Responses of fine roots and soil N availability to short-term nitrogen fertilization in a broad-leaved Korean pine mixed forest in northeastern China

Knowledge of the responses of soil nitrogen (N) availability, fine root mass, production and turnover rates to atmospheric N deposition is crucial for understanding fine root dynamics and functioning in forest ecosystems. Fine root biomass and necromass, production and turnover rates, and soil nitrate-N and ammonium-N in relation to N fertilization (50 kg N ha(-1) year(-1)) were investigated in a temperate forest over the growing season of 2010, using sequential soil cores and ingrowth cores methods. N fertilization increased soil nitrate-N by 16% (P<0.001) and ammonium-N by 6% (P<0.01) compared to control plots. Fine root biomass and necromass in 0-20 cm soil were 13% (4.61 vs. 5.23 Mg ha(-1), P<0.001) and 34% (1.39 vs. 1.86 Mg ha(-1), P<0.001) less in N fertilization plots than those in control plots. The fine root mass was significantly negatively correlated with soil N availability and nitrate-N contents, especially in 0-10 cm soil layer. Both fine root production and turnover rates increased with N fertilization, indicating a rapid underground carbon cycling in environment with high nitrogen levels. Although high N supply has been widely recognized to promote aboveground growth rates, the present study suggests that high levels of nitrogen supply may reduce the pool size of the underground carbon. Hence, we conclude that high levels of atmospheric N deposition will stimulate the belowground carbon cycling, leading to changes in the carbon balance between aboveground and underground storage. The implications of the present study suggest that carbon model and prediction need to take the effects of nitrogen deposition on underground system into account.     

Competition for nitrogen sources between European beech (Fagus sylvatica) and sycamore maple (Acer pseudoplatanus) seedlings

To investigate the short‐term consequences of direct competition between beech and sycamore maple on root N uptake and N composition, mycorrhizal seedlings of both tree species were incubated for 4 days (i.e. beech only, sycamore maple only or both together) in an artificial nutrient solution with low N availability. On the fourth day, N uptake experiments were conducted to study the effects of competition on inorganic and organic N uptake. For this purpose, multiple N sources were applied with a single label. Furthermore, fine roots were sampled and analysed for total amino acids, soluble protein, total nitrogen, nitrate and ammonium content. Our results clearly show that both tree species were able to use inorganic and organic N sources. Uptake of inorganic and organic N by beech roots was negatively affected in the presence of the competing tree species. In contrast, the presence of beech stimulated inorganic N uptake by sycamore maple roots. Both the negative effect of sycamore maple on N uptake of beech and the positive effect of beech on N uptake of sycamore maple led to an increase in root soluble protein in beech, despite an overall decrease in total N concentration. Thus, beech compensated for the negative effects of the tree competitor on N uptake by incorporating less N into structural N components, but otherwise exhibited the same strategy as the competitor, namely, enhancing soluble protein levels in roots when grown under competition. It is speculated that enhanced enzyme activities of so far unknown nature are required in beech as a defence response to inter‐specific competition.     

Deposition and canopy exchange processes in central-German beech forests differing in tree species diversity

Atmospheric deposition is an important nutrient input to forests. The chemical composition of the rainfall is altered by the forest canopy due to interception and canopy exchange. Bulk deposition and stand deposition (throughfall plus stemflow) of Na⁺, Cl^?, K⁺, Ca²⁺, Mg²⁺, PO ₄ ^3? , SO ₄ ^2? , H⁺, Mn²⁺, Al³⁺, Fe²⁺, NH ₄ ⁺ , NO ₃ ^? and N_org were measured in nine deciduous forest plots with different tree species diversity in central Germany. Interception deposition and canopy exchange rates were calculated with a canopy budget model. The investigated forest plots were pure beech (Fagus sylvatica L.) plots, three-species plots (Fagus sylvatica, Tilia cordata Mill. or T. platyphyllos Scop. and Fraxinus excelsior L.) and five-species plots (Fagus sylvatica, T. cordata or T. platyphyllos, Fraxinus excelsior, Acer platanoides L., A. pseudoplatanus L. or A. campestre L. and Carpinus betulus L.). The interception deposition of all ions was highest in pure beech plots and was negatively related to the Shannon index. The stand deposition of K⁺, Ca²⁺, Mg²⁺ and PO ₄ ^3? was higher in mixed species plots than in pure beech plots due to higher canopy leaching rates in the mixed species plots. The acid input to the canopy and to the soil was higher in pure beech plots than in mixed species plots. The high canopy leaching rates of Mn²⁺ in pure beech plots indicated differences in soil properties between the plot types. Indeed, pH, effective cation exchange capacity and base saturation were lower in pure beech plots. This may have contributed to the lower leaching rates of K⁺, Ca²⁺ and Mg²⁺ compared to the mixed species plots. However, foliar analyses indicated differences in the ion status among the tree species, which may additionally have influenced canopy exchange. In conclusion, the nutrient input to the soil resulting from deposition and canopy leaching was higher in mixed species plots than in pure beech plots, whereas the acid input was highest in pure beech plots.     

