Species pool, local assembly processes: Disentangling the mechanisms determining bacterial α- and ß-diversity during forest secondary succession.

Across ecology, and particularly within microbial ecology, there is limited understanding how the generation and maintenance of diversity. Although recent work has shown that both local assembly processes and species pools are important in structuring microbial communities, the relative contributions of these mechanisms remain an important question. Moreover, the roles of local assembly processes and species pools are drastically different when explicitly considering the potential for saturation or unsaturation, yet this issue is rarely addressed. Thus, we established a conceptual model that incorporated saturation theory into the microbiological domain to advance the understanding of mechanisms controlling soil bacterial diversity during forest secondary succession. Conceptual model hypotheses were tested by coupling soil bacterial diversity, local assembly processes and species pools using six different forest successional chronosequences distributed across multiple climate zones. Consistent with the unsaturated case proposed in our conceptual framework, we found that species pool consistently affected α-diversity, even while local assembly processes on local richness operate. In contrast, the effects of species pool on ß-diversity disappeared once local assembly processes were taken into account, and changes in environmental conditions during secondary succession led to shifts in ß-diversity through mediation of the strength of heterogeneous selection. Overall, this study represents one of the first to demonstrate that most local bacterial communities might be unsaturated, where the effect of species pool on α-diversity is robust to the consideration of multiple environmental influences, but ß-diversity is constrained by environmental selection.

Canopy nitrogen deposition enhances soil ecosystem multifunctionality in a temperate forest.

Nitrogen (N) deposition affects ecosystem functions crucial to human health and well-being. However, the consequences of this scenario for soil ecosystem multifunctionality (SMF) in forests are poorly understood. Here, we conducted a long-term field experiment in a temperate forest in China, where N deposition was simulated by adding N above and under the canopies. We discover that canopy N addition promotes SMF expression, whereas understory N addition suppresses it. SMF was regulated by fungal diversity in canopy N addition treatments, which is largely due to the strong resistance to soil acidification and efficient resource utilization characteristics of fungi. While in understory N addition treatments, SMF is regulated by bacterial diversity, which is mainly because of the strong resilience to disturbances and fast turnover of bacteria. Furthermore, rare microbial taxa may play a more important role in the maintenance of the SMF. This study provides the first evidence that N deposition enhanced SMF in temperate forests and enriches the knowledge on enhanced N deposition affecting forest ecosystems. Given the divergent results from two N addition approaches, an innovative perspective of canopy N addition on soil microbial diversity-multifunctionality relationships is crucial to policy-making for the conservation of soil microbial diversity and sustainable ecosystem management under enhanced N deposition. In future research, the consideration of canopy N processes is essential for more realistic assessments of the effects of atmospheric N deposition in forests.

Drivers of nematode diversity in forest soils across climatic zones.

Nematodes are the most abundant multi-cellular animals in soil, influencing key processes and functions in terrestrial ecosystems. Yet, little is known about the drivers of nematode abundance and diversity in forest soils across climatic zones. This is despite forests covering approximately 30% of the Earth's land surface, providing many crucial ecosystem services but strongly varying in climatic conditions and associated ecosystem properties across biogeographic zones. Here, we collected nematode samples from 13 forests across a latitudinal gradient. We divided this gradient into temperate, warm-temperate and tropical climatic zones and found that, across the gradient, nematode abundance and diversity were mainly influenced by soil organic carbon content. However, mean annual temperature and total soil phosphorus content in temperate zones, soil pH in warm-temperate zones, and mean annual precipitation in tropical zones were more important in driving nematode alpha-diversity, biomass and abundance. Additionally, nematode beta-diversity was higher in temperate than in warm-temperate and tropical zones. Together, our findings demonstrate that the drivers of nematode diversity in forested ecosystems are affected by the spatial scale and climatic conditions considered. This implies that high resolution studies are needed to accurately predict how soil functions respond if climate conditions move beyond the coping range of soil organisms.

Depth-driven responses of microbial residual carbon to nitrogen addition approaches in a tropical forest: Canopy addition versus understory addition.

