Long-term nitrogen and phosphorus addition have stronger negative effects on microbial residual carbon in subsoils than topsoils in subtropical forests.

Highly weathered lowland (sub)tropical forests are widely recognized as nitrogen (N)-rich and phosphorus (P)-poor, and the input of N and P affects soil carbon (C) cycling and storage in these ecosystems. Microbial residual C (MRC) plays a crucial role in regulating soil organic C (SOC) stability in forest soils. However, the effects of long-term N and P addition on soil MRC across different soil layers remain unclear. This study conducted a 12-year N and P addition experiment in two typical subtropical plantation forests dominated by Acacia auriculiformis and Eucalyptus urophylla trees, respectively. We measured plant C input (fine root biomass, fine root C, and litter C), microbial community structure, enzyme activity (C/N/P-cycling enzymes), mineral properties, and MRC. Our results showed that continuous P addition reduced MRC in the subsoil (20-40 cm) of both plantations (A. auriculiformis: 28.44% and E. urophylla: 28.29%), whereas no significant changes occurred in the topsoil (0-20 cm). N addition decreased MRC in the subsoil of E. urophylla (25.44%), but had no significant effects on A. auriculiformis. Combined N and P addition reduced MRC (34.63%) in the subsoil of A. auriculiformis but not in that of E. urophylla. The factors regulating MRC varied across soil layers. In the topsoil (0-10 cm), plant C input (the relative contributions to the total variance was 20%, hereafter) and mineral protection (47.2%) were dominant factors. In the soil layer of 10-20 cm, both microbial characteristics (41.3%) and mineral protection (32.3%) had substantial effects, whereas the deeper layer (20-40 cm) was predominantly regulated by microbial characteristics (37.9%) and mineral protection (18.8%). Understanding differential drivers of MRC across soil depth, particularly in deeper soil layers, is crucial for accurately predicting the stability and storage of SOC and its responses to chronic N enrichment and/or increased P limitation in (sub)tropical forests.

Microbial nitrogen and phosphorus co-limitation across permafrost region.

The status of plant and microbial nutrient limitation have profound impacts on ecosystem carbon cycle in permafrost areas, which store large amounts of carbon and experience pronounced climatic warming. Despite the long-term standing paradigm assumes that cold ecosystems primarily have nitrogen deficiency, large-scale empirical tests of microbial nutrient limitation are lacking. Here we assessed the potential microbial nutrient limitation across the Tibetan alpine permafrost region, using the combination of enzymatic and elemental stoichiometry, genes abundance and fertilization method. In contrast with the traditional view, the four independent approaches congruently detected widespread microbial nitrogen and phosphorus co-limitation in both the surface soil and deep permafrost deposits, with stronger limitation in the topsoil. Further analysis revealed that soil resources stoichiometry and microbial community composition were the two best predictors of the magnitude of microbial nutrient limitation. High ratio of available soil carbon to nutrient and low fungal/bacterial ratio corresponded to strong microbial nutrient limitation. These findings suggest that warming-induced enhancement in soil nutrient availability could stimulate microbial activity, and probably amplify soil carbon losses from permafrost areas.

Mechanism for the restoration of degraded typical steppe by nitrogen and phosphorus co-addition.

The reduction of soil nutrient content is one of the major reasons caused grassland degradation in China. Nutrient addition is thus considered as an effective measure for the restoration of degraded grasslands. However, over-fertilization can lead to decrease in plant diversity. To clarify the appropriate amount of nutrient addition and the underlying mechanism that promotes grassland restoration, we set up a nitrogen and phosphorus co-addition experiment in a degraded typical steppe of Inner Mongolia, and examined the responses at community, functional group and species levels to nutrient addition. The results showed that nutrient addition enhanced biomass while did not reduce species richness at the community level. The biomass showed a saturation response with the increases of nutrient addition, which approached saturation under the 12.0 g N·m-2, 3.8 g P·m-2 treatment. Species richness increased significantly under the lower nutrient treatments (N <9.6 g·m-2, P < 3.0 g·m-2) compared with the control, while the two high nutrient treatments did not alter species richness. At the functional group level, biomass and abundance of perennial rhizome grasses increased significantly with the increases of nutrient addition levels. Biomass and density of annuals increased significantly under high nutrient addition levels. However, the abundance and biomass of perennial bunchgrasses and perennial forbs were rarely affected. At the species level, six target species responded differently to nutrient addition. Biomass of Leymus chinensis was significantly increased due to the increase of population density and individual biomass. Biomass of Stipa grandis, Agropyron cristatum and Cleistogenes squarrosa change little. Biomass of Potentilla acaulis and Carex korshinskyi were reduced due to the decreases in individual biomass and population density, respectively. As a measure of restoring degraded grassland, nutrient addition could significantly increase biomass and species diversity, decrease biomass of the degradation indicator species, and increase biomass of perennial rhizomes grasses.