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.     

On the significance of belowground overyielding in temperate mixed forests: separating species identity and species diversity effects

Complementary soil exploration by the root systems of coexisting tree species has been hypothesised to result in a higher root biomass of mixed forests than of monocultures but the existing evidence for a belowground diversity effect in forests is scarce and not conclusive. In a species‐rich temperate broad‐leaved forest, we analysed the fine root biomass (roots ≤ 2 mm) and necromass in 100 plots differing in tree species diversity (one to three species) and species composition (all possible combinations of five species of the genera Acer, Carpinus, Fagus, Fraxinus and Tilia) which allowed us to separate possible species diversity and species identity effects on fine root biomass. We found no evidence of a positive diversity effect on standing fine root biomass and thus of overyielding in terms of root biomass. Root necromass decreased with increasing species diversity at marginal significance. Various lines of evidence indicate significant species identity effects on fine root biomass (10–20% higher fine root biomass in plots with presence of maple and beech than in plots with hornbeam; 100% higher fine root biomass in monospecific beech and ash plots than in hornbeam plots; differences significant). Ash fine roots tended to be over‐represented in the 2‐ and 3‐species mixed plots compared to monospecific ash plots pointing at apparent belowground competitive superiority of Fraxinus in this mixed forest. Our results indicate that belowground overyielding and spatial complementarity of root systems may be the exception rather than the rule in temperate mixed forests.     

Nitrogen Addition Increases Carbon Storage in Soils, But Not in Trees, in an Eastern U.S. Deciduous Forest

Forest ecosystems in most industrialized and agricultural regions receive elevated rates of atmospheric nitrogen (N) deposition from air pollution. To evaluate the effects of excess N deposition on carbon (C) and N cycling, we experimentally added N (as NH₄NO₃) to naturally-occurring, single-species plots of five different tree species that are common in the Northern Hardwood forests of northeastern North America: sugar maple (Acer saccharum Marsh), American beech (Fagus grandifolia Ehrh.), yellow birch (Betula alleghaniensis Britton), eastern hemlock (Tsuga canadensis (L.) Carr), and northern red oak (Quercus rubra L.). The experiment was performed in the Catskill Mountains of southeastern New York State, USA, and used a paired-plot design with six replicate plots per species. After 6 years of treatment, most species showed increases in foliar N concentrations in N-treated plots, but only for maple and birch were those increases statistically significant. No significant effects of the N treatment were observed on woody biomass increment or aboveground net primary production (ANPP) for any species. In the oak plots, the N treatment increased acorn production in mast years. In the soils, the N treatment was associated with a significant decline in potential N mineralization and nitrification rates in the mineral horizon but not in the forest floor, and in the mineral horizon the effect of the N treatment varied among species. The N treatment caused a significant increase in C stock, N stock and C:N ratio in the forest floor, with the largest effect in the hemlock plots. Nitrate leaching increased significantly in treated plots compared to controls. Dissolved organic carbon (DOC) in soil solution was unaffected by the N treatment, but the variation in DOC across plots was correlated with the C stock in the forest floor. These results suggest that the ANPP of these forests is not limited by N availability, but that excess N may cause accumulations of C in the forest floor, particularly in hemlock stands, perhaps through inhibition of decomposition rates or by altering phenolic chemistry of the litter. The magnitude, and sometimes the direction of the N treatment responses varied among species, suggesting that predictions of forest responses to elevated N deposition should take into account spatial and temporal variation in tree species composition.     