Canopies play an important role in nitrogen (N) redistribution in forest ecosystems, and ignoring the canopy's role might bias estimates of the ecological consequences of anthropogenic atmospheric N deposition. We investigated the effects of the approach of N addition (Canopy addition vs. Understory addition) and level of N addition (25 kg N ha-1yr-1 vs. 50 kg N ha-1yr-1) on microbial residual carbon (MRC) accumulation in topsoil and subsoil. We found that the response of MRC to both approach and level of N addition varied greatly with soil depth in a tropical forest over eight years of continuous N addition. Specifically, N addition enhanced the accumulation of fungal and total MRC and their contribution to soil organic C (SOC) pools in the topsoil, whereas it decreased the contribution of fungal and total MRC to SOC in the subsoil. The contrasting effects of N addition on MRC contribution at varying soil depths were associated with the distinct response of microbial residues production. Understory N addition showed overall greater effects on MRC accumulation than canopy N addition did. Our results suggest that the canopy plays an important role in buffering the impacts of anthropogenic atmospheric N deposition on soil C cycling in tropical forests. The depth-dependent response of microbial residues to N addition also highlights the urgent need for further studies of different response mechanisms at different soil depths.

Canopy and understory nitrogen addition have different effects on fine root dynamics in a temperate forest: implications for soil carbon storage.

Elucidating the effects of atmospheric nitrogen (N) deposition on fine root dynamics and the potential underlying mechanisms is required to understand the changes in belowground and aboveground carbon storage. However, research on these effects in forests has mostly involved direct understory addition of N and has ignored canopy interception and processing of N. Here, we conducted a field experiment comparing the effects of canopy addition of N (CAN) with those of understory addition of N (UAN) at three N-addition rates (0, 25 and 50 kg N ha-1 yr-1 ) on fine root dynamics in a temperate deciduous forest. Fine root production and biomass were significantly higher with CAN than with UAN. At the same N-addition rate, increases in fine root production with CAN were at least two-fold greater than with UAN. At the high N-addition rate and relative to the control, fine root biomass was significantly increased by CAN (by 23.5%) but was significantly decreased by UAN (by 12.2%). Our results indicate that traditional UAN may underestimate the responses of fine root dynamics to atmospheric N deposition in forest ecosystems. Canopy N processes should be considered for more realistic assessments of the effects of atmospheric N deposition in forests.

Multiyear Measurements on Δ17O of Stream Nitrate Indicate High Nitrate Production in a Temperate Forest.

Nitrification is a crucial step in ecosystem nitrogen (N) cycling, but scaling up from plot-based measurements of gross nitrification to catchments is difficult. Here, we employed a newly developed method in which the oxygen isotope anomaly (Δ17O) of nitrate (NO3-) is used as a natural tracer to quantify in situ catchment-scale gross nitrification rate (GNR) for a temperate forest from 2014 to 2017 in northeastern China. The annual GNR ranged from 71 to 120 kg N ha-1 yr-1 (average 94 ± 10 kg N ha-1 yr-1) over the 4 years in this forest. This result and high stream NO3- loss (4.2-8.9 kg N ha-1 yr-1) suggest that the forested catchment may have been N-saturated. At the catchment scale, the total N output of 10.7 kg N ha-1 yr-1, via leaching and gaseous losses, accounts for 56% of the N input from bulk precipitation (19.2 kg N ha-1 yr-1). This result indicates that the forested catchment is still retaining a large fraction of N from atmospheric deposition. Our study suggests that estimating in situ catchment-scale GNR over several years when combined with other conventional flux estimates can facilitate the understanding of N biogeochemical cycling and changes in the ecosystem N status.

Nitrogen deposition cancels out exotic earthworm effects on plant-feeding nematode communities.

