Tree species impact the terrestrial cycle of silicon through various uptakes

The quantification of silicon (Si) uptake by tree species is a mandatory step to study the role of forest vegetations in the global cycle of Si. Forest tree species can impact the hydrological output of dissolved Si (DSi) through root induced weathering of silicates but also through Si uptake and restitution via litterfall. Here, monospecific stands of Douglas fir, Norway spruce, Black pine, European beech and oak established in identical soil and climate conditions were used to quantify Si uptake, immobilization and restitution. We measured the Si contents in various compartments of the soil–tree system and we further studied the impact of the recycling of Si by forest trees on the DSi pool. Si is mainly accumulated in leaves and needles in comparison with other tree compartments (branches, stembark and stemwood). The immobilization of Si in tree biomass represents less than 15% of the total Si uptake. Annual Si uptake by oak and European beech stands is 18.5 and 23.3 kg ha^?1 year^?1, respectively. Black pine has a very low annual Si uptake (2.3 kg ha^?1 year^?1) in comparison with Douglas fir (30.6 kg ha^?1 year^?1) and Norway spruce (43.5 kg ha^?1 year^?1). The recycling of Si by forest trees plays a major role in the continental Si cycle since tree species greatly influence the uptake and restitution of Si. Moreover, we remark that the annual tree uptake is negatively correlated with the annual DSi output at 60 cm depth. The land–ocean fluxes of DSi are certainly influenced by geochemical processes such as weathering of primary minerals and formation of secondary minerals but also by biological processes such as root uptake.     

Control of Nitrous Oxide Emissions in European Beech,Norway Spruce and Scots Pine Forests

Elevated nitrogen deposition has increased tree growth, the storage of soil organic matter, and nitrate leaching in many European forests, but little is known about the effect of tree species and nitrogen deposition on nitrous oxide emission. Here we report soil N₂O emission from European beech, Scots pine and Norway spruce forests in two study areas of Germany with distinct climate, N deposition and soils. N₂O emissions and throughfall input of nitrate and ammonium were measured biweekly during growing season and monthly during dormant season over a 28 months period. Annual N₂O emission rates ranged between 0.4 and 1.3 kg N ha^?1 year^?1 among the stands and were higher in 1998 than in 1999 due to higher precipitation during the growing season of 1998. A 2-way-ANOVA revealed that N₂O fluxes were significantly higher (p<0.001) at Solling than at Unterlüß while tree species had no effect on N₂O emissions. Soil texture and the amount of throughfall explained together 94% of the variance among the stands, indicating that increasing portions of silt and clay may promote the formation of N₂O in wet forest soils. Moreover, cumulative N₂O fluxes were significantly correlated (r² = 0.60, p<0.001) with cumulative NO ₃ ^? fluxes at 10 cm depth as an indicator of N saturation, however, the slope of the regression curve indicates a rather weak effect of NO ₃ ^? fluxes on N₂O emissions. N input by throughfall was not correlated with N₂O emissions and only 1.6–3.2% of N input was released as N₂O to the atmosphere. Our results suggest that elevated N inputs have little effect on N₂O emissions in beech, spruce and pine forests.     

