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Summary: Knowledge of differential life‐history strategies in arbuscular mycorrhizal (AM) fungi is relevant for understanding the ecology of this group and its potential role in sustainable agriculture and carbon sequestration. At present, AM fungal life‐history theories often focus on differential investment into intra‐ vs extraradical structures among AM fungal taxa, and its implications for plant benefits. With this Viewpoint we aim to expand these theories by integrating a mycocentric economics‐ and resource‐based life‐history framework. As in plants, AM fungal carbon and nutrient demands are stoichiometrically coupled, though uptake of these elements is spatially decoupled. Consequently, investment in morphological structures for carbon vs nutrient uptake is not in competition. We argue that understanding the ecology and evolution of AM fungal life‐history trade‐offs requires increased focus on variation among structures foraging for the same element, that is within intra‐ or extraradical structures (in our view a 'horizontal' axis), not just between them ('vertical' axis). Here, we elaborate on this argument and propose a range of plausible life‐history trade‐offs that could lead to the evolution of strategies in AM fungi, providing testable hypotheses and creating opportunities to explain AM fungal co‐existence, and the context‐dependent effects of AM fungi on plant growth and soil carbon dynamics. [ABSTRACT FROM AUTHOR]

Background: Fungicides are an effective tool for protecting crops and maintaining a steady food supply. However, as pathogens continue to evolve, it is crucial to prolong the effectiveness of fungicides by delaying resistance development. A key strategy to achieving this is to combine or rotate fungicides with different modes of action. As fungicides lack specificity, they inevitably affect both pathogenic and non-pathogenic fungi when surrounding environments are unintentionally contaminated. Our study aims to investigate the effects of recommended application methods to prevent resistance development, specifically repeated-single fungicide, simultaneous mixture, and sequential applications on non-target soil fungi, and the subsequent impacts on important soil processes. We used fungicides with different modes of action on soil microcosms inoculated with fungi at varying levels of diversity (3, 5, and 8 species) isolated from a protected grassland. Results: We found that repeated treatments of individual isopyrazam and prothioconazole differentially inhibited fungal activity. Although mixture applications are considered more protectant against crop pathogen resistance than repeated application, our study revealed stronger negative effects of simultaneous application on saprobic fungi and consequently on soil processes. However, contrary to expectations, higher fungal diversity did not translate to improved soil function under these conditions. Conclusions: The simultaneous application of fungicides with different modes of action (MoA) has more pronounced non-target effects on soil compared to the individual or sequential application of fungicides. These non-target effects extend beyond the intended control of pathogenic fungi, impacting saprobic and beneficial soil microbes and the critical processes they drive. When fungicides are applied concurrently, microbial activities in the soil are significantly altered, even in soils with high microbial diversity. Our study emphasizes the importance of carefully considering the unintended consequences of fungicide use in agriculture. As we strive for a secure food supply, it is crucial to investigate the broader environmental impacts of these chemical interventions, including their effects on non-pathogenic fungi and overall soil health. [ABSTRACT FROM AUTHOR]

Litter decomposition is a key ecosystem process, relevant for the release and storage of nutrients and carbon in soil. Soil fungi are one of the dominant drivers of organic matter decomposition, but fungal taxa differ substantially in their functional ability to decompose plant litter. Knowledge is mostly based on observational data and subsequent molecular analyses and in vitro studies have been limited to forest ecosystems. In order to better understand functional traits of saprotrophic soil fungi in grassland ecosystems, we isolated 31 fungi from a natural grassland and performed several in vitro studies testing for i) leaf and wood litter decomposition, ii) the ability to use carbon sources of differing complexity, iii) the enzyme repertoire. Decomposition strongly varied among phyla and isolates, with Ascomycota decomposing the most and Mucoromycota decomposing the least. The phylogeny of the fungi and their ability to use complex carbon were the most important predictors for decomposition. Our findings show that it is crucial to understand the role of individual members and functional groups within the microbial community. This is an important way forward to understand the role of microbial community composition for the prediction of litter decomposition and subsequent potential carbon storage in grassland soils. [ABSTRACT FROM AUTHOR]

