl and syringyl dominating heather litter extracts. Ratios of vanillyl and syringyl concentrations characterised the SOM derived from heather and bracken better than the concentrations of the individual moieties. The analysis showed up to a depth of 5 cm under pure bracken cover, and at the interface between heather and bracken, the SOM was largely derived from bracken litter but below that depth SOM was apparently derived from heather. The use of these methods to identify the plant origin of SOM not only enables understanding the effects of changing vegetation cover on organic matter dynamics in moorland soils but could also facilitate management techniques in moorland/heathland restoration which involve the removal of comparatively nutrient-rich SOM derived from bracken. Received: 30 November 1997 / Accepted: 11 April 1998     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 2. The rhizome

The rhizome system of mature bracken (Pteridium aquilinum (L.) Kuhn) contains large reserves of both biomass (mean 8.63 kg m?² fr. wt) and buds (mean 565 m^-2) which are largely responsible for both its persistence and its often rapid rates of vegetative encroachment. Within areas such as the North York Moors the spread of bracken into areas previously dominated by heather and grass is considered undesirable because of reduced land value in terms of both agriculture and ecological diversity. In this paper we describe the morphology of bracken rhizome within a mature bracken stand, and at advancing and stationary stand margins where bracken-heather interfaces occur. Stationary margins, i.e. those where bracken is not encroaching into heather at a significant rate, often have morphological characteristics intermediate to those of a mature stand and an advancing margin. In the mature stand rhizome biomass is dominated by carbohydrate-storing long shoots which comprise 63% of the total fresh weight, whilst the majority of rhizome buds (89% of all active and 86% of all dormant buds) are found on frond-bearing short shoots. At the margins of a bracken stand the proportion of rhizome which is composed of long shoots is even greater, and that of short shoots small relative to that in the mature stand. More transitional shoots are also found at the stand margins. Hence close to the margin a greater proportion of fronds is found on transitional rhizome than is the case in more mature parts of the stand. The majority of buds on all types of rhizome are in a dormant state. The proportion of buds which are active is, however, greater on long and transitional shoots than on short shoots. Hence, a larger proportion of buds are active close to the margin where the rhizome is composed less of short shoots than is the case further into a mature stand. The differences in the morphology of bracken in a mature stand and at the stand margins which are identified here support the idea of controlling bracken at stand margins in preference to the spraying of large areas of dense, mature bracken. Morphological differences include an increased proportion of active buds, greater frequency of fronds per unit rhizome biomass, reduced biomass reserves. Improved conditions for the re-invasion and re-establishment of alternative vegetation are also available at stand margins in comparison with the centre of a dense bracken stand.     

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.     

模拟氮沉降对华西雨屏区撑绿 杂交竹凋落物分解的影响

从2008年1月至2010年1月,对华西雨屏区撑绿杂交竹(Bambusa pervariabilis × Dendrocala mopsi)人工林进行了模拟氮沉降试验,氮沉降水平分别为对照(CK, 0 g · m^-2 · a^-1)、低氮(5 g · m^-2 · a^-1)、中氮(15 g · hm^-2 · a^-1)和高氮(30 g · m^-2 · a^-1)。利用凋落袋法对杂交竹凋落叶和凋落箨进行原位分解试验,并在每月下旬定量地对各处理施氮(NH₄NO₃)。结果表明,自然状态下杂交竹凋落叶和凋落箨分解95%所需时间分别为2.9,1.5 a;氮沉降显著抑制了凋落叶的分解,在分解后期3个氮沉降处理凋落叶无灰分质量残留率均显著大于对照,氮沉降对凋落箨分解无明显影响;氮沉降显著抑制了凋落叶中木质素和纤维素的分解。杂交竹凋落叶在分解后期质量损失缓慢,处于较稳定状态,氮沉降的增加使得凋落物的残留率稳定在一个更高的水平,表明氮沉降的增加可能会使更多的凋落物残体和稳定有机质留存于杂交竹林土壤中,从而增加杂交竹林土壤碳贮存。     

The morphology of bracken (Pteridium aquilinum (L.) Kuhn) in the North York Moors—a comparison of the mature stand and the interface with heather (Calluna vulgaris (L.) Hull) 1. The fronds