Decomposition of <Superscript>15</Superscript>N-labelled beech litter and fate of nitrogen derived from litter in a beech forest

The decomposition and the fate of ¹⁵N- labelled beech litter was monitored in a beech forest (Vosges mountains, France) over 3 years. Circular plots around beech trees were isolated from neighbouring tree roots by soil trenching. After removal of the litter layer, ¹⁵N-labelled litter was distributed on the soil. Samples [labelled litter, soil (0–15 cm depths], fine roots, mycorrhizal root tips, leaves) were collected during the subsequent vegetation periods and analysed for total N and ¹⁵N concentration. Mass loss of the ¹⁵N-labelled litter was estimated using mass loss data from a litterbag experiment set up at the field site. An initial and rapid release of soluble N from the decomposing litter was balanced by the incorporation of exogenous N into the litter. Fungal N accounted for approximately 35% of the N incorporation. Over 2 years, litter N was continuously released and rates of N and mass loss were equivalent, while litter N was preferentially lost during the 3rd year. Released ¹⁵N accumulated essentially at the soil surface. ¹⁵N from the decomposing litter was rapidly (i.e. in 6 months) detected in roots and beech leaves and its level increased regularly and linearly over the course of the labelling experiment. After 3 years, about 2% of the original litter N had accumulated in the trees. ¹⁵N budgets indicated that soluble N was the main source for soil microbial biomass. Nitrogen accumulated in storage compounds was the main source of leaf N, while soil organic N was the main source of mycorrhizal N. Use of ¹⁵N-labelled beech litter as decomposing substrate allowed assessment of the fate of litter N in the soil and tree N pools in a beech forest on different time scales. Received: 3 May 1999 / Accepted: 3 January 2000     

Importance of soil nutrients in the distribution of forest communities on a large geographical scale

Aim Soil nutrient content plays a key role in plant growth through mineral nutrition and toxicity. Its impact on plant species and community distribution is studied on a large geographical scale through surrogates like topography or geology. We investigated the importance of soil pH and C:N ratio, as direct nutritional gradients, to determine, with climatic factors, the spatial distribution of plant communities over large territories. Location We studied the distribution of six beech habitats of the NATURA 2000 network throughout France (550,000 km²). Methods Models were calibrated with 2108 floristic plots classified in the NATURA 2000 system and including climatic and topographic variables and soil nutritional measurements carried out in a laboratory. Logistic regression was used to model habitat distribution according to environmental variables. Climatic layers, a digital elevation model and maps of soil pH and nitrogen content, created using plant indicator values and large floristic databases, were used to map the sites suitable for beech communities. Distribution models were evaluated with an independent set of 2091 phytosociological plots. Results pH and nitrogen supply were the key distribution drivers for four of the six beech communities on a national scale. Their use in the distribution models distinguished within homogeneous climatic territories a gradient of nutritional conditions from acidic areas, suitable for nutrient‐poor beech communities, to calcareous areas suitable for nutrient‐rich ones. Predicted maps of beech habitats fit the spatial distribution of validation plots. Main conclusions Soil pH and nitrogen supply strongly improve predictions of forest community distribution carried out with climatic variables on a broad geographical scale. They allow delineation of areas with nutritional conditions suitable for each community, as well as the realization of predictive high‐resolution maps over large areas useful for sustainable and conservation management. Nomenclature Tutin & Heywood (2001 ) Flora Europaea. Cambridge University Press, Cambridge.     

Direct and indirect effects of nitrogen deposition on species composition change in calcareous grasslands

Atmospheric nitrogen (N) deposition has been identified as a major threat to biodiversity, but field surveys of its effects have rarely focussed on sites which are actively managed to maintain characteristic species. We analysed permanent quadrat data from 106 plots in nature reserves on calcareous grassland sites in the United Kingdom collected during a survey between 1990 and 1993 and compared the data with the results from resurvey of 48 of these plots between 2006 and 2009. N deposition showed no significant spatial association with species richness, species diversity, or the frequency of species adapted to low nutrient conditions in the 1990–1993 dataset. However, temporal analysis showed that N deposition was significantly associated with changes in Shannon diversity and evenness. In plots with high rates of N deposition, there was a decrease in species diversity and evenness, a decline in the frequency of characteristic calcareous grassland species, and a lower number of rare and scarce species. As all sites had active management to maintain a high diversity and characteristic species, our results imply that even focussed management on nature conservation objectives cannot prevent adverse effects of high rates of N deposition. Structural equation modelling was used to compare different causal mechanisms to explain the observed effects. For change in Shannon diversity, direct effects of N deposition were the dominant mechanism and there was an independent effect of change in grazing intensity. In contrast, for change in herb species number, indirect effects on soil acidity, linked to both N and S deposition, were more important than direct effects of N deposition.     