Trait-based frameworks are promising tools to understand the functional consequences of community shifts in response to environmental change. The applicability of these tools to soil microbes is limited by a lack of functional trait data and a focus on categorical traits. To address this gap for an important group of soil microorganisms, we identify trade-offs underlying a fungal economics spectrum based on a large trait collection in 28 saprobic fungal isolates, derived from a common grassland soil and grown in culture plates. In this dataset, ecologically relevant trait variation is best captured by a three-dimensional fungal economics space. The primary explanatory axis represents a dense-fast continuum, resembling dominant life-history trade-offs in other taxa. A second significant axis reflects mycelial flexibility, and a third one carbon acquisition traits. All three axes correlate with traits involved in soil carbon cycling. Since stress tolerance and fundamental niche gradients are primarily related to the dense-fast continuum, traits of the 2nd (carbon-use efficiency) and especially the 3rd (decomposition) orthogonal axes are independent of tested environmental stressors. These findings suggest a fungal economics space which can now be tested at broader scales. Challenges in obtaining empirical trait data hinder the development of trait-based frameworks for soil microbes. Here, the authors analyse traits of saprobic fungal isolates from a grassland site to propose a fungal economics spectrum, suggesting a general trait framework for soil fungi. [ABSTRACT FROM AUTHOR]

The tropical Andes are a species‐rich and nitrogen‐limited system, susceptible to increased nitrogen (N) inputs from the atmosphere. However, our understanding of the impacts of increased N input on belowground systems, in particular on protists and their role in nutrient cycling, remains limited. We explored how increased N affects protists in tropical montane rainforests in Ecuador using high‐throughput sequencing (HTS) of environmental DNA from two litter layers. In addition, we manipulated the amount of arbuscular mycorrhizal fungi (AMF) and mesofauna, both playing a significant role in N cycling and interacting in complex ways with protist communities. We found that N strongly affected protist community composition in both layers, while mesofauna reduction had a stronger effect on the lower layer. Changes in concentration of the AMF marker lipid had little effect on protists. In both layers, the addition of N increased phagotrophs and animal parasites and decreased plant parasites, while mixotrophs decreased in the upper layer but increased in the lower layer. In the upper layer with higher AMF concentration, mixotrophs decreased, while in the lower layer, photoautotrophs increased and plant parasites decreased. With reduced mesofauna, phagotrophs increased and animal parasites decreased in both layers, while plant parasites increased only in the upper layer. The findings indicate that to understand the intricate response of protist communities to environmental changes, it is critical to thoroughly analyze these communities across litter and soil layers, and to include HTS. [ABSTRACT FROM AUTHOR]

Root associated fungal (RAF) communities can exert strong effects on plant communities and are potentially sensitive to shifts in soil fertility. As increased atmospheric nitrogen (N) and phosphorus (P) deposition can alter the nutrient balance in natural ecosystems, we assessed the response of RAF communities to a fertilization experiment deployed on a highly diverse Andean forest. The stand level fine root fraction was sampled after 7 years of systematic N and P additions and RAF communities were characterized by a deep sequencing approach. We expected that fertilization will enhance competition of fungal taxa for limiting nutrients, thus eliciting diversity reductions and alterations in the structure of RAF communities. Fertilization treatments did not reduce RAF richness but affected community composition. At the phylum level fertilization reduced richness exclusively among Glomeromycota. In contrast, N and P additions (alone or in combination) altered the composition of several fungal phyla. The lack of a generalized response to long‐term fertilization among RAF lineages suggests that most of these lineages will not be directly and immediately affected by the increasing rates of atmospheric N and P deposition expected for this region by 2050. [ABSTRACT FROM AUTHOR]

Summary: During the last few decades, a plethora of sequencing studies provided insight into fungal community composition under various environmental conditions. Still, the mechanisms of species assembly and fungal spread in soil remain largely unknown. While mycelial growth patterns are studied extensively, the abundant formation of asexual spores is often overlooked, though representing a substantial part of the fungal life cycle relevant for survival and dispersal. Here, we explore asexual sporulation (spore abundance, size and shape) in 32 co‐occurring soil fungal isolates under varying resource conditions, to answer the question whether resource limitation triggers or inhibits fungal investment into reproduction. We further hypothesized that trade‐offs exist in fungal investment towards growth, spore production and size. The results revealed overall increased fungal investment into spore production under resource limitations; however, effect sizes and response types varied strongly among fungal isolates. Such isolate‐specific effects were apparent in all measured traits, resulting in unique trait spaces of individual isolates. This comprehensive dataset also elucidated variability in sporulation strategies and trade‐offs with fungal growth and reproduction under resource scarcity, as only predicted by theoretical models before. The observed isolate‐specific strategies likely underpin mechanisms of co‐existence in this diverse group of saprobic soil fungi. [ABSTRACT FROM AUTHOR]