N deposition, N transformation and N leaching in acid forest soils

Nitrogen deposition, mineralisation, uptake and leaching were measured on a monthly basis in the field during 2 years in six forested stands on acidic soils under mountainous climate. Studies were conducted in three Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] plantations (D20: 20 year; D40: 40 yr; D60: 60 yr) on abandoned croplands in the Beaujolais Mounts; and two spruce (Picea abies Karst.) plantations (S45: 45 yr; S90: 90 yr) and an old beech (Fagus sylvatica L.) stand (B150: 150 yr) on ancient forest soils in a small catchment in the Vosges Mountains. N deposition in throughfall varied between 7–8 kg ha^–1 year^–1 (D20, B150, S45) and 15–21 kg ha^–1 yr^–1 (S90, D40, D60). N in annual litterfall varied between 20–29 kg ha^–1 (D40, D60, S90), and 36–43 kg ha^–1 (D20, S45, B150). N leaching below root depth varied among stands within a much larger range, between 1–9 kg ha^–1 yr^–1 (B150, S45, D60) and 28–66 kg ha^–1 yr^–1 (D40, S90, D20), with no simple relationship with N deposition, or N deposition minus N storage in stand biomass. N mineralisation was between 57–121 kg ha^–1 yr^–1 (S45, D40, S90) and between 176–209 kg ha^–1 yr^–1 in (B150, D60 and D20). The amounts of nitrogen annually mineralised and nitrified were positively related. Neither general soil parameters, such as pH, soil type, base saturation and C:N ratio, nor deposition in throughfall or litterfall were simply related to the intensity of mineralisation and/or nitrification. When root uptake was not allowed, nitrate leaching increased by 11 kg ha^–1 yr^–1 at S45, 36 kg ha^–1 yr^–1 at S90 and between 69 and 91 kg ha^–1 yr^–1 at D20, D40, B150 and D60, in relation to the nitrification rates of each plot. From this data set and recent data from the literature, we suggest that: high nitrification and nitrate leaching in Douglas-fir soils was likely related to the former agricultural land use. High nitrification rate but very low nitrate leaching in the old beech soil was related to intense recycling of mineralised N by beech roots. Medium nitrification and nitrate leaching in the old spruce stand was related to the average level of N deposition and to the deposition and declining health of the stand. Very low nitrification and N leaching in the young spruce stand were considered representative of fast growing spruce plantations receiving low N deposition on acidic soils of ancient coniferous forests. Consequently, we suggest that past land use and fine root cycling (which is dependent on to tree species and health) should be taken into account to explain the variability in the relation between N deposition and leaching in forests.     

Effects of logging on carbon dynamics of a jack pine forest in Saskatchewan,Canada

We calculated carbon budgets for a chronosequence of harvested jack pine (Pinus banksiana Lamb.) stands (0‐, 5‐, 10‐, and～29‐year‐old) and a～79‐year‐old stand that originated after wildfire. We measured total ecosystem C content (TEC), above‐, and belowground net primary productivity (NPP) for each stand. All values are reported in order for the 0‐, 5‐, 10‐, 29‐, and 79‐year‐old stands, respectively, for May 1999 through April 2000. Total annual NPP (NPP_T) for the stands (Mg C ha^?1 yr^?1±1 SD) was 0.9±0.3, 1.3±0.1, 2.7±0.6, 3.5±0.3, and 1.7±0.4. We correlated periodic soil surface CO₂ fluxes (R_S) with soil temperature to model annual R_S for the stands (Mg C ha^?1 yr^?1±1 SD) as 4.4±0.1, 2.4±0.0, 3.3±0.1, 5.7±0.3, and 3.2±0.2. We estimated net ecosystem productivity (NEP) as NPP_T minus R_H (where R_H was calculated using a Monte Carlo approach as coarse woody debris respiration plus 30–70% of total annual R_S). Excluding C losses during wood processing, NEP (Mg C ha^?1 yr^?1±1 SD) for the stands was estimated to be ?1.9±0.7, ?0.4±0.6, 0.4±0.9, 0.4±1.0, and ?0.2±0.7 (negative values indicate net sources to the atmosphere.) We also calculated NEP values from the changes in TEC among stands. Only the 0‐year‐old stand showed significantly different NEP between the two methods, suggesting a possible mismatch for the chronosequence. The spatial and methodological uncertainties allow us to say little for certain except that the stand becomes a source of C to the atmosphere following logging.     

Downward adjustment of carbon fluxes at the biochemical, leaf, and ecosystem scale in beech-spruce model communities exposed to long-term atmospheric CO2 enrichment

