Global soil nitrogen cycle pattern and nitrogen enrichment effects: Tropical versus subtropical forests

Tropical and subtropical forest biomes are a main hotspot for the global nitrogen (N) cycle. Yet, our understanding of global soil N cycle patterns and drivers and their response to N deposition in these biomes remains elusive. By a meta-analysis of 2426-single and 161-paired observations from 89 published ¹⁵ N pool dilution and tracing studies, we found that gross N mineralization (GNM), immobilization of ammonium ($I_{{\mathrm{NH}}_4}$) and nitrate ($I_{{\mathrm{NO}}_3}$), and dissimilatory nitrate reduction to ammonium (DNRA) were significantly higher in tropical forests than in subtropical forests. Soil N cycle was conservative in tropical forests with ratios of gross nitrification (GN) to $I_{{\mathrm{NH}}_4}$ (GN/$I_{{\mathrm{NH}}_4}$) and of soil nitrate to ammonium (NO₃⁻/NH₄⁺) less than one, but was leaky in subtropical forests with GN/$I_{{\mathrm{NH}}_4}$ and NO₃⁻/NH₄⁺ higher than one. Soil NH₄⁺ dynamics were mainly controlled by soil substrate (e.g., total N), but climatic factors (e.g., precipitation and/or temperature) were more important in controlling soil NO₃⁻ dynamics. Soil texture played a role, as GNM and $I_{{\mathrm{NH}}_4}$ were positively correlated with silt and clay contents, while $I_{{\mathrm{NO}}_3}$ and DNRA were positively correlated with sand and clay contents, respectively. The soil N cycle was more sensitive to N deposition in tropical forests than in subtropical forests. Nitrogen deposition leads to a leaky N cycle in tropical forests, as evidenced by the increase in GN/$I_{{\mathrm{NH}}_4}$, NO₃⁻/NH₄⁺, and nitrous oxide emissions and the decrease in $I_{{\mathrm{NO}}_3}$ and DNRA, mainly due to the decrease in soil microbial biomass and pH. Dominant tree species can also influence soil N cycle pattern, which has changed from conservative in deciduous forests to leaky in coniferous forests. We provide global evidence that tropical, but not subtropical, forests are characterized by soil N dynamics sustaining N availability and that N deposition inhibits soil N retention and stimulates N losses in these biomes.     

Effects of root diameter and root nitrogen concentration on in situ root respiration among different seasons and tree species

Knowledge of root respiration is a prerequisite for a better understanding of ecosystem carbon budget and carbon allocation. However, there are not many relevant data in the literature on direct measurements of in situ root respiration by root chamber method. Furthermore, few studies have been focused on the effects of root diameter (D _r) and root nitrogen concentration (N _r) on in situ root respiration among different seasons and tree species. To address these goals, we used a simplified root-chamber system to measure in situ root respiration rates of Acacia crassicarpa and Eucalyptus urophylla in subtropical plantations of south China. We found that the species and season variation in root respiration were affected by D _r and N _r. Also, the root respiration per unit dry mass (R _r, nmol CO₂ g⁻¹ s⁻¹) and root respiration per unit N (R _n, nmol CO₂ g N⁻¹ s⁻¹) were affected by D _r and N _r. The R _r, R _n, N _r and soil temperature sensitivity (Q ₁₀) of R _r for the two species significantly decreased with an increase of D _r. The R _r of the two species showed significant an inter-seasonal and diurnal pattern, and this trend decreased with increasing D _r. Both the R _r and Q ₁₀ of the two species increased with increasing N _r. The D _r and N _r explained 54 and 52% of the observed variation in R _r for A. crassicarpa, and 65 and 70% for E. urophylla. The R _r, N _r, and Q ₁₀ of A. crassicarpa were significantly higher than those of E. urophylla. Our results indicated that root respiration was dependent on D _r and N _r, and this dependence varied with season and plant species.     