Do increased summer precipitation and N deposition alter fine root dynamics in a Mojave Desert ecosystem?

Climate change is expected to impact the amount and distribution of precipitation in the arid southwestern United States. In addition, nitrogen (N) deposition is increasing in these regions due to increased urbanization. Responses of belowground plant activity to increases in soil water content and N have shown inconsistent patterns between biomes. In arid lands, plant productivity is limited by water and N availability so it is expected that changes in these factors will affect fine root dynamics. The objectives of this study were to quantify the effects of increased summer precipitation and N deposition on fine root dynamics in a Mojave Desert ecosystem during a 2‐year field experiment using minirhizotron measurements. Root length density, production, and mortality were measured in field plots in the Mojave Desert receiving three 25 mm summer rain events and/or 40 kg N ha^?1 yr^?1. Increased summer precipitation and N additions did not have an overall significant effect on any of the measured root parameters. However, differences in winter precipitation resulting from interannual variability in rainfall appeared to affect root parameters with root production and turnover increasing following a wet winter most likely due to stimulation of annual grasses. In addition, roots were distributed more deeply in the soil following the wet winter. Root length density was initially higher under canopies compared to canopy interspaces, but converged toward the end of the study. In addition, roots tended to be distributed more deeply into the soil in canopy interspace areas. Results from this study indicated that increased summer precipitation and N deposition in response to climate change and urbanization are not likely to affect fine root dynamics in these Mojave Desert ecosystems, despite studies showing aboveground plant physiological responses to these environmental perturbations. However, changes in the amount and possibly distribution of winter precipitation may affect fine root dynamics.     

增温对川西北亚高山高寒草甸植物群落碳、氮含量的影响

采用开顶式生长室（OTC）模拟增温实验,研究了川西北亚高山草甸植物群落碳、氮含量对温度升高的响应。由于OTC的增温作用,在整个生长季内,地温（15 cm）、地表温度和气温(30 cm)的平均值在OTC内比对照样地分别高0.28、0.46和1.4℃,OTC内土壤相对含水量也明显减少,低于对照样地5.49%。受增温及土壤含水量减少的影响,一年后,植物群落的生物量积累和碳、氮含量发生了明显的改变。除10月份OTC内地上鲜体生物量略高于对照样地外,OTC内地上鲜体生物量和根系生物量与对照样地相比,都出现了不同程度的减少;OTC内植物群落地上活体的碳浓度在整个生长季高于对照样地,而氮浓度低于对照样地;OTC内植物群落地下活根的碳浓度在整个生长季高于对照样地,并且在8月份统计检验显著,而氮浓度却低于对照样地;OTC内植物碳库在整个生长季较对照样地有不同程度的增加,增幅范围为0.90%～5.65%,而OTC内植物氮库较对照样地有不同程度的减少,减幅范围0.40%～1.28%。     

Interactive effects of elevated CO2 and nitrogen deposition on fatty acid molecular and isotope composition of above‐ and belowground tree biomass and forest soil fractions