Young beech (Fagus sylvatica) and spruce (Picea abies) trees from different provenances or genotypes were grown in competition in large model ecosystems and were exposed to two concentrations of atmospheric CO₂ (370 vs 570 μmol mol^?1), two levels of wet nitrogen deposition (7 vs 70 kg N ha^?1 yr^?1), and two native forest soils (acidic vs calcareous) for four years in open‐top chambers. The 2×2×2 factorial experimental design was fully replicated (n=4) with each CO₂×N combination applied to each soil type. Exposure to atmospheric CO₂ enrichment stimulated daytime net ecosystem CO₂ flux (NEC) as measured during sunny days in the middle of the third growing season. Nevertheless, we observed substantial down‐regulation of NEC, with larger adjustments on acidic than on calcareous soil. NEC adjustment was associated with slightly reduced leaf area index (LAI) on the acidic soil (no response on calcareous soil), enhanced soil CO₂ efflux from both substrate types, and, most importantly, with down‐regulation of CO₂ uptake at the leaf scale. Downward adjustment of light‐saturated single‐leaf photosynthesis (A) and of Rubisco was more pronounced in beech than in spruce and these species‐specific differences increased over time. By year four, A adjustment (except in one specific treatment combination in each species) had become complete in beech but had disappeared in spruce. At no time did we observe a genotype or provenance effect on the downward adjustment of carbon fluxes, and nitrogen deposition rate generally had little effect as well. Overall, our results suggest that tree species and soil quality will have profound effects on ecosystem CO₂ fluxes under continued atmospheric CO₂ enrichment.     

Influence of Tree Species on Forest Nitrogen Retention in the Catskill Mountains, New York, USA

This study examines the effect of four tree species on nitrogen (N) retention within forested catchments of the Catskill Mountains, New York (NY). We conducted a 300-day ¹⁵N field tracer experiment to determine how N moves through soil, microbial, and plant pools under different tree species and fertilization regimes. Samples were collected from single-species plots of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga canadensis L.), red oak (Quercus rubra L.), and sugar maple (Acer saccharum Marsh). Using paired plots we compared the effects of ambient levels of N inputs (11 kg N/ha/y) to additions of 50 kg N/ha/y that began 1.5 years prior to and continued throughout this experiment. Total plot ¹⁵N recovery (litter layer, organic and mineral soil to 12 cm, fine roots, and aboveground biomass) did not vary significantly among tree species, but the distribution of sinks for ¹⁵N within the forest ecosystem did vary. Recovery in the forest floor was significantly lower in sugar maple stands compared to the other species. ¹⁵Nitrogen recovery was 22% lower in the fertilized plots compared to the ambient plots and red oak stands had the largest drop in ¹⁵N recovery as a result of N fertilization. Aboveground biomass became a significantly greater ¹⁵N sink with fertilization, although it retained less than 1% of the tracer addition. These results indicate that different forest types vary in the amount of N retention in the forest floor, and that forest N retention may change depending upon N inputs.     

Site and temporal variation of soil respiration in European beech, Norway spruce, and Scots pine forests

Global warming and changes in rainfall amount and distribution may affect soil respiration as a major carbon flux between the biosphere and the atmosphere. The objectives of this study were to investigate the site to site and interannual variation in soil respiration of six temperate forest sites. Soil respiration was measured using closed chambers over 2 years under mature beech, spruce and pine stands at both Solling and Unterlüß, Germany, which have distinct climates and soils. Cumulative annual CO₂ fluxes varied from 4.9 to 5.4 Mg C ha^?1 yr^?1 at Solling with silty soils and from 4.0 to 5.9 Mg C ha^?1 yr^?1 at Unterlüß with sandy soils. With one exception soil respiration rates were not significantly different among the six forest sites (site to site variation) and between the years within the same forest site (interannual variation). Only the respiration rate in the spruce stand at Unterlüß was significant lower than the beech stand at Unterlüß in both years. Soil respiration rates of the sandy sites at Unterlüß were limited by soil moisture during the rather dry and warm summer 1999 while soil respiration at the silty Solling site tended to increase. We found a threshold of ?80 kPa at 10 cm depth below which soil respiration decreased with increasing drought. Subsequent wetting of sandy soils revealed high CO₂ effluxes in the stands at Unterlüß. However, dry periods were infrequent, and our results suggest that temporal variation in soil moisture generally had little effect on annual soil respiration rates. Soil temperature at 5 cm and 10 cm depth explained 83% of the temporal variation in soil respiration using the Arrhenius function. The correlations were weaker using temperature at 0 cm (r² = 0.63) and 2.5 cm depth (r² = 0.81). Mean Q₁₀ values for the range from 5 to 15 °C increased asymptotically with soil depth from 1.87 at 0 cm to 3.46 at 10 cm depth, indicating a large uncertainty in the prediction of the temperature dependency of soil respiration. Comparing the fitted Arrhenius curves for same tree species from Solling and Unterlüß revealed higher soil respiration rates for the stands at Solling than in the respective stands at Unterlüß at the same temperature. A significant positive correlation across all sites between predicted soil respiration rates at 10 °C and total phosphorus content and C‐to‐N ratio of the upper mineral soil indicate a possible effect of nutrients on soil respiration.     