Soil ecological stoichiometry provides powerful theories to integrate the complex interplay of element cycling and microbial communities into biogeochemical models. One essential assumption is that microbes maintain stable C:N:P (carbon:nitrogen:phosphorus) ratios independent of resource supply, although such homeostatic regulations have rarely been assessed in individual microorganisms. Here, we report an unexpected high flexibility in C:N and C:P values of saprobic fungi along nutrient supply gradients, overall ranging between 7‐126 and 20‐1488, respectively, questioning microbial homeostasis. Fungal N:P varied comparatively less due to simultaneous reductions in mycelial N and P contents. As a mechanism, internal recycling processes during mycelial growth and an overall reduced N and P uptake appear more relevant than element storage. The relationships among fungal stoichiometry and growth disappeared in more complex media. These findings affect our interpretation of stoichiometric imbalances among microbes and soils and are highly relevant for developing microbial soil organic carbon and nitrogen models. [ABSTRACT FROM AUTHOR]

Summary: Anthropogenic atmospheric deposition can increase nutrient supply in the most remote ecosystems, potentially affecting soil biodiversity. Arbuscular mycorrhizal fungal (AMF) communities rapidly respond to simulated soil eutrophication in tropical forests. Yet the limited spatio‐temporal extent of such manipulations, together with the often unrealistically high fertilization rates employed, impedes generalization of such responses.We sequenced mixed root AMF communities within a seven year‐long fully factorial nitrogen (N) and phosphorus (P) addition experiment, replicated at three tropical montane forests in southern Ecuador with differing environmental characteristics. We hypothesized: strong shifts in community composition and species richness after long‐term fertilization, site‐ and clade‐specific responses to N vs P additions depending on local soil fertility and clade life history traits respectively.Fertilization consistently shifted AMF community composition across sites, but only reduced richness of Glomeraceae. Compositional changes were mainly driven by increases in P supply while richness reductions were observed only after combined N and P additions.We conclude that moderate increases of N and P exert a mild but consistent effect on tropical AMF communities. To predict the consequences of these shifts, current results need to be supplemented with experiments that characterize local species‐specific AMF functionality. [ABSTRACT FROM AUTHOR]

Summary: The dependency of microbial activity on nutrient availability in soil is only partly understood, but highly relevant for nutrient cycling dynamics. In order to achieve more insight on microbial adaptations to nutrient limiting conditions, precise physiological knowledge is needed. Therefore, we developed an experimental system assessing traits of 16 saprobic fungal isolates in nitrogen (N) limited conditions. We tested the hypotheses that (1) fungal traits are negatively affected by N deficiency to a similar extent and (2) fungal isolates respond in a phylogenetically conserved fashion. Indeed, mycelial density, spore production and fungal activity (respiration and enzymatic activity) responded similarly to limiting conditions by an overall linear decrease. By contrast, mycelial extension and hyphal elongation peaked at lowest N supply (C:N 200), causing maximal biomass production at intermediate N contents. Optimal N supply rates differed among isolates, but only the extent of growth reduction was phylogenetically conserved. In conclusion, growth responses appeared as a switch from explorative growth in low nutrient conditions to exploitative growth in nutrient‐rich patches, as also supported by responses to phosphorus and carbon limitations. This detailed trait‐based pattern will not only improve fungal growth models, but also may facilitate interpretations of microbial responses observed in field studies. [ABSTRACT FROM AUTHOR]

In tropical ecosystems, interactions between arbuscular mycorrhizal fungi (AMF) and other organisms have been little studied, but may be of significant importance for understanding the role of AMF in decomposition processes and nutrient cycling. In this study, we used ingrowth cores to investigate the impacts of regular rotation of the cores, defaunation and nitrogen addition on AMF, microbial biomass and microarthropods in the fermentation/humus (F/H) and litter (L) layers of an Ecuadorian montane tropical rainforest. AMF were substantially reduced in the F/H layer (to 34% of initial), while in the L layer they remained constant during the experiment. Overall, microorganisms and microarthropods were largely independent of AMF hyphae and their exudates, however, defaunation strongly affected the recovery of their communities. Nitrogen addition increased the quality of litter material and beneficially affected microbial communities thereby increasing decomposition rates, but did not impact AMF abundance and microarthropod communities. These findings suggest that the cutoff of the carbon supply from the plant to the fungal mycelium was not compensated by switching resources in the F/H layer, underlining a strong association between AMF and living roots. While in the L layer, AMF likely competed with saprotrophic microorganisms for litter-derived resources at intermediate stages of decomposition pointing to indirect contributions of AMF to decomposition processes. Overall, the results support the view that root-derived resources are important in fueling soil food webs, but also indicate that in the studied montane rainforest these resources are only available close to roots and not channeled distant to roots via AMF. [ABSTRACT FROM AUTHOR]