不同温度条件下杉木、桤木和火力楠细根分解对土壤活性有机碳的影响

通过室内培养试验,研究了不同温度(9 ℃、14 ℃、24 ℃和28 ℃)条件下桤木、杉木和火力楠细根分解对土壤活性有机碳的影响.结果表明,不同树种细根的分解率不同,树种间差异显著,大小依次为火力楠>桤木>杉木.细根分解率随着培养温度的增加而增大,随着培养时间的延长而降低.添加细根的种类、培养温度和培养时间均对实验系统中土壤微生物碳和水溶性有机碳的含量产生影响.3个树种细根分解使土壤微生物碳和水溶性有机碳含量显著高于对照,大小依次为火力楠>桤木>杉木>对照; 培养中期以及中等培养温度条件下细根分解对应着较高的土壤微生物碳和水溶性有机碳含量.细根分解对土壤易氧化碳含量无显著影响.     

Fine root decomposition rates do not mirror those of leaf litter among temperate tree species

Elucidating the function of and patterns among plant traits above ground has been a major research focus, while the patterns and functioning of belowground traits remain less well understood. Even less well known is whether species differences in leaf traits and their associated biogeochemical effects are mirrored by differences in root traits and their effects. We studied fine root decomposition and N dynamics in a common garden study of 11 temperate European and North American tree species (Abies alba, Acer platanoides, Acer pseudoplatanus, Carpinus betulus, Fagus sylvatica, Larix decidua, Picea abies, Pseudotsuga menziesii, Quercus robur, Quercus rubra and Tilia cordata) to determine whether leaf litter and fine root decomposition rates are correlated across species as well as which species traits influence microbial decomposition above versus below ground. Decomposition and N immobilization rates of fine roots were unrelated to those of leaf litter across species. The lack of correspondence of above- and belowground processes arose partly because the tissue traits that influenced decomposition and detritus N dynamics different for roots versus leaves, and partly because influential traits were unrelated between roots and leaves across species. For example, while high hemicellulose concentrations and thinner roots were associated with more rapid decomposition below ground, low lignin and high Ca concentrations were associated with rapid aboveground leaf decomposition. Our study suggests that among these temperate trees, species effects on C and N dynamics in decomposing fine roots and leaf litter may not reinforce each other. Thus, species differences in rates of microbially mediated decomposition may not be as large as they would be if above- and belowground processes were working in similar directions (i.e., if faster decomposition above ground corresponded to faster decomposition below ground). Our results imply that studies that focus solely on aboveground traits may obscure some of the important mechanisms by which plant species influence ecosystem processes.     

Stream carbon and nitrogen supplements during leaf litter decomposition: contrasting patterns for two foundation species

Leaf litter decomposition plays a major role in nutrient dynamics in forested streams. The chemical composition of litter affects its processing by microorganisms, which obtain nutrients from litter and from the water column. The balance of these fluxes is not well known, because they occur simultaneously and thus are difficult to quantify separately. Here, we examined C and N flow from streamwater and leaf litter to microbial biofilms during decomposition. We used isotopically enriched leaves (¹³C and ¹⁵N) from two riparian foundation tree species: fast-decomposing Populus fremontii and slow-decomposing Populus angustifolia, which differed in their concentration of recalcitrant compounds. We adapted the isotope pool dilution method to estimate gross elemental fluxes into litter microbes. Three key findings emerged: litter type strongly affected biomass and stoichiometry of microbial assemblages growing on litter; the proportion of C and N in microorganisms derived from the streamwater, as opposed to the litter, did not differ between litter types, but increased throughout decomposition; gross immobilization of N from the streamwater was higher for P. fremontii compared to P. angustifolia, probably as a consequence of the higher microbial biomass on P. fremontii. In contrast, gross immobilization of C from the streamwater was higher for P. angustifolia, suggesting that dissolved organic C in streamwater was used as an additional energy source by microbial assemblages growing on slow-decomposing litter. These results indicate that biofilms on decomposing litter have specific element requirements driven by litter characteristics, which might have implications for whole-stream nutrient retention.     