Effects of soil freezing disturbance on soil solution nitrogen, phosphorus, and carbon chemistry in a northern hardwood ecosystem

Reductions in snow cover undera warmer climate may cause soil freezing eventsto become more common in northern temperateecosystems. In this experiment, snow cover wasmanipulated to simulate the late development ofsnowpack and to induce soil freezing. Thismanipulation was used to examine the effects ofsoil freezing disturbance on soil solutionnitrogen (N), phosphorus (P), and carbon (C)chemistry in four experimental stands (twosugar maple and two yellow birch) at theHubbard Brook Experimental Forest (HBEF) in theWhite Mountains of New Hampshire. Soilfreezing enhanced soil solution Nconcentrations and transport from the forestfloor. Nitrate (NO₃ ^–) was thedominant N species mobilized in the forestfloor of sugar maple stands after soilfreezing, while ammonium (NH₄ ⁺) anddissolved organic nitrogen (DON) were thedominant forms of N leaching from the forestfloor of treated yellow birch stands. Rates ofN leaching at stands subjected to soil freezingranged from 490 to 4,600 mol ha^–1yr^–1, significant in comparison to wet Ndeposition (530 mol ha^–1 yr^–1) andstream NO₃ ^– export (25 mol ha^–1yr^–1) in this northern forest ecosystem. Soil solution fluxes of P_i from the forestfloor of sugar maple stands after soil freezingranged from 15 to 32 mol ha^–1 yr^–1;this elevated mobilization of P_i coincidedwith heightened NO₃ ^– leaching. Elevated leaching of P_i from the forestfloor was coupled with enhanced retention ofP_i in the mineral soil Bs horizon. Thequantities of P_i mobilized from the forestfloor were significant relative to theavailable P pool (22 mol ha^–1) as well asnet P mineralization rates in the forest floor(180 mol ha^–1 yr^–1). Increased fineroot mortality was likely an important sourceof mobile N and P_i from the forest floor,but other factors (decreased N and P uptake byroots and increased physical disruption of soilaggregates) may also have contributed to theenhanced leaching of nutrients. Microbialmortality did not contribute to the acceleratedN and P leaching after soil freezing. Resultssuggest that soil freezing events may increaserates of N and P loss, with potential effectson soil N and P availability, ecosystemproductivity, as well as surface wateracidification and eutrophication.     

Modeling to discern nitrogen fertilization impacts on carbon sequestration in a Pacific Northwest Douglas‐fir forest in the first‐postfertilization year

This study investigated how nitrogen (N) fertilization with 200 kg N ha^?1 of urea affected ecosystem carbon (C) sequestration in the first‐postfertilization year in a Pacific Northwest Douglas‐fir (Pseudotsuga menziesii) stand on the basis of multiyear eddy‐covariance (EC) and soil‐chamber measurements before and after fertilization in combination with ecosystem modeling. The approach uses a data‐model fusion technique which encompasses both model parameter optimization and data assimilation and minimizes the effects of interannual climatic perturbations and focuses on the biotic and abiotic factors controlling seasonal C fluxes using a prefertilization 9‐year‐long time series of EC data (1998–2006). A process‐based ecosystem model was optimized using the half‐hourly data measured during 1998–2005, and the optimized model was validated using measurements made in 2006 and further applied to predict C fluxes for 2007 assuming the stand was not fertilized. The N fertilization effects on C sequestration were then obtained as differences between modeled (unfertilized stand) and EC or soil‐chamber measured (fertilized stand) C component fluxes. Results indicate that annual net ecosystem productivity in the first‐post‐N fertilization year increased by～83%, from 302 ± 19 to 552 ± 36 g m^?2 yr^?1, which resulted primarily from an increase in annual gross primary productivity of～8%, from 1938 ± 22 to 2095 ± 29 g m^?2 yr^?1 concurrent with a decrease in annual ecosystem respiration (R_e) of～5.7%, from 1636 ± 17 to 1543 ± 31 g m^?2 yr^?1. Moreover, with respect to respiration, model results showed that the fertilizer‐induced reduction in R_e (～93 g m^?2 yr^?1) principally resulted from the decrease in soil respiration R_s (～62 g m^?2 yr^?1).     