The activity and spread of exotic earthworms often are spatially correlated with N deposition because both arise from human activities. Exotic earthworms, in turn, can also greatly affect soil abiotic and biotic properties, as well as related ecological processes. Previous studies showed, for example, that earthworms can counteract the detrimental effects of plant-feeding nematodes on plant growth. However, potential interactive effects of N deposition and exotic earthworms on ecosystems are poorly understood. We explored the changes in density of plant-feeding nematodes in response to the presence of exotic earthworms, and whether these changes are altered by elevated N deposition in a two-factorial field mesocosm experiment at the Heshan National Field Research Station of Forest Ecosystem, in southern China. Our results show that earthworm addition marginally significantly increased the density of exotic earthworms and significantly increased the mass of earthworm casts. The total density of plant-feeding nematodes was not significantly affected by exotic earthworms or N deposition. However, exotic earthworms tended to increase the density of plant-feeding nematode taxa that are less detrimental to plant growth (r-strategists), while they significantly reduced the density of more harmful plant-feeding nematodes (K-strategists). Importantly, these earthworm effects were restricted to the ambient N deposition treatment, and elevated N deposition cancelled out the earthworm effect. Although exotic earthworms and N deposition interactively altered foliar N : P ratio in the target tree species, this did not result in significant changes in shoot and root biomass in the short term. Overall, our study indicates that N deposition can cancel out exotic earthworm-induced reductions in the density of harmful plant-feeding nematodes. These results suggest that anthropogenic N deposition can alter biotic interactions between exotic and native soil organisms with potential implications for ecosystem functioning.

Tree diversity regulates soil respiration through elevated tree growth in a microcosm experiment.

Soil respiration is an essential component of carbon (C) cycling in terrestrial ecosystems. Despite increasing awareness of the significance of aboveground-belowground interactions, little is known about tree diversity effects on soil respiration and the underlying mechanisms. Here, we conducted a 105-days microcosm experiment in a climate chamber to test the effects of tree diversity (1, 2, and 4 species) on soil respiration. We expected tree diversity to affect soil respiration through changes in tree growth and surface litter decomposition (indicated by the litter mass loss). Our results show that soil respiration varied among the four focal tree species and increased with tree species richness. Path analysis revealed tree growth as the main mechanism driving soil respiration and explaining the tree diversity effect in this short-term experiment. Our results indicate that tree diversity and particular tree traits regulate C dynamics through balancing C storage (plant productivity) and C release (soil respiration).

Increased precipitation alters the effects of nitrogen deposition on soil bacterial and fungal communities in a temperate forest.

Anthropogenic nitrogen (N) deposition and increased precipitation are known to alter soil microbial communities. However, the combined effects of elevated N deposition and increased precipitation on soil microbial community dynamics and co-occurrence networks in temperate forests remain elusive. In this study, we conducted a field manipulation experiment by applying N solution and water to the forest canopy to simulate natural N deposition and increased precipitation in a temperate forest. We collected samples in the litter layer, organic soil layer, and mineral soil layer in 2018-2019 after 6-7 years of N and water treatments, and explored how elevated N deposition and increased precipitation regulate soil microbial diversity, community composition, and co-occurrence networks in different soil layers and at different sampling times. We found that the effects of N deposition and increased precipitation on soil microbial communities varied with soil layers and sampling times. Compared to the ambient environment, single canopy N addition (CN) or single canopy water addition (CW) did not affect bacterial Shannon diversity in the mineral soil layer in 2018, but the combined canopy N and water additions (CNW) decreased it in this layer at this time. CN increased fungal OTU richness in the organic and mineral soil layers in 2018; however, CW and CNW did not have an effect on it in the same layer at the same time. CW and CNW, but not CN, significantly affected bacterial and fungal community compositions in the litter layer in 2018 and in the organic soil layer in 2019. In contrast, CN, but not CW or CNW, significantly affected fungal community composition in the litter layer in 2019. CNW exhibited higher complexities of bacterial and fungal co-occurrence networks than CN and the ambient environment, indicating increased precipitation can strengthen the effect of N deposition on the complexity of bacterial and fungal co-occurrence networks. Our findings suggest that increased precipitation alters the effects of atmospheric N deposition on soil bacterial and fungal communities in this temperate forest, depending on soil layer and sampling time. Moreover, both bacterial and fungal community compositions are sensitive to increased precipitation, but the bacterial community composition is more sensitive to N deposition than the fungal community composition in the organic and mineral soil layers in this forest.

Zea mays cultivation, biochar, and arbuscular mycorrhizal fungal inoculation influenced lead immobilization.

