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.

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.

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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.

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.

Responses of earthworm Metaphire vulgaris gut microbiota to arsenic and nanoplastics contamination.

The growing contamination of arsenic and plastics has severely effects on the soil fauna health, including shifts of gut microbiota community. A few studies have focused on effects of microplastics and metal(loid) in soil and fauna gut microbiome. However, the environmental effect of nanoplastics and arsenic on the earthworm gut microbiota, especially on arsenic biotransformation in the gut, remain largely unknown. Here, a microcosm study was performed to explore the effects of nanoplastics and arsenic on the microbiota characteristics in earthworm Metaphire vulgaris gut using Illumina high throughput sequencing, and to investigate changes in the gut microbiota-mediated arsenic biotransformation genes (ABGs) by using high-throughput quantitative PCR. Our results demonstrated that the concentration of arsenic in the earthworm body tissues after exposure to arsenic and nanoplastics was significantly lower from that with arsenic alone exposure. Moreover, the clearly different bacterial community was observed in the soil compared with the earthworm gut, which was dominated by Proteobacteria, Actinobacteria, and Firmicutes at phylum level. Arsenic exposure significantly disturbed bacterial community structure in the earthworm gut, but exposure to nanoplastics did not induce gut microbiota changes. More interestingly, nanoplastics can relieve adverse effect of arsenic on the gut microbiota possibly by adsorbing arsenic. In addition, a total of 16 ABGs were detected, and predominant genes involved in arsenic reduction and transport process were observed in the earthworm guts. In short, this study provides a new picture of the effects of nanoplastics and arsenic on the gut microbiota and arsenic biotransformation in soil fauna gut.

Drought promotes soil phosphorus transformation and reduces phosphorus bioavailability in a temperate forest.

Drought can substantially alter ecosystem functions, especially biogeochemical cycles of key nutrients. As an essential but often limiting nutrient, P plays a central role in critical ecosystem processes (i.e. primary productivity). However, little is known about how drought can affect the soil phosphorus (P) cycle and its bioavailability in forest ecosystems. Here, we conducted a four-year field drought experiment using throughfall reduction approach to examine how drought can alter soil P dynamics and bioavailability in a warm temperate forest. We found that the P held in calcium phosphate was significantly decreased under drought, which was accompanied by the increases of inorganic and organic P bound with secondary minerals (Fe/Al oxides). These drought-induced P transformations can be well explained by the soil pH. The significant decline in soil pH under drought can drive the solubilization of P held in calcium phosphate. Our study further showed that drought directly decreased soil P bioavailability and altered the potential mechanisms of the replenishment of inorganic P into the soil solution. The potential of the inorganic P release driven by protons was reduced, while inorganic P release potentials driven by enzyme and organic acid were increased under drought. Therefore, our results strongly suggested that drought can significantly alter the soil P biogeochemical cycles and change the biological mechanisms underlying P bioavailability.

Earthworm Pontoscolex corethrurus stimulated soil CO2 emission by enhancing substrate availability rather than changing microbiota community structure.

Earthworms may regulate carbon (C) mineralization through a top-down process by altering microbiota community and/or a bottom-up process by providing basic conditions such as mineralizable substrates. However, these two roles performed by earthworms have not been separately quantified. Here, we focused on how the pantropical widespread earthworm, Pontoscolex corethrurus, affected CO2 emission by changing soil microbiota community and substrate availability in infertile subtropical soils. Two experiments were performed. Firstly, we conducted a 3-year field experiment wherein P. corethrurus population was manipulated by electrical shocking. The two treatments were earthworm reduction (ER) and earthworm addition (EA). Thereby, we tried to understand CO2 emission pattern through the earthworm-induced changes in soil microbiota community and C-related enzyme activities in field plots. Secondly, a cross-soil inoculation microcosm experiment was conducted to partition the contributions of earthworm-regulated microbiota community and substrate quality to CO2 emission. The four treatments were 1) autoclaved ER soil + ER microbiota, 2) autoclaved EA soil + ER microbiota, 3) autoclaved ER soil + EA microbiota, and 4) autoclaved EA soil + EA microbiota. We found that, in the field experiment, earthworm addition changed soil microbiota community structure, but increased CO2 emission despite decreasing soil ß-glucosidase activity by 12%. In the cross-inoculation experiment, the ß-glucosidase activity and CO2 emission (1-day incubation) in EA soils was 65.5% and 35.5% greater than that in the ER soils, respectively; the cumulative CO2 emission (30-day incubation) in EA soils was also significantly greater than that in ER soils. However, no significant effects of microbiota inoculation on soil CO2 emissions were observed. These results suggested that the earthworm-enhanced substrate availability, rather than the earthworm-induced changing in the microbiota community structure and enzymatic activities, played a key role in C mineralization. This study implies that P. corethrurus occupies a "bottom niche" in infertile subtropical soils.