Plant and soil natural abundance <Emphasis Type="Italic">δ</Emphasis><Superscript>15</Superscript>N: indicators of relative rates of nitrogen cycling in temperate forest ecosystems

Watersheds within the Catskill Mountains, New York, receive among the highest rates of nitrogen (N) deposition in the northeastern United States and are beginning to show signs of N saturation. Despite similar amounts of N deposition across watersheds within the Catskill Mountains, rates of soil N cycling and N retention vary significantly among stands of different tree species. We examined the potential use of δ ¹⁵N of plants and soils as an indicator of relative forest soil N cycling rates. We analyzed the δ ¹⁵N of foliage, litterfall, bole wood, surface litter layer, fine roots and organic soil from single-species stands of American beech (Fagus grandifolia), eastern hemlock (Tsuga canadensis), red oak (Quercus rubra), and sugar maple (Acer saccharum). Fine root and organic soil δ ¹⁵N values were highest within sugar maple stands, which correlated significantly with higher rates of net mineralization and nitrification. Results from this study suggest that fine root and organic soil δ ¹⁵N can be used as an indicator of relative rates of soil N cycling. Although not statistically significant, δ ¹⁵N was highest within foliage, wood and litterfall of beech stands, a tree species associated with intermediate levels of soil N cycling rates and forest N retention. Our results show that belowground δ ¹⁵N values are a better indicator of relative rates of soil N cycling than are aboveground δ ¹⁵N values.     

温带两种林型对氮沉降的再分配及其生长季动态与影响因子

氮(N)沉降对森林生态系统的结构与功能具有重要的影响,而森林对到达林地的N沉降量及其分配格局的影响尚不清楚。量化了2012—2013年5—10月两个生长季蒙古栎林和杂木林的林内树干径流和穿透雨及其林外大气降雨总氮(TN)、可溶性氮(DN)和颗粒态氮(PN)沉降通量的季节动态,旨在比较两种林型对N沉降的再分配格局及其季节变化,分析影响其变异的主要因子。结果表明:林外大气降雨、蒙古栎林林内、杂木林林内(树干径流+穿透雨)TN沉降平均值分别为:8.49、15.97、13.39 kg hm~(-2)a~(-1),其中DN分别占其TN的76.35%、82.79%和75.02%,PN分别占其TN的26.35%、17.21%和24.98%,蒙古栎林和杂木林林内穿透雨TN沉降量分别占其TN的95.5%和94.5%。蒙古栎林和杂木林冠层淋溶TN沉降量分别为7.48kg hm~(-2)a~(-1)和4.90 kg hm~(2)a~(-1);其中,前者的DN高于后者,但PN呈相反趋势。两种林型的N沉降组分具有明显的季节动态:沉降量均集中在生长季中期(6—8月),生长季前期和末期较低。林外降雨量分别与林外大气降雨、蒙古栎林和杂木林林内的树干径流和穿透雨中的TN、DN浓度呈显著负指数函数关系(P0.001)。连续降雨天数对蒙古栎林、杂木林林内TN、DN浓度的影响表现为连续降雨2 d以内为富集作用,之后为稀释作用。研究表明林冠对大气氮沉降有显著富集作用,其富集强度及时间动态与森林类型和降雨特征有关,建议氮沉降试验应考虑林冠的富集效应。     

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

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

Isotopic evidence for the occurrence of biological nitrification and nitrogen deposition processing in forest canopies