Ecosystem processes and the organisms involved are generally limited by the availability of one or more elements in soil, an important phenomenon to consider for our understanding of ecosystem functioning and future changes. Especially in tropical forests, typically growing on nutrient depleted soils, nitrogen (N), phosphorus (P) or other limitations are assumed. However, large‐scale nutrient manipulation experiments revealed complex site‐specific patterns and several authors raised the need for novel approaches to reveal deeper mechanistic insights on limitation patterns, especially concerning soil microbial activity.In order to gain such deeper knowledge, based on a short review of previous small‐scale studies focusing on soil micro‐organisms, we developed an experimental approach which controls for common biases, including indirect treatment effects, addition of co‐elements or nutrient pulses. Using this technique, we tested the hypotheses that fungi growing in tropical forest soils are mainly limited by P and that N versus P limitations shift along an altitudinal gradient.Mesh bags of 38 μm filled with sand were amended with weak ion exchange resins loaded with N, P or potassium (K) and buried underneath the litter layer at three altitudinal sites in southern Ecuador. After a period of four months, the in‐growth of fungal hyphae was quantified, phospholipid fatty acids were analysed for a subset of samples, and chemical properties were determined.In line with the first hypothesis, hyphal abundance was increased in P‐amended mesh bags, indicating P limitation. However, this pattern was not significantly shifted along the altitudinal gradient. By contrast, N addition increased hyphal abundance at the lowest site, compared to significant reductions at 2,000 and 3,000 m—not only in fungi but also in bacterial abundance as indicated by PLFA analyses, contradicting common soil‐age hypotheses. Decreased nutrient immobilization and fungal in‐growth at higher elevations suggest slow microbial activity, including nitrification, which may have caused toxic ammonium accumulation.The experimental design offers a promising tool to provide more mechanistic and soil‐focused analyses specifically targeting microbes, which in this system strongly supported the hypothesis of primary fungal P limitation in tropical forest soils. A plain language summary is available for this article. [ABSTRACT FROM AUTHOR]

The socio-economic values of fertile and carbon-rich Dark Earth soils are well described from the Amazon region. Very recently, Dark Earth soils were also identified in tropical West Africa, with comparable beneficial soil properties and plant growth-promoting effects. The impact of this management technique on soil microbial communities, however, is less well understood, especially with respect to the ecologically relevant group of arbuscular mycorrhizal (AM) fungi. Thus, we tested the hypotheses that (1) improved soil quality in African Dark Earth (AfDE) will increase soil microbial biomass and shift community composition and (2) concurrently increased nutrient availability will negatively affect AM fungal communities. Microbial communities were distinct in AfDE in comparison to adjacent sites, with an increased fungal:bacterial ratio of 71%, a pattern mainly related to shifts in pH. AM fungal abundance and diversity, however, did not differ despite clearly increased soil fertility in AfDE, with 3.7 and 1.7 times greater extractable P and total N content, respectively. The absence of detrimental effects on AM fungi, often seen following applications of inorganic fertilizers, and the enhanced role of saprobic fungi relevant for mineralization and C sequestration support previous assertions of this management type as a sustainable alternative agricultural practice. [ABSTRACT FROM AUTHOR]

Global changes in nutrient deposition rates are likely to have profound effects on plant communities, particularly in the nutrient-limited systems of the tropics. We studied the effects of increased nutrient availability on the seedlings of six tree species in montane forests of southern Ecuador in situ. After five years of continued N, P, or N+P addition, naturally grown seedlings of each of the two most common species at each elevation (1000, 2000, and 3000 m asl) were harvested for analyses of leaf morphology, nutrient content, herbivory, and tissue biomass allocation. Most species showed increased foliar N and P concentrations after addition of each respective element. Leaf tissue N:P ratios of >20 in the control plants of all species suggest that P is more growth-limiting in these forests than N. Leaf morphological responses to nutrient addition were species and nutrient specific, with some species (Hedyosmum purparescens, Graffenrieda emarginata) exhibiting increased specific leaf area (SLA), and others (Graffenrieda harlingii) increased leaf area ratios (LAR). Pouteria torta (1000 m) had lower SLA and LAR after P addition. Increased herbivory was only evident in G. emarginata (after N and N+P addition). Only the species from 3000 m asl modified biomass allocation after nutrient addition. In general, N and N+P addition more strongly affected the species studied at the upper elevations, whereas P addition had a similar range of effects on the species at all elevations. We conclude that the responses of the studied tropical montane forest tree seedlings to chronic N and P addition are highly species-specific and that successful adaptation to increased nutrient availability will depend on species-specific morphological and physiological plasticity. [ABSTRACT FROM AUTHOR]