The encroachment of bracken (Pteridium aquilinum (L.) Kuhn) into areas previously dominated by heather represents a threat to the ecology, agricultural economy and landscape value of many UK upland areas, including the moorland of the North York Moors National Park. The morphology of bracken, within a mature stand and at several bracken-heather interfaces, has been studied at a number of sites within the National Park. Differences have been found in the frond growth of bracken in a mature stand, at stationary stand margins, and at advancing stand margins where bracken is encroaching into heather. Frequency of fronds present on bracken rhizome growing at a stationary stand margin close to the interface with heather (1–2 m behind the boundary) are approximately the same as those found within a mature stand. At advancing margins (again 1–2 m behind the boundary), maximum frond densities were often found to exceed those present in either a mature stand or at a stationary margin. Frond numbers decline rapidly at the stand margins as distance from the stand increases. This is especially true where the front is stationary and bracken is not encroaching into heather at a significant rate. Maximum frond heights in a mature stand consistently exceed those at stand margins (even 1–2 m into the stand) and are greater at stationary margins than at advancing margins. Outlying fronds at the edges of bracken stands are generally present in greater numbers, and further into the area dominated by heather, where the margin is advancing. Heights of outlying fronds fall as distance from the bracken stand increases, as does stipe length. Fronds at the edges of bracken stands emerge each spring before those further into the stand and are therefore particularly vulnerable to frost damage. Outlying fronds are not, however, the first to emerge. Early emerging fronds reach their maximum height and eventually become senescent before later emerging fronds. Whilst most fronds emerge before the end of June a few fronds continue to emerge throughout the summer. Frond densities close to the edges of bracken stands (1–2 m into the stand) are comparable to those in a mature stand. At advancing stand margins frond densities generally exceed those in a mature stand, suggesting that a large number of potential entry points for foliage-applied herbicides are available for bracken control at the stand margins. The ratio of potential uptake points to biomass of rhizome is also greatest at the edges of the stand, and the canopy 1–2 m into the stand is usually almost completely closed. It is possible therefore, the efficacy of herbicides could be improved by the use of small scale applications, using tractors or hand-held sprayers, close to the margins of bracken stands.     

Profile distribution of organic carbon,iron, aluminium and manganese in soils under bracken and heather

Summary The vertical distribution of extractable iron, aluminium, and manganese in soil profiles on a heather-bracken transect indicates a less advanced stage of leaching under the bracken than under the heather. Evidence that the bracken is advancing into the heather is presented, and implies that the leaching process, like the accumulation of a mor humus layer, is reversed by the presence of the bracken.     

Response of macrophyte litter decomposition in global blue carbon ecosystems to climate change

Blue carbon ecosystems (BCEs) are important nature-based solutions for climate change-mitigation. However, current debates question the reliability and contribution of BCEs under future climatic-scenarios. The answer to this question depends on ecosystem processes driving carbon-sequestration and -storage, such as primary production and decomposition, and their future rates. We performed a global meta-analysis on litter decomposition rate constants (k) in BCEs and predicted changes in carbon release from 309 studies. The relationships between k and climatic factors were examined by extracting remote-sensing data on air temperature, sea-surface temperature, and precipitation aligning to the decomposition time of each experiment. We constructed global numerical models of litter decomposition to forecast k and carbon release under different scenarios. The current k averages at 27 ± 3 × 10⁻² day⁻¹ for macroalgae were higher than for seagrasses (1.7 ± 0.2 × 10⁻² day⁻¹), mangroves (1.6 ± 0.1 × 10⁻² day⁻¹) and tidal marshes (5.9 ± 0.5 × 10⁻³ day⁻¹). Macrophyte k increased with both air temperature and precipitation in intertidal BCEs and with sea surface temperature for subtidal seagrasses. Above a temperature threshold for vascular plant litter at ~25°C and ~20°C for macroalgae, k drastically increased with increasing temperature. However, the direct effect of high temperatures on k are obscured by other factors in field experiments compared with laboratory experiments. We defined “fundamental” and “realized” temperature response to explain this effect. Based on relationships for realized temperature response, we predict that proportions of decomposed litter will increase by 0.9%–5% and 4.7%–28.8% by 2100 under low- (2°C) and high-warming conditions (4°C) compared to 2020, respectively. Net litter carbon sinks in BCEs will increase due to higher increase in litter C production than in decomposition by 2100 compared to 2020 under RCP 8.5. We highlight that BCEs will play an increasingly important role in future climate change-mitigation. Our findings can be leveraged for blue carbon accounting under future climate change scenarios.     