This study examines the role of tree canopies in processing atmospheric nitrogen (N_dep) for four forests in the United Kingdom subjected to different N_dep: Scots pine and beech stands under high N_dep (HN, 13–19 kg N ha^?1 yr^?1), compared to Scots pine and beech stands under low N_dep (LN, 9 kg N ha^?1 yr^?1). Changes of NO₃‐N and NH₄‐N concentrations in rainfall (RF) and throughfall (TF) together with a quadruple isotope approach, which combines δ¹⁸O, Δ¹⁷O and δ¹⁵N in NO₃^? and δ¹⁵N in NH₄⁺, were used to assess N transformations by the canopies. Generally, HN sites showed higher NH₄‐N and NO₃‐N concentrations in RF compared to the LN sites. Similar values of δ¹⁵N‐NO₃^? and δ¹⁸O in RF suggested similar source of atmospheric NO₃^? (i.e. local traffic), while more positive values for δ¹⁵N‐NH₄⁺ at HN compared to LN likely reflected the contribution of dry NH_x deposition from intensive local farming. The isotopic signatures of the N‐forms changed after interacting with tree canopies. Indeed, ¹⁵N‐enriched NH₄⁺ in TF compared to RF at all sites suggested that canopies played an important role in buffering dry N_dep also at the low N_dep site. Using two independent methods, based on δ¹⁸O and Δ¹⁷O, we quantified for the first time the proportion of NO₃^? in TF, which derived from nitrification occurring in tree canopies at the HN site. Specifically, for Scots pine, all the considered isotope approaches detected biological nitrification. By contrast for the beech, only using the mixing model with Δ¹⁷O, we were able to depict the occurrence of nitrification within canopies. Our study suggests that tree canopies play an active role in the N cycling within forest ecosystems. Processing of N_dep within canopies should not be neglected and needs further exploration, with the combination of multiple isotope tracers, with particular reference to Δ¹⁷O.     

Forest and agricultural land-use-dependent CO2 exchange in Thuringia, Germany

Eddy covariance was used to measure the net CO₂ exchange (NEE) over ecosystems differing in land use (forest and agriculture) in Thuringia, Germany. Measurements were carried out at a managed, even‐aged European beech stand (Fagus sylvatica, 70–150 years old), an unmanaged, uneven‐aged mixed beech stand in a late stage of development (F. sylvatica, Fraxinus excelsior, Acer pseudoplantanus, and other hardwood trees, 0–250 years old), a managed young Norway spruce stand (Picea abies, 50 years old), and an agricultural field growing winter wheat in 2001, and potato in 2002. Large contrasts were found in NEE rates between the land uses of the ecosystems. The managed and unmanaged beech sites had very similar net CO₂ uptake rates (～?480 to ?500 g C m^?2 yr^?1). Main differences in seasonal NEE patterns between the beech sites were because of a later leaf emergence and higher maximum leaf area index at the unmanaged beech site, probably as a result of the species mix at the site. In contrast, the spruce stand had a higher CO₂ uptake in spring but substantially lower net CO₂ uptake in summer than the beech stands. This resulted in a near neutral annual NEE (?4 g C m^?2 yr^?1), mainly attributable to an ecosystem respiration rate almost twice as high as that of the beech stands, despite slightly lower temperatures, because of the higher elevation. Crops in the agricultural field had high CO₂ uptake rates, but growing season length was short compared with the forest ecosystems. Therefore, the agricultural land had low‐to‐moderate annual net CO₂ uptake (?34 to ?193 g C m^?2), but with annual harvest taken into account it will be a source of CO₂ (+97 to +386 g C m^?2). The annually changing patchwork of crops will have strong consequences on the regions' seasonal and annual carbon exchange. Thus, not only land use, but also land‐use history and site‐specific management decisions affect the large‐scale carbon balance.     

The Role of Wood Ants (Formica rufa group) in Carbon and Nutrient Dynamics of a Boreal Norway Spruce Forest Ecosystem