Long-term phosphorus addition alleviates CO2 and N2O emissions via altering soil microbial functions in secondary rather primary tropical forests.

Tropical forests, where the soils are nitrogen (N) rich but phosphorus (P) poor, have a disproportionate influence on global carbon (C) and N cycling. While N deposition substantially alters soil C and N retention in tropical forests, whether P input can alleviate these N-induced effects by regulating soil microbial functions remains unclear. We investigated soil microbial taxonomy and functional traits in response to 10-year independent and interactive effects of N and P additions in a primary and a secondary tropical forest in Hainan Island. In the primary forest, N addition boosted oligotrophic bacteria and phosphatase and enriched genes responsible for C-, P-mineralization, nitrification and denitrification, suggesting aggravated P limitation while N excess. This might stimulate P excavation via organic matter mineralization, and enhance N losses, thereby increasing soil CO2 and N2O emissions by 86% and 110%, respectively. Phosphorus and NP additions elevated C-mining enzymes activity mainly due to intensified C limitation, causing 82% increase in CO2 emission. In secondary forest, P and NP additions reduced phosphatase activity, enriched fungal copiotrophs and increased microbial biomass, suggesting removal of nutrient deficiencies and stimulation of fungal growth. Meanwhile, soil CO2 emission decreased by 25% and N2O emission declined by 52-82% due to alleviated P acquisition from organic matter decomposition and increased microbial C and N immobilization. Overall, N addition accelerates most microbial processes for C and N release in tropical forests. Long-term P addition increases C and N retention via reducing soil CO2 and N2O emissions in the secondary but not primary forest because of strong C limitation to microbial N immobilization. Further, the seasonal and annual variations in CO2 and N2O emissions should be considered in future studies to test the generalization of these findings and predict and model dynamics in greenhouse gas emissions and C and N cycling.

Contrasting effects of nitrogen and phosphorus additions on nitrogen competition between coniferous and broadleaf seedlings.

Nitrogen (N) is a major element limiting plant growth and metabolism. Nitrogen addition can influence plant growth, N uptake, and species interactions, while phosphorus (P) addition may affect N acquisition. However, knowledge of how nutrient availability influences N uptake and species interactions remains limited and controversial. Here, pot experiments were conducted for 14 months, in which conifers (Pinus massoniana and Pinus elliottii) and broadleaved trees (Michelia maudiae and Schima superba) were planted in monoculture or mixture, and provided additional N and P in a full-factorial design. Nitrogen addition increased the biomass, but P addition did not significantly affect the biomass of the four subtropical species. Combined N and P (NP) addition had no additive effect on plant biomass over N addition. Total plant biomass was significantly positively correlated to root traits (branching intensity and root tissue density) and leaf traits (net photosynthetic rate, stomatal conductance, and transpiration rate), but negatively correlated to root diameter in response to nutrient addition. Plant uptake rates of NH4+ or NO3- were not altered by N addition, but P or NP additions decreased NH4+ uptake rates and increased NO3- uptake rates. Neighboring conifers significantly inhibited NH4+ and NO3- uptake rates of the two broadleaf species, but neighboring broadleaves had no effects on the N uptake rates of pine species. The effects of nutrient additions on interspecific interactions differed among species. Nitrogen addition altered the interaction of P. elliottii and M. maudiae from neutral to competition, while P addition altered the interaction of P. massoniana and M. maudiae from neutral to favorable effects. Increasing nutrient availability switched the direction of interspecific interaction in favor of pines. This study provides insights into forest management for productivity improvement and optimizing the selection of broadleaf species regarding differences in soil fertility of subtropical plantations.