Effects of temperature elevation on the population dynamics of the upland heather psyllid Strophingia ericae (Curtis) (Homoptera: Psylloidea)

1 Cloches permeable to gas and water were used to simulate the effect of climate warming on upland populations of the heather psyllid Strophingia ericae at Moor House National Nature Reserve in the North Pennines, UK.
2 The cloches produced an average warming of about 1 °C in the heather canopy over a period of one year.
3 The density of S. ericae increased markedly in cloches within a few months of erection.
4 Species composition and numbers of potential predators were similar inside cloches and in control plots.
5 In the two year life-cycle of S. ericae , the warming effect advanced the phenology from the second to third instar in the first winter, but in the second winter, fifth instar nymphs did not moult prematurely to adult.
6 The density of S. ericae was higher on Calluna vulgaris at its boundary with Juncus squarrosus than in the pure C. vulgaris sward (in both cloched and control plots).     

模拟氮沉降对两种竹林不同凋落物组分分解过程养分释放的影响

利用原位分解袋法研究了华西雨屏区苦竹(Pleioblastus amarus)和撑绿杂交竹(Bambusa pervariabilis × Dendrocala mopsi)人工林几种凋落物组分在模拟氮沉降下分解过程中养分释放状态,试验周期为2 a。氮沉降水平分别为对照(CK, 0 g · m^-2 · a^-1)、低氮(5 g · m^-2 · a^-1)、中氮(15 g · m^-2 · a^-1)和高氮(30 g · m^-2 · a^-1),每月下旬定量地对各处理施氮(NH₄NO₃)。结果表明,苦竹林和杂交竹林凋落物主要由凋落叶、凋落箨和凋落枝组成,其中凋落叶约占80%;两个竹种凋落物在分解过程中养分元素释放的种间差异主要与初始养分元素含量有关;凋落物养分元素初始含量对元素释放模式和最终净释放率的大小具有重要的决定作用;目前,这两种竹林生态系统土壤氮输入主要以大气氮沉降(8.24 g · m^-2 · a^-1)为主,同时凋落物氮输入(苦竹和杂交竹林分别为1.93,5.07 g · m^-2 · a^-1)也是一个重要途径;模拟氮沉降对苦竹凋落物碳、磷、钾、钙元素和杂交竹凋落物碳、氮、磷、钾、钙、镁元素释放的抑制作用较弱,处理与对照之间元素总释放率差异一般小于10%;氮沉降显著抑制了苦竹林凋落物氮元素释放,减小幅度为19.0%-27.2%,但由于氮沉降增加对土壤肥力的直接改良作用,氮沉降的增加并不会因为凋落物分解速率的降低造成植物生长所需养分供应的减少;从短期来看,在氮沉降继续增加的情况下,该地区这类竹林生态系统的碳吸存能力仍可能会因为N沉降对植物生长的促进作用而增加。     

Macroalgal detrital systems: an overlooked ecological niche for heterotrophic nitrogen fixation

Diazotrophic macroalgal associations (DMAs) can contribute fixed nitrogen (N) to the host macroalgae. Biological nitrogen fixation (BNF) rates investigated using acetylene reduction assays with living macroalgae surrounding Santa Catalina Island were low (maximum: 36 nmol N × g⁻¹ (dw) × h⁻¹) and probably insufficient towards helping meet macroalgal N demand. However, DMAs were observed during periods of low nitrate availability in Southern California coastal waters, highlighting the potential importance of diazotrophs during N depleted conditions. Eleven long-term (16–32 days) litter bag decomposition experiments with various macroalgae, especially those with high (> 10) C:N ratios, resulted in much higher BNF rates (maximum: 693 nmol N × g⁻¹ (dw) × h⁻¹) than observed with living macroalgae. BNF rates were lower at the beginning of macroalgal decomposition but rapidly increased during the second phase before declining towards the end of decomposition. Labile carbon availability is likely influencing BNF rates throughout macroalgal degradation and limits BNF in the final decomposition stage. Comparable dark and light BNF rates with most macroalgae surveyed suggest macroalgal detrital systems are an overlooked, potentially global, niche for heterotrophic N₂ fixation. Lastly, suppressed BNF rates with sodium molybdate additions highlight the prevalence of sulfate reducing diazotrophs.     