Plant cultivation can influence the immobilization of heavy metals in soil. However, the roles of soil amendments and microorganisms in crop-based phytoremediation require further exploration. In this study, we evaluated the impact of Zea mays L. cultivation, biochar application, and arbuscular mycorrhizal fungi (AMF) inoculation on soil lead (Pb) immobilization. Our results indicated that biochar addition resulted in a significant, 42.00%, reduction in AMF colonization. Plant cultivation, AMF inoculation, and biochar addition all contributed to enhanced Pb immobilization, as evidenced by decreased levels of diethylenetriaminepentaacetic acid- and CaCl2-extractable Pb in the soil. Furthermore, soil subjected to plant cultivation with AMF and biochar displayed reduced concentrations of bioavailable Pb. Biochar addition altered the distribution of Pb fractions in the soil, transforming the acid-soluble form into the relatively inert reducible and oxidizable forms. Additionally, biochar, AMF, and their combined use promoted maize growth parameters, including height, stem diameter, shoot and root biomass, and phosphorus uptake, while simultaneously reducing the shoot Pb concentration. These findings suggest a synergistic effect in Pb phytostabilization. In summary, despite the adverse impact of biochar on mycorrhizal growth, cultivating maize with the concurrent use of biochar and AMF emerges as a recommended and effective strategy for Pb phytoremediation.IMPORTANCEHeavy metal contamination in soil is a pressing environmental issue, and phytoremediation has emerged as a sustainable approach for mitigating this problem. This study sheds light on the potential of maize cultivation, biochar application, and arbuscular mycorrhizal fungi (AMF) inoculation to enhance the immobilization of Pb in contaminated soil. The findings demonstrate that the combined use of biochar and AMF during maize cultivation can significantly improve Pb immobilization and simultaneously enhance maize growth, offering a promising strategy for sustainable and effective Pb phytoremediation practices. This research contributes valuable insights into the field of phytoremediation and its potential to address heavy metal pollution in agricultural soils.

Distinct effects of canopy vs understory and organic vs inorganic N deposition on root resource acquisition strategies of subtropical Moso bamboo plants.

Atmospheric nitrogen (N) deposition inevitably alters soil nutrient status, subsequently prompting plants to modify their root morphology (i.e., adopting a do-it-yourself strategy), mycorrhizal symbioses (i.e., outsourcing strategy), and root exudation (i.e., nutrient-mining strategy) linking with resource acquisition. However, how N deposition influences the integrated pattern of these resource-acquisition strategies remains unclear. Furthermore, most studies in forest ecosystems have focused on understory N and inorganic N deposition, neglecting canopy-associated processes (e.g., N interception and assimilation) and the impacts of organic N on root functional traits. In this study, we compared the effects of canopy vs understory, organic vs inorganic N deposition on eight root functional traits of Moso bamboo plants. Our results showed that N deposition significantly decreased arbuscular mycorrhizal fungi (AMF) colonization, altered root exudation rate and root foraging traits (branching intensity, specific root area, and length), but did not influence root tissue density and N concentration. Moreover, the impacts of N deposition on root functional traits varied significantly with deposition approach (canopy vs. understory), form (organic vs. inorganic), and their interaction, showing variations in both intensity and direction (positive/negative). Furthermore, specific root area and length were positively correlated with AMF colonization under canopy N deposition and root exudation rate in understory N deposition. Root trait variation under understory N deposition, but not under canopy N deposition, was classified into the collaboration gradient and the conservation gradient. These findings imply that coordination of nutrient-acquisition strategies dependent on N deposition approach. Overall, this study provides a holistic understanding of the impacts of N deposition on root resource-acquisition strategies. Our results indicate that the evaluation of N deposition on fine roots in forest ecosystems might be biased if N is added understory.