Atmospheric carbon dioxide (CO₂) and reactive nitrogen (N) concentrations have been increasing due to human activities and impact the global carbon (C) cycle by affecting plant photosynthesis and decomposition processes in soil. Large amounts of C are stored in plants and soils, but the mechanisms behind the stabilization of plant‐ and microbial‐derived organic matter (OM) in soils are still under debate and it is not clear how N deposition affects soil OM dynamics. Here, we studied the effects of 4 years of elevated (¹³C‐depleted) CO₂ and N deposition in forest ecosystems established in open‐top chambers on composition and turnover of fatty acids (FAs) in plants and soils. FAs served as biomarkers for plant‐ and microbial‐derived OM in soil density fractions. We analyzed above‐ and belowground plant biomass of beech and spruce trees as well as soil density fractions for the total organic C and FA molecular and isotope (δ¹³C) composition. FAs did not accumulate relative to total organic C in fine mineral fractions, showing that FAs are not effectively stabilized by association with soil minerals. The δ¹³C values of FAs in plant biomass increased under high N deposition. However, the N effect was only apparent under elevated CO₂ suggesting a N limitation of the system. In soil fractions, only isotope compositions of short‐chain FAs (C₁₆₊₁₈) were affected. Fractions of ‘new’ (experimental‐derived) FAs were calculated using isotope depletion in elevated CO₂ plots and decreased from free light to fine mineral fractions. ‘New’ FAs were higher in short‐chain compared to long‐chain FAs (C_20?30), indicating a faster turnover of short‐chain compared to long‐chain FAs. Increased N deposition did not significantly affect the quantity of ‘new’ FAs in soil fractions, but showed a tendency of increased amounts of ‘old’ (pre‐experimental) C suggesting that decomposition of ‘old’ C is retarded by high N inputs.     

Factors controlling decomposition rates of fine root litter in temperate forests and grasslands

Background and aims

Fine root decomposition contributes significantly to element cycling in terrestrial ecosystems. However, studies on root decomposition rates and on the factors that potentially influence them are fewer than those on leaf litter decomposition. To study the effects of region and land use intensity on fine root decomposition, we established a large scale study in three German regions with different climate regimes and soil properties. Methods In 150 forest and 150 grassland sites we deployed litterbags (100 μm mesh size) with standardized litter consisting of fine roots from European beech in forests and from a lowland mesophilous hay meadow in grasslands. In the central study region, we compared decomposition rates of this standardized litter with root litter collected on-site to separate the effect of litter quality from environmental factors.

Results

Standardized herbaceous roots in grassland soils decomposed on average significantly faster (24?±?6 % mass loss after 12 months, mean ± SD) than beech roots in forest soils (12?±?4 %; p?Conclusions Grasslands, which have higher fine root biomass and root turnover compared to forests, also have higher rates of root decomposition. Our results further show that at the regional scale fine root decomposition is influenced by environmental variables such as soil moisture, soil temperature and soil nutrient content. Additional variation is explained by root litter quality.     

Nitrate nutrition ofDeschampsia flexuosa (L.) Trin. in relation to nitrogen deposition in Sweden

Summary Current and maximally induced nitrate reductase activity (NRA), total-N, nitrate, K, P, Ca, Mg, Mo and sucrose in leaves ofDeschampsia flexuosa was measured three times during the vegetation period in forests along a deposition gradient (150 km) in south Sweden, in north Sweden where the nitrogen deposition is considerably lower, and at heavily N-fertilized plots. In addition, the interaction between nitrogen nutrition and light was studied along transects from clearings into forest in both south and north Sweden. Plants from sites with high nitrogen deposition had elevated current NRA compared to plants from less polluted sites, indicating high levels of available soil nitrate at the former. Current NRA and total N concentration in grass from sites with high deposition resembled those found at heavily N-fertilized plots. Under such circumstances, the ratio current NRA: maximally induced NRA as well as the concentration of nitrate was high, while the concentration of sucrose was low. This suggests that the grass at these sites was already utilizing a large portion of its capacity to assimilate nitrate. Light was found to play an important role in the assimilation of nitrate; leaf concentration of sucrose was found to be negatively correlated with both nitrate and total N. Consequently, grass growing under dense canopies in south Sweden is not able to dilute N by increasing growth. The diminished capacity of the grass to assimilate nitrate will increase leaching losses of N from forests approaching N saturation.     

Evaluation of different approaches for modelling individual tree seedling height growth

We compared different approaches for modelling height growth of individual beech seedlings in a controlled factorial experiment as well as in field data from naturally regenerated beech seedlings under the canopy of overstorey mature beech trees. Several competition indices, a model of overstorey fine root density, relative photosynthetically active radiation (PAR) values, and soil water values were used in these approaches. In the factorial experiment relative PAR and soil water content were measured and used for the prediction of seedlings height growth. In the field experiment this was done by using relative PAR and estimated fine root biomass as a surrogate for below ground resource availability. The latter approach was compared with a model where we used various competition indices representing the impact of overstorey trees on beech seedlings. Our results suggested that (1) models which combine resource based growth functions are suitable for the prediction of individual height growth of beech seedlings. Resource based models offer the opportunity to investigate on the independent multiplicative effect of irradiance and water supply and their interactions on tree seedlings. It was (2) shown that a combined model could be used not only to predict individual height growth of beech seedlings in a controlled experiment but also in the field. The model parameters of a pure light response function for the controlled factorial experiment are comparable to those obtained in the field study. The results showed (3) that the precision of predicting beech seedlings height growth is comparable between the model types tested within this study. Approximately half of the observed variation in seedlings relative height growth rate could be explained. However, the simple competition index approach provides no information on the environmental factors constraining tree seedlings growth; whereas the multiplicative combined models can be used to get a better understanding of growth dynamics in the field.     