Effect of tree species and crown pruning on root length and soil water content in semi-arid agroforestry

Soil cores were taken to estimate root length prior to transplanting and after 60 days growth of a dry season sorghum crop in an agroforestry experiment in a semi-arid region of north-east Nigeria. The experiment compared sorghum grown alone and with two tree species (Acacia nilotica subsp adstringens and Prosopis juliflora) and one management treatment (pruning of tree crowns). Data on soil water content were collected from 6 days before and 20, 60 and 110 days after sorghum transplanting. The main findings were: (i) Per unit root length, A. nilotica had a more negative effect on sorghum above and below ground than P. juliflora. This appeared to be correlated with greater rates of water extraction from layers of soil shared with crop roots; (ii) Crown pruning substantially reduced the competitive effect of P. juliflora on crop yield but did not affect the impact of A. nilotica on intercropped sorghum. Since the impact of pruning on tree-crop competition varies with species, tree species selection and management will be a key factor in determining the feasibility of dryland agroforestry systems.     

Intrinsic effects of species on leaf litter and root decomposition: a comparison of temperate grasses from North and South America

A simple model of water uptake by vegetation is used to aid the discrimination of plant water sources determined with isotope data. In the model, water extracted from different soil depths depends on the leaf–soil potential difference, a root distribution function and a lumped hydraulic conductance parameter. Measurements of plant transpiration rate, and soil and leaf water potentials are used to estimate the value of the conductance parameter. Isotopic ratios in soil water and xylem are then used to constrain the root distribution. The model is applied to field measurements of transpiration, leaf water potential and ¹⁸O composition of xylem water on Corymbia clarksoniana, Lophostemon suaveolens, Eucalpytus platyphylla and Melaleuca viridiflora, and soil water potential and ¹⁸O composition of soil water to 8.5 m depth, in an open woodland community, Pioneer Valley, North Queensland. Estimates of the water uptake from various depths below the surface are determined for each species. At the time of sampling, the proportion of groundwater extracted by the trees ranged from 100% for C. clarksoniana to <15% for L. suaveolens and E. platyphylla. The advantages of the model over the traditional approach to determining sources of water used by plants using isotope methods are that it: (1) permits more quantitative assessments of the proportion of water sourced from different depths, (2) can deal with gradational soil water isotope profiles (rather than requiring distinct values for end-members), and (3) incorporates additional data on plant water potentials and is based on simple plant physiological processes.     

Context-dependent tree species effects on soil nitrogen transformations and related microbial functional genes

Although it is generally accepted that tree species can influence nutrient cycling processes in soils, effects are not consistently found, nor are the mechanisms behind tree species effects well understood. Our objectives were to gain insights into the mechanism(s) underlying the effects of tree species on soil nitrogen cycling processes, and to determine the consistency of tree species effects across sites. We compared N cycling in soils beneath six tree species (ash, sycamore maple, lime, beech, pedunculate oak, Norway spruce) in common garden experiments planted 42 years earlier at three sites in Denmark with distinct land-use histories (forest and agriculture). We measured: (1) net and gross rates of N transformations using the ¹⁵N isotope pool-dilution method, (2) soil microbial community composition through qPCR of fungal ITS, bacterial and archaeal 16S, and (3) abundance of functional genes associated with N cycling processes—for nitrification the archaeal and bacterial ammonia-monooxygenase genes (amoA AOA and amoA AOB, respectively) and for denitrification, the nitrate reductase genes nirK and nirS. Carbon concentrations were higher in soils under spruce than under broadleaves, so N transformation rates were standardized per g soil C. Soil NH₄⁺ parameters (gross ammonification, gross NH₄⁺ consumption, net ammonification (net immobilization in this case), and NH₄⁺ concentrations, per g C) were all lowest in soils under spruce. Soils under spruce also had the lowest gene abundance of bacteria, bacterial:fungal ratio, denitrifying microorganisms, ammonia-oxidizing archaea and ammonia-oxidizing bacteria. Differences in N-cycling processes and organisms among the five broadleaf species were smaller. The ‘spruce effect’ on soil microbes and N transformations appeared to be driven by its acidifying effect on soil and tighter N cycling, which occurred at the previously forested sites but not at the previously agricultural site. We conclude that existing characteristics of soils, including those resulting from previous land use, mediate the effects of tree species on the soil microbial communities and activities that determine rates of N-cycling processes.     