Abstract: Soil fungi and bacteria are the key players in the transformation and processing of carbon and nutrients in terrestrial ecosystems, yet controls on their abundance and activity are not well understood. Based on stoichiometric principles, soil microbial processes are expected to be limited by mineral nutrients, which are particularly scarce in often highly weathered tropical forest soils. Such limitation is directly relevant for the fate of soil carbon and global element cycles, but its extent and nature have never been assessed systematically across the tropical biome. Here, we address the relative importance of nitrogen, phosphorus, and other nutrients in limiting soil microbial biomass and process rates in tropical forests. We conducted an in‐depth literature review and a meta‐analysis of the available nutrient addition experiments in tropical forests worldwide. Our synthesis showed predominant and general phosphorus limitation of a variety of microbial processes across tropical forests, and additional nitrogen limitation in tropical montane forests. The apparent widespread microbial phosphorus limitation needs to be accounted for in the understanding and prediction of biogeochemical cycles in tropical forests and their future functioning. Other mineral nutrients or carbon may modify the importance of phosphorus, but more experimental studies are urgently needed. [ABSTRACT FROM AUTHOR]

Nitrogen (N) and phosphorus (P) deposition are increasing worldwide largely due to increased fertilizer use and fossil fuel combustion. Most work with N and P deposition in natural ecosystems has focused on temperate, highly industrialized, regions. Tropical regions are becoming more developed, releasing large amounts of these nutrients into the atmosphere. Nutrient enrichment in nutrient-poor systems such as tropical montane forest can represent a relatively large shift in nutrient availability, especially for sensitive microorganisms such as arbuscular mycorrhizal fungi (AMF). These symbiotic fungi are particularly critical, given their key role in ecosystem processes affecting plant community structure and function. To better understand the consequences of nutrient deposition in plant communities, a long-term nutrient addition experiment was set up in a tropical montane forest in the Andes of southern Ecuador. In this study, we investigated the impacts of 7 years of elevated N and P on AMF root colonization potential (AMF-RCP) through a greenhouse bait plant method in which we quantified root colonization. We also examined the relationship between AMF-RCP and rarefied tree diversity. After 7 years of nutrient addition, AMF-RCP was negatively correlated with soil P, positively correlated with soil N, and positively correlated with rarefied tree diversity. Our results show that AMF in this tropical montane forest are directly affected by soil N and P concentrations, but may also be indirectly impacted by shifts in rarefied tree diversity. Our research also highlights the need to fully understand the benefits and drawbacks of using different sampling methods (e.g., AMF-RCP versus direct root sampling) to robustly examine AMF-plant interactions in the future. [ABSTRACT FROM AUTHOR]

Community analyses of arbuscular mycorrhizal fungi ( AMF) using ribosomal small subunit ( SSU) or internal transcribed spacer ( ITS) DNA sequences often suffer from low resolution or coverage. We developed a novel sequencing based approach for a highly resolving and specific profiling of AMF communities., We took advantage of previously established AMF-specific PCR primers that amplify a c. 1.5-kb long fragment covering parts of SSU, ITS and parts of the large ribosomal subunit ( LSU), and we sequenced the resulting amplicons with single molecule real-time ( SMRT) sequencing., The method was applicable to soil and root samples, detected all major AMF families and successfully discriminated closely related AMF species, which would not be discernible using SSU sequences. In inoculation tests we could trace the introduced AMF inoculum at the molecular level. One of the introduced strains almost replaced the local strain(s), revealing that AMF inoculation can have a profound impact on the native community., The methodology presented offers researchers a powerful new tool for AMF community analysis because it unifies improved specificity and enhanced resolution, whereas the drawback of medium sequencing throughput appears of lesser importance for low-diversity groups such as AMF. [ABSTRACT FROM AUTHOR]