ß-diversity in temperate grasslands is driven by stronger environmental filtering of plant species with large genomes.

Elucidating mechanisms underlying community assembly and biodiversity patterns is central to ecology and evolution. Genome size (GS) has long been hypothesized to potentially affect species' capacity to tolerate environmental stress and might therefore help drive community assembly. However, its role in driving ß-diversity (i.e., spatial variability in species composition) remains unclear. We measured GS for 161 plant species and community composition across 52 sites spanning a 3200-km transect in the temperate grasslands of China. By correlating the turnover of species composition with environmental dissimilarity, we found that resource filtering (i.e., environmental dissimilarity that includes precipitation, and soil nitrogen and phosphorus concentrations) affected ß-diversity patterns of large-GS species more than small-GS species. By contrast, geographical distance explained more variation of ß-diversity for small-GS than for large-GS species. In a 10-year experiment manipulating levels of water, nitrogen, and phosphorus, adding resources increased plant biomass in species with large GS, suggesting that large-GS species are more sensitive to the changes in resource availability. These findings highlight the role of GS in driving community assembly and predicting species responses to global change.

Nitrogen enrichment alters multiple dimensions of grassland functional stability via changing compositional stability.

Anthropogenic nutrient enrichment is known to alter the composition and functioning of plant communities. However, how nutrient enrichment influences multiple dimensions of community- and ecosystem-level stability remains poorly understood. Using data from a nitrogen (N) and phosphorus (P) addition experiment in a temperate semi-arid grassland that experienced a natural drought, we show that N enrichment, not P enrichment, decreased grassland functional and compositional temporal stability, resistance and recovery but increased functional and compositional resilience. Compositional stability and species asynchrony, rather than species diversity, were identified as key determinants of all dimensions of grassland functional stability, except for recovery. Whereas grassland functional recovery was decoupled from compositional recovery, N enrichment altered other dimensions of functional stability primarily through changing their corresponding compositional stability dimensions. Our findings highlight the need to examine ecological stability at the community level for a more mechanistic understanding of ecosystem dynamics in the face of environmental change.

Phosphorus addition regulates the growth of Chinese fir by changing needle nitrogen fractions in growing and dormant seasons.

Forest productivity is generally limited by nutrient scarcity. This study aims to reveal seasonal interactions among leaf carbon (C), nitrogen (N) fractions and tree growth driven by nutrient addition in a subtropical forest. Here, a field nutrient addition experiment was conducted with six treatments, namely, +N5 (5 g N m-2 yr-1), +N10 (10 g N m-2 yr-1), +P5 (5 g P m-2 yr-1), +N5 + P5, +N10 + P5, and control (N0 + P0). C fractions (structural and non-structural carbohydrates) and N fractions (soluble N, nucleic N and protein N) in needles as well as tree growth indicated by basal area increment (BAI) were measured in growing and dormant seasons. Total N and protein N in old needles were significantly increased by P addition, while no significant differences of non-structural carbohydrates in young (<1-year old) and old needles (>1-year old) were detected among the treatments in both seasons. N and P addition increased the structural carbohydrates of old needles in dormant season. P addition decreased and increased tree growth in growing and dormant seasons, respectively. The variation of BAI was explained 18.3 % by total N and 17.8 % by protein N in growing season, and was explained 33.9 % by total N and 34.2 % by protein N in dormant season. Our study suggested that the P addition effect on Chinese fir growth mostly depends on needle N fractions. This study highlights tree seasonal growth driven by nutrient alteration might be characterized by leaf N fractions rather than C fractions in subtropical forests.

Variabilities in autumn cyanobacterial responses to ecosystem external enrichments based on nutrient addition bioassay in Pengxi River, Three Gorges Reservoir, China.