Fine root architecture, morphology, and biomass of different branch orders of two Chinese temperate tree species

We have limited understanding of architecture and morphology of fine root systems in large woody trees. This study investigated architecture, morphology, and biomass of different fine root branch orders of two temperate tree species from Northeastern China—Larix gmelinii Rupr and Fraxinus mandshurica Rupr —by sampling up to five fine root branch orders three times during the 2003 growing season from two soil depths (i.e., 0–10 and.10–20 cm). Branching ratio (R _b) differed with the level of branching: R _b values from the fifth to the second order of branching were approximately three in both species, but markedly higher for the first two orders of branching, reaching a value of 10.4 for L. gmelinii and 18.6 for F. mandshurica. Fine root diameter, length, SRL and root length density not only had systematic changes with root order, but also varied significantly with season and soil depth. Total biomass per order did not change systematically with branch order. Compared to the second, third and/or fourth order, the first order roots exhibited higher biomass throughout the growing season and soil depths, a pattern related to consistently higher R _b values for the first two orders of branching than the other levels of branching. Moreover, the differences in architecture and morphology across order, season, and soil depth between the two species were consistent with the morphological disparity between gymnosperms and angiosperms reported previously. The results of this study suggest that root architecture and morphology, especially those of the first order roots, should be important for understanding the complexity and multi-functionality of tree fine roots with respect to root nutrient and water uptake, and fine root dynamics in forest ecosystems.     

The effects of clipping and soil moisture on leaf and root morphology and root respiration in two temperate and two tropical grasses

Numerous studies have explored the effect of environmental conditions on a number of plant physiological and structural traits, such as photosynthetic rate, shoot versus root biomass allocation, and leaf and root morphology. In contrast, there have been a few investigations of how those conditions may influence root respiration, even though this flux can represent a major component of carbon (C) pathway in plants. In this study, we examined the response of mass-specific root respiration (μmol CO₂ g⁻¹ s⁻¹), shoot and root biomass, and leaf photosynthesis to clipping and variable soil moisture in two C₃ (Festuca idahoensis Elmer., Poa pratensis L.) and two C₄ (Andropogon greenwayi Napper, and Sporobolus kentrophyllus K. Schum.) grass species. The C₃ and C₄ grasses were collected in Yellowstone National Park, USA and the Serengeti ecosystem, Africa, respectively, where they evolved under temporally variable soil moisture conditions and were exposed to frequent, often intense grazing. We also measured the influence of clipping and soil moisture on specific leaf area (SLA), a trait associated with moisture conservation, and specific root length (SRL), a trait associated with efficiency per unit mass of soil resource uptake. Clipping did not influence any plant trait, with the exception that it reduced the root to shoot ratio (R:S) and increased SRL in P. pratensis. In contrast to the null effect of clipping on specific root respiration, reduced soil moisture lowered specific root respiration in all four species. In addition, species differed in how leaf and root structural traits responded to lower available soil moisture. P. pratensis and A. greenwayi increased SLA, by 23% and 33%, respectively, and did not alter SRL. Conversely, S. kentrophyllus increased SRL by 42% and did not alter SLA. F. idahoensis responded to lower available soil moisture by increasing both SLA and SRL by 38% and 33%, respectively. These responses were species-specific strategies that did not coincide with photosynthetic pathway (C₃/C₄) or growth form. Thus, mass-specific root respiration responded uniformly among these four grass species to clipping (no effect) and increased soil moisture stress (decline), whereas the responses of other traits (i.e., R:S ratio, SLA, SRL) to the treatments, especially moisture availability, were species-specific. Consequently, the effects of either clipping or variation in soil moisture on the C budget of these four different grasses species were driven primarily by the plasticity of R:S ratios and the structural leaf and root traits of individual species, rather than variation in the response of mass-specific root respiration.     

Macro- and micronutrient effects on decomposition of leaf litter from two tropical tree species: inferences from a short-term laboratory incubation

While a large number of studies have investigated the effects of macronutrients such as nitrogen (N) or phosphorus (P) on litter decomposition, recent studies suggest that micronutrients including zinc (Zn) may also limit decomposition rates. Our goal was to compare the effects of nutrient addition on decomposition of two leaf litter types from tropical dry forest trees in a short-term laboratory microcosm experiment. Single nutrients (N, P, Zn, potassium, magnesium, and nickel) were applied to leaf litter in solution at low or high concentrations (to mimic in situ availability or to alleviate nutrient limitation, respectively), and decomposition was assessed as final mass remaining and carbon dioxide mineralization. Both mass remaining and CO₂ mineralization were affected by nutrient identity and concentration, and these effects varied by species. In general, P and Zn addition increased decomposition, Mg and N inhibited it, and K and Ni had no significant effects. Future studies should consider the interactions between decomposition processes, decomposer communities, and a wider range of macro- and micronutrients.     