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As regulators of the surface land processes, soil fauna communities are the vital foundations for healthy terrestrial ecosystems. Soil fauna have been studied in China for more than 70 years. Great progresses have been achieved in exploring soil fauna species composition and geographical distribution patterns. Soil fauna eco-geography, as a bridge between soil fauna geographic patterns and ecosystem services, has a new development opportunity with the deep recognition of soil fauna ecological functions. Soil fauna eco-geography research could be partitioned into four dimensions including the spatio-temporal patterns of: 1) the apparent characteristics of soil fauna community, such as species composition, richness and abundance; 2) the intrinsic characteristics of soil fauna community, such as dietary and habits; 3) soil fauna-related biotic and abiotic interactions especially those indicating drivers of soil fauna community structure or shaping the roles of soil fauna in ecosystems; and 4) soil fauna-related or -regulated key ecological processes. Current studies focus solely on soil fauna themselves and their geographical distributions. To link soil fauna geography more closely with ecosystem services, we suggested that: 1) converting the pure biogeography studies to those of revealing the spatio-temporal patterns of the soil fauna-related or regulated key relationships and ecological processes;2) expanding the temporal and spatial scales in soil fauna geographical research;3) exploring the integrated analysis approach for soil fauna-related data with multi-scales, multi-factors, and multi-processes;and 4) establishing standard reference systems for soil fauna eco-geographical researches. Hence, the change patterns of ecological niche of soil fauna communities could be illustrated, and precision mani-pulations of soil fauna communities and their ecological functions would become implementable, which finally contributes to ecosystem health and human well-being.

Bioactive molecules in soil ecosystems: masters of the underground.

Complex biological and ecological processes occur in the rhizosphere through ecosystem-level interactions between roots, microorganisms and soil fauna. Over the past decade, studies of the rhizosphere have revealed that when roots, microorganisms and soil fauna physically contact one another, bioactive molecular exchanges often mediate these interactions as intercellular signal, which prepare the partners for successful interactions. Despite the importance of bioactive molecules in sustainable agriculture, little is known of their numerous functions, and improving plant health and productivity by altering ecological processes remains difficult. In this review, we describe the major bioactive molecules present in below-ground ecosystems (i.e., flavonoids, exopolysaccharides, antibiotics and quorum-sensing signals), and we discuss how these molecules affect microbial communities, nutrient availability and plant defense responses.

Deciphering roles of microbiota in arsenic biotransformation from the earthworm gut and skin.

Biotransformation mediated by microbes can affect the biogeochemical cycle of arsenic. However, arsenic biotransformation mediated by earthworm-related microorganisms has not been well explored, especially the role played by earthworm skin microbiota. Herein, we reveal the profiles of arsenic biotransformation genes (ABGs) and elucidate the microbial communities of the earthworm gut, skin, and surrounding soil from five different soil environments in China. The relative abundance of ABGs in the earthworm skin microbiota, which were dominated by genes associated with arsenate reduction and transport, was approximately three times higher than that in the surrounding soil and earthworm gut microbiota. The composition and diversity of earthworm skin microbiota differed significantly from those of the soil and earthworm gut, comprising a core bacterial community with a relative abundance of 96% Firmicutes and a fungal community with relative abundances of 50% Ascomycota and 13% Mucoromycota. In addition, stochastic processes mainly contributed to the microbial community assembly across all samples. Moreover, fungal genera such as Vishniacozyma and Oomyces were important mediators of ABGs involved in the biogeochemical cycle of arsenic. This is the first study to investigate earthworm skin as a reservoir of microbial diversity in arsenic biotransformation. Our findings broaden the current scientific knowledge of the involvement of earthworms in the arsenic biogeochemical cycle.

Effect of Arbuscular Mycorrhiza Fungus Diversispora eburnea Inoculation on Lolium perenne and Amorpha fruticosa Growth, Cadmium Uptake, and Soil Cadmium Speciation in Cadmium-Contaminated Soil.

Cadmium (Cd) pollution has become aggravated during the past decades of industrialization, severely endangering human health through its entry into the food chain. While it is well understood that arbuscular mycorrhizal fungi (AMF) have a strong ability to regulate plant growth and Cd uptake, studies investigating how they affect soil Cd speciation and influence Cd uptake are limited. We designed a pot experiment comprising two AMF-inoculant groups (inoculation with Diversispora eburnea or no inoculation), three Cd concentration levels (0, 5, and 15 mg/kg), and two plant species (Lolium perenne and Amorpha fruticosa) to study the effect of AMF Diversispora eburnea on plant growth, Cd uptake, and Cd speciation in the soil. The results revealed that L. perenne exhibited higher productivity and greater Cd uptake than A. fruticosa, regardless of AMF D. eburnea inoculation. However, AMF D. eburnea significantly altered soil Cd speciation by increasing the proportion of exchangeable Cd and decreasing residual Cd, resulting in Cd enrichment in the plant root organs and the elimination of Cd from the polluted soils. Our experiments demonstrate that inoculating plants with AMF D. eburnea is an effective alternative strategy for remediating Cd-contaminated soil.