Nutrient availability, is a crucial anthropogenic stressor promoting freshwater eutrophication and rapid expansion of harmful algal blooms (HABs), deteriorating water quality and threatening public health worldwide. The estimation of the HABs community responses to diel changes in the nutrients while characterizing the ecosystem growth limiting factors, is key to prudent watershed management. The present study investigated the short-term variabilities in autumn cyanobacterial responses to the external nutrient inputs into the Pengxi River using the nutrient addition bioassay approach. Results reveal phytoplankton community structure dominated by the cyanobacteria: Anabaena and Aphanizomenon spp. (relative abundance = 46.20% equilibrium abundance), followed by the diatoms, out of which Lindayia bodaniica, are preponderant. Nutrient enrichment triggered strong variabilities in dominance and successions among the cyanobacterial group, with maximum dominance (76.34%) exhibited by the Aphanizomenon sp. upon NH4 addition. Fe enrichment led to the succession of cyanobacteria, Leptolyngbya tenuis, which was below the detectable limit in the control, indicating the role of Fe in its proliferation. Studies on nutrient limitation demonstrated P/NH4 co-limited ecosystem, with P as the primary and NH4, a secondary limiting factor. The nitrate preference index (NO3-RPI = 0.991) shows a high preference for NH4 while NO3 constitutes the bulk of the ecosystem TN. Considering the elevated NO3 concentration, we posit that a shift in the phytoplankton community structure from cyanobacteria to diatoms dominated ecosystem, is expected following Fe depletion and a further stretch on the current ecosystem NH4 limitation. The study provides useful and first-ever insights for nutrient reduction in the middle Three Gorges Reservoir (TGR) before the onset of the heavy HABs during spring in the Pengxi River.

Delayed autumnal leaf senescence following nutrient fertilization results in altered nitrogen resorption.

Increased atmospheric nitrogen (N) deposition could create an imbalance between N and phosphorus (P), which may substantially impact ecosystem functioning. Changes in autumnal phenology (i.e., leaf senescence) and associated leaf nutrient resorption may profoundly impact plant fitness and productivity. However, we know little about how and to what extent nutrient addition affects leaf senescence in tree species, or how changes in senescence may influence resorption. We thus investigated the impacts of N and P addition on leaf senescence and leaf N resorption in 2-year-old larch (Larix principisrupprechtii) seedlings in northern China. Results showed that nutrient addition (i.e., N, P or N + P addition) significantly delayed autumnal leaf senescence, and decreased leaf N resorption efficiency (NRE) and proficiency (NRP), particularly in the N and N + P treatments. Improved leaf N concentrations were correlated with delayed leaf senescence, as indicated by the positive relationship between mature leaf N concentrations and the timing of leaf senescence. Following nutrient addition, larch seedlings shifted toward delayed onset, but more rapid, leaf senescence. Additionally, we observed an initial negative correlation between the timing of leaf senescence and NRE and NRP, followed by a positive correlation, indicating delayed and less efficient remobilization during the early stages of senescence, followed by accelerated resorption in the later stages. However, the latter effect was potentially impaired by the increased risk of early autumn frost damage, thus failed to fully compensate for the negative effects observed during the early stages of senescence. Improved soil P availability increased leaf N resorption and thus weakened the negative impact of delayed leaf senescence on leaf N resorption, so P addition had no significant impact on leaf N resorption. Overall, our findings clarify the relationship between nutrient addition-resorption and the linkage with leaf senescence, and would have important implications for plant nutrient conservation strategy and nutrient cycling.

Soil phosphorus drives plant trait variations in a mature subtropical forest.