Effects of tree species on soil organic carbon density: A common garden experiment of five temperate tree species

Aims Forest trees alter litter inputs, turnover and rhizospheric activities, modify soil physical, chemical and biological properties, and consequently affect soil organic carbon (SOC) storage and carbon sink strength. That how to select appropriate tree species in afforestation, reforestation and management practices is critical to enhancing forest carbon sequestration. The objective of this study was to determine the effects of tree species on SOC density and vertical distributions.Methods A common garden experiment with the same climate, soil, and management history was established in Maoershan Forest Ecosystem Station, Northeast China, in 2004. The experimental design was a completely randomized arrangement with twenty 25 m × 25 m plots, consisting of monocultures of five tree species, including white birch (Betula platyphylla), Manchurian walnut (Juglans mandshurica), Manchurian ash (Fraxinus mandshurica), Dahurian larch (Larix gmelinii), and Mongolian pine (Pinus sylvestris var. mongolica), each with four replicated plots. A decade after the establishment (2013-2014), we measured carbon density and related factors (i.e., bulk density, total nitrogen concentration, microbial biomass carbon, microbial biomass nitrogen, pH value) in soils of the 0-40 cm depth for these monocultures. Important findings Results showed that tree species significantly influenced the SOC density in the 0-40 cm depth (p < 0.05). SOC density in the 0-10 cm depth varied from 2.79 to 3.08 kg·m^-2, in the order of walnut > ash> birch > larch > pine, in the 10-20 cm depth from 1.56 to 2.19 kg·m^-2, in the order of pine > walnut > ash > birch > larch, in the 20-30 cm depth from 1.17 to 2.10 kg·m^-2, and in the 20-40 cm depth from 0.84 to 1.43 kg·m^-2. The greatest SOC density occurred in the birch stands in the 20-40 cm depth. The vertical distributions of SOC density varied with tree species. The percentage of SOC in the 0-10 cm depth over the total SOC in the soil profile was significantly higher in the walnut and larch stands than in others, while the percentage of SOC in the 20-40 cm depth over the total SOC was highest in the birch stands. SOC concentration and soil bulk density differed significantly among the stands of different tree species, and were negatively correlated. SOC density was positively correlated with soil microbial biomass and soil pH in the walnut, ash, and larch stands, and with total nitrogen density in all the stands. We conclude that tree species modifies soil properties and microbial activity, thereby influencing SOC density, and that different patterns of vertical distributions of SOC density among monocultures of different tree species may be attributed to varying SOC controls at each soil depth.     

东北温带森林常见树种粗根分解过程及调控因子

粗根是森林生态系统中重要的碳库和养分库,对生态系统的碳和养分循环起着重要的作用。但目前人们对于影响粗根分解的主要因素以及粗根分解模式的研究较少。采用埋袋法对东北温带森林常见的10个树种（黄檗、胡桃楸、水曲柳、色木槭、红松、落叶松、白桦、春榆、紫缎、蒙古栎）的粗根（5-10 mm）进行了为期1年的分解实验研究,来探索粗根分解和养分释放的动态变化规律。研究结果表明：黄檗、胡桃楸、水曲柳、色木槭、红松、落叶松、白桦、春榆、紫缎、蒙古栎粗根年分解系数分别为0.826、0.897、0.477、0.341、0.358、0.264、0.244、0.593、0.458、0.227。由此可见,胡桃楸分解速率最快,蒙古栎分解速率最慢。在粗根分解过程中,不同调控因子对根系分解的影响不同。研究结果表明,粗根的分解速率与根系的初始C/N比例呈显著负相关（P<0.0001）,与初始木质素含量呈负相关（P<0.0001）,与初始非结构性碳水化合物（NSC）含量呈正相关（P<0.0001）。初始C/N、木质素含量与非结构性碳水化合物含量分别可以解释所研究的10个树种粗根分解速率的68%、20%与65%。研究结论对于预测粗根参与的碳循环与养分释放具有重要意义。     