Effects of heather and oat supplementation on gastrointestinal nematode infections and performance of grazing Cashmere goats

This work aimed to evaluate the effects of tannin-containing heather (Calluna vulgaris, Erica spp.) and energy (oats, Avena sativa) supplementation, combined or not, on feed intake, gastrointestinal nematode infections and performance of goats grazing mountain grasslands. Two successive experiments were established across one grazing season on four paddocks. The first (late April to early August, Period 1) involved two treatments, i.e., supplementation with heather (+H) vs. non-supplementation (−H), each randomly allocated to two paddocks. The second (mid-August to mid-November, Period 2) consisted on four treatments in a 2 × 2 factorial design, i.e., supplementation with heather and oats (+H+O), only heather (+H−O), only oats (−H+O), and no supplementation (−H−O). Results from Period 1 indicated that faecal nematode egg counts (FEC) were lower in +H goats (50% less in August), achieving more favourable live weight (LW) and body condition changes than −H goats. Total dry matter intake (DMI) in June was similar between treatments, with heather accounting for 0.21 of +H goat diets. Kids FEC and LW gains were not affected by heather supplementation. In Period 2, both heather and oat supplementation reduced FEC (45 and 61%, respectively) and improved goat performance, without an interaction between both factors. Total DMI in October was higher in +H+O compared to other treatments (48 vs. 30 g DM kg LW^−0.75 d⁻¹; P < 0.01). Rumen ammonia concentration was lower (P < 0.001) in +H than in −H goats, and in +O than in −O goats, whereas that of volatile fatty acids (VFA) was not affected by treatments, though the molar proportions of some VFA were modified. It is concluded that the combination of both supplements (tannins and energy) contributes to reduce gastrointestinal nematode parasitism and increases goat performance, allowing a lower dependence on conventional chemotherapy.     

Effect of prostaglandin F2α on propulsive activity of the isolated segmental colon of the guinea-pig

The effects of prostaglandin F_2α (PGF_2α) on propulsive activity in segments of isolated colon and on isolated strips of guinea-pig colon were investigated.Using experimental conditions under which spontaneous propulsive activity was negligible, PGF_2α (5×10⁻⁸×1×10⁻⁶M), added to the bathing medium, increased propulsive activity in a concentration dependent manner. This increase of propulsive activity was abolished in the presence of atropine or tetrodotoxin (1×10⁻⁷g/ml).The contractions produced by PGF_2α(5×10⁻⁷ − 1×10⁻⁵M) in isolated longitudinal and circular smooth muscle strips of guinea-pig colon were unaffected in the presence of atropine or tetrodotoxin (1×10⁻⁷g/ml).From these results it is concluded that under the conditions employed in this study propulsive activity stimulated by PGF_2α may depend on the contractions of both muscle layers and stimulation of the peristalic reflex.     

Differential effects of ammonium and nitrate deposition on fen phanerogams and bryophytes

Question: High atmospheric nitrogen (N) deposition has been shown to affect productivity and species composition of terrestrial ecosystems. This study focused on the differential effects of the two inorganic N forms in atmospheric deposition (i.e. ammonium and nitrate). Methods and location: Nutrient addition experiments were carried out during 4 years in a mesotrophic fen in a low‐deposition area in Ireland. In a factorial design, plots were fertilized with ammonium and/or nitrate, in two doses comparable with 35 and 70 kg N ha^?1 y^?1 and compared with an unfertilized control. Results: Vascular plant biomass as well as bryophyte biomass were not affected by N dose but showed significantly different responses to the N form. In the ammonium‐fertilized plots, vascular plant biomass was higher and moss biomass was lower than the control, while nitrate additions had no effect. Vascular plant species density was high (16 species per 0.49 m²) and was not affected by any of the treatments; bryophyte species density was also high (seven species per 0.04 m²) but showed a significant decrease upon ammonium fertilization. Conclusion: The vulnerability of the mesotrophic vegetation to enhanced atmospheric N deposition depends strongly on the N form. If N would be mainly deposited as NO_x, no detrimental effects on the vegetation will occur. If, however, the deposition is mainly in the form of NH_y, the bryophyte vegetation will be seriously damaged, while the vascular plant vegetation will show an increased biomass production with possible shifts in dominance from Carex and herb species to grasses and shrubs.     