Earth system models are implementing soil phosphorus dynamic and plant functional traits to predict functional changes in global forests. However, the linkage between soil phosphorus and plant traits lacks empirical evidence, especially in mature forests. Here, we examined the soil phosphorus constraint on plant functional traits in a mature subtropical forest based on observations of 9943 individuals from 90 species in a 5-ha forest dynamic plot and 405 individuals from 15 species in an adjacent 10-year nutrient-addition experiment. We first confirmed a pervasive phosphorus limitation on subtropical tree growth based on leaf N:P ratios. Then, we found that soil phosphorus dominated multidimensional trait variations in the 5-ha forest dynamic plot. Soil phosphorus content explained 44% and 53% of the variance in the traits defining the main functional space across species and communities, respectively. Lastly, we found much stronger phosphorus effects on most plant functional traits than nitrogen at both species and community levels in the 10-year nutrient-addition experiment. This study provides evidence for the consistent pattern of soil phosphorus constraint on plant trait variations between the species and community levels in a mature evergreen broadleaf forest in the East Asian monsoon region. These findings shed light on the predominant role of soil phosphorus on plant functional trait variations in mature subtropical forests, providing new insights for models to incorporate soil phosphorus constraint in predicting future vegetation dynamics.

Study on nutrient limitation of phytoplankton growth in Xiangxi Bay of the Three Gorges Reservoir, China.

After the impoundment of the Three Gorges Reservoir (TGR), algal blooms in the sidearm tributaries have resulted from increasing nutrient loads along the major tributaries. Field sampling and in situ nutrient addition bioassay were implemented to examine the nutrient limitation of phytoplankton growth and bloom initiation during autumn in Xiangxi Bay of the TGR. Result shows that P is the primary limiting nutrient for algal growth and bloom in Xiangxi Bay during autumn. The treatment involving the combination of N, P and Si had a significant (p < .05) additional effect on the growth of phytoplankton. The N, P, Si combined treatment increased growth by 10-50% relative to the N and P treatments from day 1 to day 4, respectively. Trace metal additions involving Fe, Zn, Mn, and Cu and/or in combination with N, P, and Si initially resulted in an extremely low growth rate which later increased significantly (p < .05) towards the end of the study. The present study provides an insight into the responses of different phytoplankton taxa in autumn under nutrient conditions in the tributary bay. The nutrient limitation study is recognized as the first step to mitigating the bloom while proposing an effective nutrient control strategy. The outcome of which can provide the basis for formulating sustainable watershed management. Multiple nutrients reductions with P as primary concern are required for a lasting management solution to the risk of bloom in the TGR.

The Impact of Warming and Nutrients on Algae Production and Microcystins in Seston from the Iconic Lake Lesser Prespa, Greece.

Lake Lesser Prespa and its adjacent pond, Vromolimni in Greece, is a shallow freshwater system and a highly protected area hosting an exceptional biodiversity. The occurrence of microcystins (MCs) producing cyanobacterial blooms in these waters during recent years can be harmful to the wildlife. We tested the hypothesis that both cyanobacterial biomass and MCs are strongly influenced by nutrients (eutrophication) and warming (climate change). Lake and pond water was collected from two sites in each water body in 2013 and incubated at three temperatures (20 °C, 25 °C, 30 °C) with or without additional nutrients (nitrogen +N, phosphorus +P and both +N and +P). Based on both biovolume and chlorophyll-a concentrations, cyanobacteria in water from Lesser Prespa were promoted primarily by combined N and P additions and to a lesser extent by N alone. Warming seemed to yield more cyanobacteria biomass in these treatments. In water from Vromolimni, both N alone and N+P additions increased cyanobacteria and a warming effect was hardly discernible. MC concentrations were strongly increased by N and N+P additions in water from all four sites, which also promoted the more toxic variant MC-LR. Hence, both water bodies seem particularly vulnerable to further N-loading enhancing MC related risks.

Individual and combined effects of multiple global change drivers on terrestrial phosphorus pools: A meta-analysis.