Spatial structure and genetic diversity of two tropical tree species with contrasting breeding systems and different ploidy levels

Analyses of the spatial distribution pattern, spatial genetic structure and of genetic diversity were carried out in two tropical tree species with contrasting breeding systems and different ploidy levels using a 50-ha demographic plot in a lowland dipterocarp forest in Peninsular Malaysia. Shorea leprosula is a diploid and predominantly outcrossed species, whereas S. ovalis ssp. sericea is an autotetraploid species with apomictic mode of reproduction. Genetic diversity parameters estimated for S. leprosula using microsatellite were consistently higher than using allozyme. In comparisons with S. leprosula and other tropical tree species, S. ovalis ssp. sericea also displayed relatively high levels of genetic diversity. This might be explained by the lower pressure of genetic drift due to tetrasomic inheritance, and for autotetraploids each locus can accommodate up to four different alleles and this allows maintenance of more alleles at individual loci. The observed high levels of genetic diversity in S. ovalis ssp. sericea can also be due to a random retention of more heterogeneous individuals in the past, and the apomictic mode of reproduction might be an evolutionary strategy, which allows the species to maintain high levels of genetic diversity. The spatial distribution pattern analyses of both species showed significant levels of aggregation at small and medium but random distribution at the big diameter-class. The decrease in magnitude of spatial aggregation from small- to large-diameter classes might be due to compensatory mortality during recruitment and survival under competitive thinning process. Spatial genetic structure analyses for both species revealed significant spatial genetic structure for short distances in all the three diameter-classes. The magnitude of spatial genetic structure in both species was observed to be decreasing from smaller- to larger-diameter classes. The high spatial genetic structuring observed in S. ovalis ssp. sericea at the small-diameter class is due primarily to limited seed dispersal and apomictic mode of reproduction. The similar observation in S. leprosula, however, can be explained by limited seed and pollen dispersal, which supports further the fact that the species is pollinated by weak fliers, mainly of Thrips and Megalurothrips in the lowland dipterocarp forest.     

Early effects of temperate agroforestry practices on soil organic matter and microbial enzyme activity

Plant and Soil - A field experiment was conducted to evaluate the effects of alley cropping systems on microbial activity and soil organic matter (SOM) pools. We hypothesized that enzyme activity...     

Influence of tree species composition on soil and soil solution properties in two mixed spruce-beech stands with contrasting history in Southern Germany

The properties of the soil and soil solution of two mixed spruce-beech forests in Southern Germany were investigated in order to identify non-additive effects of tree species compared to the monocultures. At five subplots in each of the two monocultures and at 10 subplots in each of the two mixed stands humus morphology, topsoil acidity as well as nitrate and sulphate concentrations in seepage water below the rooting zone were measured. At the Höglwald site, an 8×8 m sampling grid was also installed. Tree species composition within a 10 m circle surrounding each sampling point was correlated with the soil properties and the soil solution properties using the method of breakpoint estimation as well as a linear and a cubic model. At both sites, thickness and acidity of the forest floor as well as sulphate and nitrate concentrations in seepage water were significantly higher in the spruce monocultures than in the beech monocultures. Non-linear patterns in the correlation between both the thickness and the acidity of the forest floor and tree species composition occurred at both sites. In addition, nitrate concentration in the seepage water showed a non-linear correlation pattern at the Höglwald site. The correlation patterns were site specific and depended on stand history. While the influence of spruce at the Höglwald site was greater than expected from spruce monocultures, the opposite was found at the Schongau site. Interaction effects on nitrate concentration were absent at the Schongau site. Interaction effects on sulphate concentrations were absent at both sites. Current knowledge about the complex processes and patterns in mixed species stands is still limited. At least for certain properties, mixed species stands cannot be treated as a summation of the corresponding monocultures.     

Aquatic macroinvertebrate assemblages associated with two floating macrophyte species of contrasting root systems in a tropical wetland

Limnology - Aquatic macrophytes play an important role in structuring biotic communities. A comparative study of macroinvertebrate community structures associated with Salvinia auriculata Aublet...