Litter inputs exert greater influence over soil respiration and its temperature sensitivity than roots in a coniferous forest in north-south transition zone.

The changes in carbon inputs of litter and roots to forest soils caused by climate change will result in a serious cascade effect on soil respiration and its temperature sensitivity (Q10). To differentiate and quantify the effects of surface litter and living roots on soil respiration and Q10, and further explore the role of abiotic factors and microbial properties on soil respiration and Q10, a short-term (two years) detritus input and removal treatment experiment was conducted in a coniferous forest of central China. Soil temperature, soil moisture, C/N, microbial biomass and community composition were analyzed to explore the drive mechanisms of soil respiration and Q10 in response to carbon inputs. The results showed that litter addition increased soil respiration by 22 %, while litter or roots removal did not affect soil respiration, which might be ascribed to the "priming effects" mediated by fresh plant litter. We also found that litter addition increased Q10, while litter removal decreased Q10. Litter addition significantly enhanced the microbial biomass for any single functional group and altered soil microbial community composition. Structural equation model further proved that microbial biomass and community composition exerted stronger impacts on Q10 than do soil abiotic factors. Soil moisture, microbial biomass and community structure were main factors in predicting soil respiration. The study highlights the important role of litter inputs compared with living roots in carbon cycling in short-term and deepens our understanding on the complex relationships among soil respiration, soil micro-environment and microbial community composition.

Responses of soil microbial and nematode communities to aluminum toxicity in vegetated oil-shale-waste lands.

Both soil nematodes and microorganisms have been shown to be sensitive bioindicators of soil recovery in metal-contaminated habitats; however, the underlying processes are poorly understood. We investigated the relationship among soil microbial community composition, nematode community structure and soil aluminum (Al) content in different vegetated aluminum-rich ecosystems. Our results demonstrated that there were greater soil bacterial, fungal and arbuscular mycorrhizal fungal biomass in Syzygium cumini plantation, greater abundance of soil nematodes in Acacia auriculiformis plantation, and greater abundance of soil predatory and herbivorous nematodes in Schima wallichii plantation. The concentration of water-soluble Al was normally greater in vegetated than non-vegetated soil. The residual Al and total Al concentrations showed a significant decrease after planting S. cumini plantation onto the shale dump. Acid extractable, reducible and oxidisable Al concentrations were greater in S. wallichii plantation. Stepwise linear regression analysis suggests the concentrations of water-soluble Al and total Al content explain the most variance associated with nematode assembly; whereas, the abundance of early-successional nematode taxa was explained mostly by soil moisture, soil organic C and total N rather than the concentrations of different forms of Al. In contrast, no significant main effects of either Al or soil physico-chemical characteristics on soil microbial biomass were observed. Our study suggests that vegetation was the primary driver on soil nematodes and microorganisms and it also could regulate the sensitivity of bio-indicator role mainly through the alteration of soil Al and physico-chemical characteristics, and S. cumini is effective for amending the Al contaminated soils.

Canopy and Understory Nitrogen Addition Alters Organic Soil Bacterial Communities but Not Fungal Communities in a Temperate Forest.