Human activity-induced global change drivers have dramatically changed terrestrial phosphorus (P) dynamics. However, our understanding of the interactive effects of multiple global change drivers on terrestrial P pools remains elusive, limiting their incorporation into ecological and biogeochemical models. We conducted a meta-analysis using 1751 observations extracted from 283 published articles to evaluate the individual, combined, and interactive effects of elevated CO2, warming, N addition, P addition, increased rainfall, and drought on P pools of plant (at both single-plant and plant-community levels), soil and microbial biomass. Our results suggested that (1) terrestrial P pools showed the most sensitive responses to the individual effects of warming and P addition; (2) P pools were consistently stimulated by P addition alone or in combination with simultaneous N addition; (3) environmental and experimental setting factors such as ecosystem type, climate, and latitude could significantly influence both the individual and combined effects; and (4) the interactive effects of two-driver pairs across multiple global change drivers are more likely to be additive rather than synergistic or antagonistic. Our findings highlighting the importance of additive interactive effects among multiple global change drivers on terrestrial P pools would be useful for incorporating P as controls on ecological processes such as photosynthesis and plant growth into ecosystem models used to analyze effects of multiple drivers under future global change.

Long-term simulated nitrogen deposition alters the plant cover dynamics of a Mediterranean rosemary shrubland in Central Spain through defoliation.

Nitrogen (N) deposition due to anthropogenic pollution is a major driver of the global biodiversity loss. We studied the effect of experimental N and phosphorus (P) fertilization (0, 10, 20, and 50 kg N ha-1 year-1 and 14 kg P ha-1 year-1 over the background deposition levels) on plant cover dynamics of a rosemary (Rosmarinus officinalis L.) shrubland after 8 years of nutrient addition in a semiarid Mediterranean ecosystem from Central Spain. We specifically aimed at testing whether N deposition has the potential to influence the observed expanding trend of woody vegetation into areas dominated by grassland, biological soil crusts, and bare soil. Our results show that N addition loads above 10 kg N ha-1 year-1 reverted the cover dynamics of shrubs. Under N addition conditions, N was no longer a limiting nutrient and other elements, especially P and calcium, determined the seasonal growth of young twigs. Interestingly, N fertilization did not inhibit the growth of young shoots; our estimates point to a reduced rosemary leaf lifespan that is driving individuals to death. This may be triggered by long-term accumulation of N compounds in leaves, suggesting the need to consider the old organs and tissues in long-lived perennial plants, where N toxicity effects could be more mediated by accumulation processes. Shrublands are a widely distributed ecosystem type in biodiverse Mediterranean landscapes, where shrubs play a key role as nurse plants. Therefore, the disappearance of shrublands may accelerate the biodiversity loss associated with other global change drivers, hamper the recruitment of seedlings of woody species, and, as a consequence, accelerate desertification.

C, N and P fertilization in an Amazonian rainforest supports stoichiometric dissimilarity as a driver of litter diversity effects on decomposition.

Plant leaf litter generally decomposes faster as a group of different species than when individual species decompose alone, but underlying mechanisms of these diversity effects remain poorly understood. Because resource C : N : P stoichiometry (i.e. the ratios of these key elements) exhibits strong control on consumers, we supposed that stoichiometric dissimilarity of litter mixtures (i.e. the divergence in C : N : P ratios among species) improves resource complementarity to decomposers leading to faster mixture decomposition. We tested this hypothesis with: (i) a wide range of leaf litter mixtures of neotropical tree species varying in C : N : P dissimilarity, and (ii) a nutrient addition experiment (C, N and P) to create stoichiometric similarity. Litter mixtures decomposed in the field using two different types of litterbags allowing or preventing access to soil fauna. Litter mixture mass loss was higher than expected from species decomposing singly, especially in presence of soil fauna. With fauna, synergistic litter mixture effects increased with increasing stoichiometric dissimilarity of litter mixtures and this positive relationship disappeared with fertilizer addition. Our results indicate that litter stoichiometric dissimilarity drives mixture effects via the nutritional requirements of soil fauna. Incorporating ecological stoichiometry in biodiversity research allows refinement of the underlying mechanisms of how changing biodiversity affects ecosystem functioning.

Strategic enhancement of algal biomass, nutrient uptake and lipid through statistical optimization of nutrient supplementation in coupling Scenedesmus obliquus-like microalgae cultivation and municipal wastewater treatment.