根系去除改变了毛竹林土壤酶活性对氮磷添加的响应

亚热带是我国氮和磷沉降的热点地区,森林生态系统碳（C）、氮（N）、磷（P）循环对N和P沉降的响应受到广泛关注。根系作为森林土壤碳和养分持续供给的重要来源,其输入变化在N和P沉降下通过影响土壤酶活性进而调控土壤碳和养分循环过程。以毛竹林为研究对象,选择硝酸铵和磷酸二氢钠分别进行N和P添加,共设置4个处理：对照、N添加、P添加和N+P添加,并结合壕沟处理,探究毛竹林土壤C、N、P循环相关酶活性对氮磷添加和根系去除的响应,并分析它们与土壤和细根化学性质之间的关系。结果表明：土壤C、N循环相关酶活性整体上对根系去除的响应比氮磷添加更敏感,这主要是因为根系去除显著降低土壤全氮和铵态氮含量,但短期氮磷添加并未显著改变调控土壤酶活性分异的土壤有机碳、全氮和铵态氮含量。与C、N循环相关酶活性不同,土壤P循环相关酶对根系去除和磷添加处理均呈现负响应,这可能是因为阻断根系P吸收和补充土壤P元素降低了微生物的P开采作用。但研究区土壤微生物仍受到P限制作用,这是因为在P缺乏毛竹林生态系统,根系比微生物具有更强的P吸收利用能力。研究结果强调了不同功能土壤酶活性对土壤养分添加和根系输入变化响应的敏感性差异,为预测低P毛竹林生态系统土壤C和养分循环提供依据。     

干热河谷失稳性坡面植物碳氮磷化学计量特征对土壤性质的响应

以干热河谷阴坡和阳坡典型失稳性坡面不同区段(稳定区、失稳区和堆积区)的草本植物为研究对象,分析植物地上和地下碳(C)、氮(N)、磷(P)含量及其化学计量特征对土壤性质的响应机制。结果表明:失稳性坡面植物地上C含量(320.59 g·kg^-1)、N含量(12.15 g·kg^-1)和C/P(25.37)均显著高于地下(分别为254.01 g·kg^-1、6.12 g·kg^-1、13.43),C/N则表现为地下(43.09)显著高于地上(31.90)。阳坡植物地上和地下C含量、N/P均表现为稳定区和失稳区显著高于堆积区,N含量则表现为失稳区显著高于堆积区;阴坡植物地上N、P含量及地下P含量均表现为堆积区显著高于稳定区和失稳区,地下C含量则反之。植物地上生长主要受P限制,根系生长主要受N限制并随区段向下呈逐渐增加趋势;阴坡植物生长受P的限制程度高于阳坡,N限制弱于阳坡。土壤含水量(SWC)是影响植物地上C、N、P含量变化的重要因子,影响值分别为28.8%、20.8%、19.9%,土壤有机碳(SOC)则显著影响植物地下C、N含量,影响值分别为49.5%、22.1%,植物地下N含量的变化主要受土壤pH值的影响(24.3%)。坡向、区段及土壤因子均显著影响植物养分的分配,SWC和SOC是主要影响因子,同时还受土壤pH值的影响。     

Decomposition of roots and twigs: Effects of wood type (beech and ash), diameter,site of exposure and macrofauna exclusion