Nitrogen deposition and plant species interact to influence soil carbon stabilization

Anthropogenic nitrogen (N) deposition effects on soil organic carbon (C) decomposition remain controversial, while the role of plant species composition in mediating effects of N deposition on soil organic C decomposition and long‐term soil C sequestration is virtually unknown. Here we provide evidence from a 5‐year grassland field experiment in Minnesota that under elevated atmospheric CO₂ concentration (560 ppm), plant species determine whether N deposition inhibits the decomposition of soil organic matter via inter‐specific variation in root lignin concentration. Plant species producing lignin‐rich litter increased stabilization of soil C older than 5 years, but only in combination with elevated N inputs (4 g m^?2 year^?1). Our results suggest that N deposition will increase soil C sequestration in those ecosystems where vegetation composition and/or elevated atmospheric CO₂ cause high litter lignin inputs to soils.     

滇中亚高山地带性植被凋落物分解对模拟氮沉降的响应

模拟氮(N)沉降对凋落物分解特征的影响对研究森林生态系统物质循环响应大气N沉降的内在机理和应对N沉降全球化具有重要意义。从2018年2月至2019年1月,对滇中亚高山常绿阔叶林(Evergreen broad-leaf forest)和高山栎林(Quercus semecarpifolia forest)两种地带性植被进行模拟N沉降试验,利用尼龙网袋法对两种林型凋落叶和凋落枝进行原位分解试验,N沉降处理水平分别为对照CK(Control check, 0 g N m^-2 a^-1)、低氮LN(Low nitrogen, 5 g N m^-2 a^-1)、中氮MN(Medium nitrogen, 15 g N m^-2 a^-1)和高氮HN(High nitrogen, 30 g N m^-2 a^-1)。结果表明:常绿阔叶林凋落叶和凋落枝分解率分别为44.84%和21.96%,均高于高山栎林的35.97%(凋落叶)和17.5...     

Tree ring responses to elevated CO2 and increased N deposition in Picea abies

Four- to seven-year-old spruce trees (Picea abies) were exposed to three CO₂ concentrations (280, 420 and 560 cm³ m^?3) and three rates of wet N deposition (0, 30 and 90 kg ha^?1 year^?1) for 3 years in a simulated montane forest climate. Six trees from each of six clones were grown in competition in each of nine 100 × 70 × 36 cm model ecosystems with nutrient-poor natural forest soil. Stem dises were analysed using X-ray densitometry. The radial stem increment was not affected by [CO₂] but increased with increasing rates of N deposition. Wood density was increased by [CO₂], but decreased by N deposition. Wood-starch concentration increased, and wood nitrogen concentration decreased with increasing [CO₂], but neither was affected by N deposition. The lignin concentration in wood was affected by neither [CO₂] nor N deposition. Our results suggest that, under natural growth conditions, rising atmospheric [CO₂] will not lead to enhanced radial stem growth of spruce, but atmospheric N deposition will, and in some regions is probably already doing so. Elevated [CO₂], however, will lead to denser wood unless this effect is compensated by massive atmospheric N deposition. If can be speculated that greater wood density under elevated [CO₂] may alter the mechanical properties of wood, and higher ratios of C/N and lignin/N in wood grown at elevated [CO₂] may affect nutrient cycles of forest ecosystems.     

Impact of sheep grazing on nutrient budgets of dry heathlands

Questions: What effect does sheep grazing have on the nutrient budgets of heathlands? Can grazing compensate for atmospheric nutrient loads in heathland ecosystems? What are the conclusions for heathland management? Location: Lüneburg Heath, NW Germany. Methods: During a one-year grazing experiment (stocking rate 1.1 sheep/ha) nutrient balances for N, Ca, K, Mg and P were calculated by quantifying input rates (atmospheric deposition, sheep excrement) and output rates (biomass removal, leaching). Results: Atmospheric nutrient deposition amounted to 22.8 kg.ha⁻¹.a⁻¹ for N and < 0.2 kg.ha⁻¹.a⁻¹ for P. Sheep excrement increased the inputs for N and P by ca. 3.5 and 0.2 kg.ha⁻¹.a⁻¹, respectively. Grazing reduced N- and P-stores in the above-ground biomass by 25.6 and 1.9 kg.ha⁻¹.a⁻¹, respectively. N-and P-losses via leaching amounted to 2.2 and < 0.2 kg.ha⁻¹.a⁻¹. Output:input ratios for P were high, indicating that grazing severely affected P-budgets of heaths. Conclusions: Our results suggest that sheep grazing has the potential to compensate for atmospheric nutrient loads (particularly for current N deposition rates). However, in the long term the combination of elevated N-deposition and P-loss due to grazing may cause a shift from N-(co-) limited to more P-(co-) limited plant growth. To counteract an aggravation of P-deficiency in the long term, grazing may be combined with management measures that affect P-budgets to a lesser extent (e.g. prescribed burning).     