Aim: Little is known about how N and P co-limited ecosystems respond to single nutrient enrichment. This work assesses the susceptibility of above- and belowground ecosystem components and of their linkages in an N and P co-limited pasture to N- and P-enrichment. We tested if the plants' responses can be explained by the concept of serially linked nutrients introduced by Ågren (Ecol Lett 7:185-191, ). In this concept, the control of the growth rate by one nutrient is assumed to depend on the control of a different cellular process by another nutrient. Methods: We investigated the responses of shoot and root biomass and C:N:P stoichiometry of the grass Setaria sphacelata (Schumach.) to moderate N, P, and N + P application over 5 years. In addition, the effects of nutrient enrichment on soil nutrient pools, on arbuscular mycorrhizal fungi (AMF) as well as on microbial biomass, activity, and community structure (phospholipid fatty acids: PLFA) were tested. In order to evaluate the importance of different factors explaining microbial responses, we applied a likelihood-based information-theoretic approach. Results: The application of N + P increased aboveground grass biomass (+61 %). Root biomass was stimulated by P-treatment (+45 %). Grass C:N:P stoichiometry responded by altering the P-uptake (P-treatment) or by translocating P from shoot to root (N-treatment). In particular, root C:N and C:P stoichiometry decreased in P- and in N-treatment. Extractable fractions of soil C, N, and P were significantly affected by nutrient enrichment. P application increased the biomass of Gram-positive bacteria (+22 %) and the abundance of AMF (+46 %), however, results of the IT-approach suggested indirect effects of nutrient enrichment on microbes. Conclusions: The responses of the N and P co-limited pasture to particular nutrient enrichment support the concept of serially linked nutrients. The present study provides evidence for the fundamental importance of P for controlling resource allocation of plants in responses to nutrient enrichment. Resource allocation of the grass rather than direct effects of nutrient additions drives changes in AMF, microbial biomass, community structure, and activity. [ABSTRACT FROM AUTHOR]

Increased nitrogen (N) depositions expected in the future endanger the diversity and stability of ecosystems primarily limited by N, but also often co-limited by other nutrients like phosphorus (P). In this context a nutrient manipulation experiment (NUMEX) was set up in a tropical montane rainforest in southern Ecuador, an area identified as biodiversity hotspot. We examined impacts of elevated N and P availability on arbuscular mycorrhizal fungi (AMF), a group of obligate biotrophic plant symbionts with an important role in soil nutrient cycles. We tested the hypothesis that increased nutrient availability will reduce AMF abundance, reduce species richness and shift the AMF community toward lineages previously shown to be favored by fertilized conditions. NUMEX was designed as a full factorial randomized block design. Soil cores were taken after 2 years of nutrient additions in plots located at 2000 m above sea level. Roots were extracted and intraradical AMF abundance determined microscopically; the AMF community was analyzed by 454-pyrosequencing targeting the large subunit rDNA. We identified 74 operational taxonomic units (OTUs) with a large proportion of Diversisporales. N additions provoked a significant decrease in intraradical abundance, whereas AMF richness was reduced significantly by N and P additions, with the strongest effect in the combined treatment (39% fewer OTUs), mainly influencing rare species. We identified a differential effect on phylogenetic groups, with Diversisporales richness mainly reduced by N additions in contrast to Glomerales highly significantly affected solely by P. Regarding AMF community structure, we observed a compositional shift when analyzing presence/absence data following P additions. In conclusion, N and P additions in this ecosystem affect AMF abundance, but especially AMF species richness; these changes might influence plant community composition and productivity and by that various ecosystem processes. [ABSTRACT FROM AUTHOR]

Tropical regions are facing increasing atmospheric inputs of nutrients, which will have unknown consequences for the structure and functioning of these systems. Here, we show that Neotropical montane rainforests respond rapidly to moderate additions of N (50 kg ha-1 yr-1) and P (10 kg ha-1 yr-1). Monitoring of nutrient fluxes demonstrated that the majority of added nutrients remained in the system, in either soil or vegetation. N and P additions led to not only an increase in foliar N and P concentrations, but also altered soil microbial biomass, standing fine root biomass, stem growth, and litterfall. The different effects suggest that trees are primarily limited by P, whereas some processes--notably aboveground productivity--are limited by both N and P. Highly variable and partly contrasting responses of different tree species suggest marked changes in species composition and diversity of these forests by nutrient inputs in the long term. The unexpectedly fast response of the ecosystem to moderate nutrient additions suggests high vulnerability of tropical montane forests to the expected increase in nutrient inputs. [ABSTRACT FROM AUTHOR]