Wood ants (Formica rufa group) are regarded as keystone species in boreal and mountain forests of Europe and Asia by their effect on ecosystem carbon (C) and nutrient pools and fluxes. To quantify the impact of their activity on boreal forest ecosystems, C, nitrogen (N), phosphorus (P), potassium (K) and calcium (Ca) pools and fluxes in wood ant nests (WAN), and soil were assessed along a 5-, 30-, 60-, and 100-year-old Norway spruce (Picea abies L. Karsten) dominated successional gradient in eastern Finland. Amounts of C and nutrients in WAN increased with stand age, but contained less than 1% of total C and nutrient pools in these stands. The CO₂-efflux from nests was also insignificant, as compared to CO₂-efflux from the forest floor. Annually, the amount of C brought by wood ants into their nests as honeydew, prey and nest-building materials ranged from 2.7 to 49.3 kg ha^?1 C, but this is only 0.1–0.7% of the combined net primary production of trees and understorey in boreal forests. The difference between wood ant nest C inputs and outputs was very small in the younger-aged stands, and increased in the older stands. Carbon accumulation rates in nests over a 100 year period are estimated to be less than 10 kg ha^?1 a^?1. In contrast to C, annual inputs of N, P, and K are larger compared to wood ant nest nutrient pool size, ranging from 3 to 6% of the annual tree stand and understorey uptake. This indicates a more rapid turnover and transport of N, P, and K out of WAN, and suggests that wood ants increase the cycling rate of these nutrients in boreal forests.     

The natural abundance of 15N in litter and soil profiles under six temperate tree species: N cycling depends on tree species traits and site fertility

Aims

We investigated the influence of tree species on the natural ¹⁵N abundance in forest stands under elevated ambient N deposition.

Methods

We analysed δ¹⁵N in litter, the forest floor and three mineral soil horizons along with ecosystem N status variables at six sites planted three decades ago with five European broadleaved tree species and Norway spruce.

Results

Litter δ¹⁵N and ¹⁵N enrichment factor (δ¹⁵N_litter–δ¹⁵N_soil) were positively correlated with N status based on soil and litter N pools, nitrification, subsoil nitrate concentration and forest growth. Tree species differences were also significant for these N variables and for the litter δ¹⁵N and enrichment factor. Litter from ash and sycamore maple with high N status and low fungal mycelia activity was enriched in ¹⁵N (+0.9 delta units) relative to other tree species (European beech, pedunculate oak, lime and Norway spruce) even though the latter species leached more nitrate.

Conclusions

The δ¹⁵N pattern reflected tree species related traits affecting the N cycling as well as site fertility and former land use, and possibly differences in N leaching. The tree species δ¹⁵N patterns reflected fractionation caused by uptake of N through mycorrhiza rather than due to nitrate leaching or other N transformation processes.     

The phosphorus status of northern hardwoods differs by species but is unaffected by nitrogen fertilization

Northern hardwood forests in the eastern US exhibit species-specific influences on nitrogen (N) cycling, suggesting that their phosphorus (P) cycling characteristics may also vary by species. These characteristics are increasingly important to understand in light of evidence suggesting that atmospheric N deposition has increased N availability in the region, potentially leading to phosphorus limitation. We examined how P characteristics differ among tree species and whether these characteristics respond to simulated N deposition (fertilization). We added NH₄NO₃ fertilizer (50 kg ha^?1 year^?1) to single-species plots of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.), eastern hemlock (Tsuga canadensis (L.) Carr.), American beech (Fagus grandifolia Ehrh.), and yellow birch (Betula alleghaniensis Britt.), in the Catskill Mountains, New York from 1997 to 2007. Species differences were observed in foliar, litter and root P concentrations, but all were unaffected by a cumulative N fertilization of 550 kg/ha. Similarly, measures of soil P availability and biotic P sufficiency differed by species but were unaffected by N fertilization. Results suggest species exhibit unique relationships to P as well as N cycles. We found little evidence that N fertilization leads to increased P limitation in these northern hardwood forests. However, species such as sugar maple and red oak may be sufficient in P, whereas beech and hemlock may be less sufficient and therefore potentially more sensitive to future N-stimulated P limitation.     

Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Plant and soil responses of an alpine steppe on the Tibetan Plateau to multi-level nitrogen addition

Background

Although plant growth in alpine steppes on the Tibetan Plateau has been suggested to be sensitive to nitrogen (N) addition, the N limitation conditions of alpine steppes remain uncertain.

Methods

After 2 years of fertilization with NH₄NO₃ at six rates (0, 10, 20, 40, 80 and 160 kg N ha^?1 yr^?1), the responses of plant and soil parameters as well as N₂O fluxes were measured.