The impact of elevated CO2, increased nitrogen availability and biodiversity on plant tissue quality and decomposition

Elevated CO₂, increased nitrogen (N) deposition and increasing species richness can increase net primary productivity (NPP). However, unless there are comparable changes in decomposition, increases in productivity will most likely be unsustainable. Without comparable increases in decomposition nutrients would accumulate in dead organic matter leading to nutrient limitations that could eventually prohibit additional increases in productivity. To address this issue, we measured aboveground plant and litter quality and belowground root quality, as well as decomposition of aboveground litter for one and 2‐year periods using in situ litterbags in response to a three‐way factorial manipulation of CO₂ (ambient vs. 560 ppm), N deposition (ambient vs. the addition of 4 g N m⁻² yr⁻¹) and plant species richness (one, four, nine and 16 species) in experimental grassland plots. Litter chemistry responded to the CO₂, N and plant diversity treatments, but decomposition was much less responsive. Elevated CO₂ induced decreases in % N and % lignin in plant tissues. N addition led to increases in % N and decreases in % lignin. Increasing plant diversity led to decreases in % N and % lignin and an increase in % cellulose. In contrast to the litter chemistry changes, elevated CO₂ had a much lower impact on decomposition and resulted in only a 2.5% decrease in carbon (C) loss. Detectable responses were not observed either to N addition or to species richness. These results suggest that global change factors such as biodiversity loss, elevated CO₂ and N deposition lead to significant changes in tissue quality; however, the response of decomposition is modest. Thus, the observed increases in productivity at higher diversity levels and with elevated CO₂ and N fertilization are not matched by an increase in decomposition rates. This lack of coupled responses between production and decomposition is likely to result in an accumulation of nutrients in the litter pool which will dampen the response of NPP to these factors over time.     

Comparison of the effects of canopy and understory nitrogen addition on xylem growth of two dominant species in a warm temperate forest,China

Understanding the effect of increasing atmospheric nitrogen (N) deposition on xylem growth of trees is critical to predict tree growth and carbon sequestration under global change. Canopy N addition (CAN) is generally believed to realistically simulate atmospheric N deposition on terrestrial ecosystems given it takes all processes of N deposition from forest canopy to belowground into account. However, whether CAN is more effective in reflecting the effect of atmospheric N deposition on xylem growth of trees than understory N addition (UAN) has been rarely reported. To address the question, we conducted a CAN vs. UAN experimental study to weekly monitor xylem growth of two dominant broadleaf species (Quercus acutissima Carruth. and Quercus variabilis Blume) in a warm temperate forest of China during 2014–2015. Weekly xylem increment during the two years was measured. Mixed-effects models were used to quantify the effects of N addition on xylem growth and detect the differences among treatments. We found that CAN of 50 kg N ha⁻¹ yr⁻¹ plays a more significant role in promoting xylem growth of Q. acutissima than UAN of 50 kg N ha⁻¹ yr⁻¹, and significantly enhanced the formation of differentiating xylem (zones of radial enlarging and wall-thickening cells) of Q. acutissima in the early growing season (April-June) and the rate of xylem increment, but no significant difference in xylem increment of Q. variabilis was detected between CAN50 and UAN50. This is the first study to quantitatively demonstrate that previous UAN studies may have underestimated the effects of atmospheric N deposition on tree growth by ignoring the N interception through forest canopy. Furthermore, our study also suggested a species-specific response of xylem growth to N addition. Under a certain amount of atmospheric N deposition in the future, the xylem increment of Q. acutissima may be superior to that of Q. variabilis.