Carbon loss and nitrogen dynamics in beech roots (Fagus sylvatica L.), beech twigs and ash roots (Fraxinus excelsior L.) of 0–3, 3–10 and 10–40 mm diameter were investigated during 36 months of exposure in litter bags of 1 and 4 mm mesh. Four experiments were set up: (1) Beech and ash roots (three size classes) were placed in a soil depth of ca 5 cm in a beechwood on limestone; (2) beech twigs (three size classes) were placed on the soil surface of the beechwood; (3) beech roots (3–10 mm) were placed on the soil surface of the beechwood: (4) beech twigs (3–10 mm) were placed on the soil surface of four sites representing different stages of secondary succession (wheat field, 13 year old fallow, ca 50 year old fallow, beechwood). Ash roots generally lost more C than beech roots. Loss in C of ash roots was similar for each of the size classes, whereas in beech roots and beech twigs C loss was in the order large roots > medium roots > small roots. Beech roots (3–10 mm) placed on the soil surface lost considerably less C than beech twigs (3–10 mm). Decomposition of beech twigs varied among ecosystems but generally did not follow clear patterns with successional stages. The fit of linear vs exponential models of decay is compared and in most materials exponential models fitted the data better. In each of the wood materials an accumulation of N occurred. Irrespective of wood type, root and twig diameter, mineralization of N of wood materials placed in the beechwood started uniformly after 12 months. Multiple regression analysis indicated a negative relationship between initial N content and C loss in beech roots and twigs but not in ash roots. The analysis also indicated a significant influence of the degree of white rot and of the amount of mineral soil deposited in the litter bags on C loss of certain wood materials. Generally, mesh size affected C loss and N dynamics only slightly, which is attributed to the comparatively short exposure time.     

Chronic N deposition alters root respiration‐tissue N relationship in northern hardwood forests

Specific root respiration rates typically increase with increasing tissue N concentration. As a result, it is often assumed that external factors inducing greater root N concentration, such as chronic N deposition, will lead to increased respiration rates. However, enhanced N availability also alters root biomass, making the ecosystem‐level consequences on whole‐root‐system respiration uncertain. The objective of this study was to determine the effects of chronic experimental N deposition on root N concentrations, specific respiration rates, and biomass for four northern hardwood forests in Michigan. Three of the six measurement plots at each location have received experimental N deposition (3 g ‐N m^?2 yr^?1) since 1994. We measured specific root respiration rates and N concentrations of roots from four size classes (<0.5, 0.5–1, 1–2, and 2–10 mm) at three soil depths (0–10, 10–30, and 30–50 cm). Root biomass data for the same size classes and soil depths was used in combination with specific respiration rates to assess the response of whole‐root‐system respiration. Root N and respiration rate were greater for smaller diameter roots and roots at shallow depths. In addition, root N concentrations were significantly greater under chronic N deposition, particularly for larger diameter roots. Specific respiration rates and root biomass were unchanged for all depths and size classes, thus whole‐root‐system respiration was not altered by chronic N deposition. Higher root N concentrations in combination with equivalent specific respiration rates under experimental N deposition resulted in a lower ratio of respiration to tissue N. These results indicate that relationships between root respiration rate and N concentration do not hold if N availability is altered significantly. For these forests, use of the ambient respiration to N relationship would over‐predict actual root system respiration for the chronic N deposition treatment by 50%.     

Nitrogen supply affects arbuscular mycorrhizal colonization of Artemisia vulgaris in a phosphate-polluted field site

Root colonization by arbuscular mycorrhizal fungi (AMF) was investigated in industrially polluted grassland characterized by exceptionally high phosphorus levels (up to 120 g kg(-1) soil). Along a pollution-induced nitrogen gradient, soil and tissue element concentrations of Artemisia vulgaris plants and their mycorrhizal status were determined. Additionally, we compared mycorrhization rates and above-ground biomass of A. vulgaris at N-fertilized and control plots in the N-poor area. Despite high soil and tissue P concentrations, plants from N-deficient plots, which were characterized by low tissue N concentrations and N : P ratios, were strongly colonized by AMF, whereas at a plot with comparable P levels, but higher soil and plant N concentrations and N : P ratios, mycorrhization rates were significantly lower. Correlation analyses revealed a negative relationship between percentage root colonization of A. vulgaris by AMF and both tissue N concentration and N : P ratio. Accordingly, in the fertilization experiment, control plants had higher mycorrhization rates than N-fertilized plants, whereas the species attained higher biomass at N-fertilized plots. The results suggest that N deficiency stimulates root colonization by AMF in this extraordinarily P-rich field site.