Results

At the vegetation level, N addition resulted in an increase in the aboveground N pool from 0.5?±?0.1 g m^?2 in the control plots to 1.9?±?0.2 g m^?2 in the plots at the highest N input rate. The aboveground C pool, biomass N concentration, foliar δ¹⁵N, soil NO₃ ^?-N and N₂O flux were also increased by N addition. However, as the N fertilization rate increased from 10 kg N ha^?1 yr^?1 to 160 kg N ha^?1 yr^?1, the N-use efficiency decreased from 12.3?±?4.6 kg C kg N^?1 to 1.6?±?0.2 kg C kg N^?1, and the N-uptake efficiency decreased from 43.2?±?9.7 % to 9.1?±?1.1 %. Biomass N:P ratios increased from 14.4?±?2.6 in the control plots to 20.5?±?0.8 in the plots with the highest N input rate. Biomass N:P ratios, N-uptake efficiency and N-use efficiency flattened out at 40 kg N ha^?1 yr^?1. Above this level, soil NO₃ ^?-N began to accumulate. The seasonal average N₂O flux of growing season nonlinearly increased with increased N fertilization rate and linearly increased with the weighted average foliar δ¹⁵N. At the species level, N uptake responses to relative N availability were species-specific. Biomass N concentration of seven out of the eight non-legume species increased significantly with N fertilization rates, while Kobresia macrantha and the one legume species (Oxytropics glacialis) remained stable. Both the non-legume and the legume species showed significant ¹⁵N enrichment with increasing N fertilization rate. All non-legume species showed significant increased N:P ratios with increased N fertilization rate, but not the legume species.

Conclusions

Our findings suggest that the Tibetan alpine steppes might be N-saturated above a critical N load of 40 kg N ha^?1 yr^?1. For the entire Tibetan Plateau (ca. 2.57 million km²), a low N deposition rate (10 kg N ha^?1 yr^?1) could enhance plant growth, and stimulate aboveground N and C storage by at least 1.1?±?0.3 Tg N yr^?1 and 31.5?±?11.8 Tg C yr^?1, respectively. The non-legume species was N-limited, but the legume species was not limited by N.     

Relationships between tree dimension and coarse root biomass in mixed stands of European beech (Fagus sylvatica L.) and Norway spruce (Picea abies[L.] Karst.)

Relationships between tree parameters above ground and the biomass of the coarse root system were examined in six mixed spruce-beech stands in the Solling Mountain region in northwest Germany. The selected stands were located on comparable sites and covered an age range of 44 to 114 years. Coarse roots (d?\ge?2 mm) of 42 spruce and 27 beech trees were sampled by excavating the entire root system. A linear model with logarithmic transformation of the variables was developed to describe the relationship between the coarse root biomass (CRB, dry weight) and the corresponding tree diameter at breast height (DBH). The coefficients of determination (R ²) attained values between 0.92 for spruce and 0.94 for beech; the logarithmic standard deviation values were between 0.29 and 0.43. A significantly different effect of tree species on the model estimates could not be detected by an analysis of co-variance (ANCOVA). For spruce, the derived relationships were similar to those reported in previous studies, but not for beech. Biomass partitioning in the tree compartments above and below ground differs significantly between spruce (coarse root/shoot ratio 0.16±0.06) and beech (coarse root/shoot ratio 0.10±0.03) in the mixed stands. These results are similar to those given in other studies involving pure spruce and beech stands on comparable sites in the region, although the ratios of pure stands in other regions growing under different site conditions are somewhat higher. Comparing trees of the same DBH classes, root/shoot ratios of spruce are 1.2 to 3 times higher than those of beech. Dominant spruce trees (DBH>60 cm) attained the highest ratios, suppressed beech trees (DBH<10 cm) the lowest. Site conditions of varying climate and soils and interspecific tree competition are likely to affect root/shoot ratio and DBH-coarse root biomass relationships. The greater variability in beech compared with spruce indicates a high 'plasticity' and adaptability of beech carbon allocation. Thus, the derived equations are useful for biomass estimates of coarse roots involving trees of different ages in mixed stands of spruce and beech in the Solling Mountains. However, application of these relationships to stands in other regions would need further testing.