Supplementing proper nutrients could be a strategy for enhancing algal biomass, nutrients uptake and lipid accumulation in the coupling system of biodiesel production and municipal wastewater treatment. However, there is scant information reporting systematic studies on screening and optimization of key supplemented components in the coupling system. The main factors were scientifically screened and optimized using statistical methods. Plackett-Burman design (PBD) was used to explore the roles of added nutrient factors, whereas response surface methodology (RSM) was employed for optimization. Based on the statistic analysis, the optimum added TP and FeCl3·6H2O concentrations for Scenedesmus obliquus-like microalgae growth, nutrients uptake and lipid accumulation were 4.41 mg L(-1) and 6.48 mg L(-1), respectively. The corresponding biomass, lipid content and TN/TP removal efficiency were 1.46 g L(-1), 36.26% and >99%. The predicted value agreed well with the experimental value, as determined by validation experiments, which confirmed the availability and accuracy of the model.

Nitrogen cycling in canopy soils of tropical montane forests responds rapidly to indirect N and P fertilization.

Although the canopy can play an important role in forest nutrient cycles, canopy-based processes are often overlooked in studies on nutrient deposition. In areas of nitrogen (N) and phosphorus (P) deposition, canopy soils may retain a significant proportion of atmospheric inputs, and also receive indirect enrichment through root uptake followed by throughfall or recycling of plant litter in the canopy. We measured net and gross rates of N cycling in canopy soils of tropical montane forests along an elevation gradient and assessed indirect effects of elevated nutrient inputs to the forest floor. Net N cycling rates were measured using the buried bag method. Gross N cycling rates were measured using (15) N pool dilution techniques. Measurements took place in the field, in the wet and dry season, using intact cores of canopy soil from three elevations (1000, 2000 and 3000 m). The forest floor had been fertilized biannually with moderate amounts of N and P for 4 years; treatments included control, N, P, and N + P. In control plots, gross rates of NH4 (+) transformations decreased with increasing elevation; gross rates of NO3 (-) transformations did not exhibit a clear elevation trend, but were significantly affected by season. Nutrient-addition effects were different at each elevation, but combined N + P generally increased N cycling rates at all elevations. Results showed that canopy soils could be a significant N source for epiphytes as well as contributing up to 23% of total (canopy + forest floor) mineral N production in our forests. In contrast to theories that canopy soils are decoupled from nutrient cycling in forest floor soil, N cycling in our canopy soils was sensitive to slight changes in forest floor nutrient availability. Long-term atmospheric N and P deposition may lead to increased N cycling, but also increased mineral N losses from the canopy soil system.

Multi-nutrient vs. nitrogen-only effects on carbon sequestration in grassland soils.

Human activities have greatly increased the availability of biologically active forms of nutrients [e.g., nitrogen (N), phosphorous (P), potassium (K), magnesium (Mg)] in many soil ecosystems worldwide. Multi-nutrient fertilization strongly increases plant productivity but may also alter the storage of carbon (C) in soil, which represents the largest terrestrial pool of organic C. Despite this issue is important from a global change perspective, key questions remain on how the single addition of N or the combination of N with other nutrients might affect C sequestration in human-managed soils. Here, we use a 19-year old nutrient addition experiment on a permanent grassland to test for nutrient-induced effects on soil C sequestration. We show that combined NPKMg additions to permanent grassland have 'constrained' soil C sequestration to levels similar to unfertilized plots whereas the single addition of N significantly enhanced soil C stocks (N-only fertilized soils store, on average, 11 t C ha(-1) more than unfertilized soils). These results were consistent across grazing and liming treatments suggesting that whilst multi-nutrient additions increase plant productivity, soil C sequestration is increased by N-only additions. The positive N-only effect on soil C content was not related to changes in plant species diversity or to the functional composition of the plant community. N-only fertilized grasslands show, however, increases in total root mass and the accumulation of organic matter detritus in topsoils. Finally, soils receiving any N addition (N only or N in combination with other nutrients) were associated with high N losses. Overall, our results demonstrate that nutrient fertilization remains an important global change driver of ecosystem functioning, which can strongly affect the long-term sustainability of grassland soil ecosystems (e.g., soils ability to deliver multiple ecosystem services).