Atmospheric nitrogen (N) deposition is known to alter soil microbial communities, but how canopy and understory N addition affects soil bacterial and fungal communities in different soil layers remains poorly understood. Conducting a 6-year canopy and understory N addition experiment in a temperate forest, we showed that soil bacterial and fungal communities in the organic layer exhibited different responses to N addition. The main effect of N addition decreased soil bacterial diversity and altered bacterial community composition in the organic layer, but not changed fungal diversity and community composition in all layers. Soil pH was the main factor that regulated the responses of soil bacterial diversity and community composition to N addition, whereas soil fungal diversity and community composition were mainly controlled by soil moisture and nutrient availability. In addition, compared with canopy N addition, the understory N addition had stronger effects on soil bacterial Shannon diversity and community composition but had a weaker effect on soil bacteria richness in the organic soil layer. Our study demonstrates that the bacterial communities in the organic soil layer were more sensitive than the fungal communities to canopy and understory N addition, and the conventional method of understory N addition might have skewed the effects of natural atmospheric N deposition on soil bacterial communities. This further emphasizes the importance of considering canopy processes in future N addition studies and simultaneously evaluating soil bacterial and fungal communities in response to global environmental changes.

Differential Mechanisms Drive Species Loss Under Artificial Shade and Fertilization in the Alpine Meadow of the Tibetan Plateau.

Fertilization is an effective management strategy to promote community biomass but can simultaneously reduce species diversity in many grassland systems. Shifts in competition for resources have been proposed to explain the decline in plant species diversity due to fertilization, yet the underlying mechanism driving species loss remains controversial. This uncertainty may be driven by variation in aboveground and belowground resource availability. However, experiments simultaneously manipulating both light availability and soil nutrients are rare. Using a 6-year field experiment to manipulate light availability (via shade cloth) and soil nutrients (via fertilizer addition), we tested this resource competition hypothesis in a species-rich alpine meadow by examining the variation of species traits associated with the capacity of light acquisition within these treatments. Our results showed that artificial shade decreased community biomass accumulation whereas fertilization increased it. In contrast, both shade and fertilization reduced species diversity. Extinction of non-Gramineae species (e.g., Fabaceae and Cyperaceae) was the main reason for species diversity decline. Species loss can be explained by the limitation of light availability and predicted by species traits associated with light acquisition capability under fertilization and low light tolerance under artificial shade. Specifically, fertilization eliminated species with lower stature and artificial shade exterminated species with the higher light compensation point (LCP). The findings suggest that light availability is consistently important for plant growth and that low competitiveness for light under fertilization and intolerance of low light conditions under artificial shade trigger species loss process in the alpine meadow. Our experiment helps clarify the mechanisms of how artificial shade and fertilization decreased species diversity and highlight that LCP, which tends to be neglected by most of the studies, is one of the vital drivers in determining species coexistence.

Long-term water use efficiency and non-structural carbohydrates of dominant tree species in response to nitrogen and water additions in a warm temperate forest.

Nitrogen (N) deposition tends to accompany precipitation in temperate forests, and vegetation productivity is mostly controlled by water and N availability. Many studies showed that tree species response to precipitation or N deposition alone influences, while the N deposition and precipitation interactive effects on the traits of tree physiology, especially in non-structural carbohydrates (NSCs) and long-term water use efficiency (WUE), are still unclear. In this study, we measured carbon stable isotope (δ13C), total soluble sugar and starch content, total phenols, and other physiological traits (e.g., leaf C:N:P stoichiometry, lignin, and cellulose content) of two dominant tree species (Quercus variabilis Blume and Liquidambar formosana Hance) under canopy-simulated N deposition and precipitation addition to analyze the changes of long-term WUE and NSC contents and to explain the response strategies of dominant trees to abiotic environmental changes. This study showed that N deposition decreased the root NSC concentrations of L. formosana and the leaf lignin content of Q. variabilis. The increased precipitation showed a negative effect on specific leaf area (SLA) and a positive effect on leaf WUE of Q. variabilis, while it increased the leaf C and N content and decreased the leaf cellulose content of L. formosana. The nitrogen-water interaction reduced the leaf lignin and total phenol content of Q. variabilis and decreased the leaf total phenol content of L. formosana, but it increased the leaf C and N content of L. formosana. Moreover, the response of L. formosana to the nitrogen-water interaction was greater than that of Q. variabilis, highlighting the differences between the two dominant tree species. The results showed that N deposition and precipitation obviously affected the tree growth strategies by affecting the NSC contents and long-term WUE. Canopy-simulated N deposition and precipitation provide a new insight into the effect of the nitrogen-water interaction on tree growth traits in a temperate forest ecosystem, enabling a better prediction of the response of dominant tree species to global change.