Exotic earthworms maintain soil biodiversity by altering bottom-up effects of plants on the composition of soil microbial groups and nematode communities.

Bottom-up effects of plants on soil communities can be modified by the activity of exotic earthworms, by altering resource availability for soil food webs through feeding, burrowing, and casting activities. The present study explored effects of plants (planting of shrubs) on soil micro-food webs (composition of soil microbial and nematode communities), and whether these effects were altered by the activity of exotic earthworms (exotic earthworms addition). Planted shrubs resulted in a non-significant increase of bacterial biomass and significantly increased the abundance of different nematode trophic groups and total nematode biomass, indicating that planted shrubs had significant bottom-up effects on soil bacteria and nematodes. Planted shrubs decreased nematode diversity, evenness, and richness, but increased nematode dominance in the plots where the abundance of exotic earthworms was not amended. By contrast, these effects of shrub presence on soil biodiversity were not found in the plots that received exotic earthworms. In addition, planted shrubs increased the total energy flux to the nematode community. By contrast, the elevated activity of exotic earthworms mitigated the increase in total energy flux to nematodes in the presence of shrubs, and increased the ratio of fungal to bacterial PLFAs. Both of these changes indicate reduced energy flux in the plots with added exotic earthworms. Nematode diversity decreased, while nematode dominance increased with increasing total energy flux to nematodes, probably because few species benefited from high energy flux. Our study indicates that exotic earthworms can maintain soil biodiversity by reducing the energy flux through soil food webs.

An alternative approach to reduce algorithm-derived biases in monitoring soil organic carbon changes.

Quantifying soil organic carbon (SOC) changes is a fundamental issue in ecology and sustainable agriculture. However, the algorithm-derived biases in comparing SOC status have not been fully addressed. Although the methods based on equivalent soil mass (ESM) and mineral-matter mass (EMMM) reduced biases of the conventional methods based on equivalent soil volume (ESV), they face challenges in ensuring both data comparability and accuracy of SOC estimation due to unequal basis for comparison and using unconserved reference systems. We introduce the basal mineral-matter reference systems (soils at time zero with natural porosity but no organic matter) and develop an approach based on equivalent mineral-matter volume (EMMV). To show the temporal bias, SOC change rates were recalculated with the ESV method and modified methods that referenced to soils at time t1 (ESM, EMMM, and EMMV-t1) or referenced to soils at time zero (EMMV-t0) using two datasets with contrasting SOC status. To show the spatial bias, the ESV- and EMMV-t0-derived SOC stocks were compared using datasets from six sites across biomes. We found that, in the relatively C-rich forests, SOC accumulation rates derived from the modified methods that referenced to t1 soils and from the EMMV-t0 method were 5.7%-13.6% and 20.6% higher than that calculated by the ESV method, respectively. Nevertheless, in the C-poor lands, no significant algorithmic biases of SOC estimation were observed. Finally, both the SOC stock discrepancies (ESV vs. EMMV-t0) and the proportions of this unaccounted SOC were large and site-dependent. These results suggest that although the modified methods that referenced to t1 soils could reduce the biases derived from soil volume changes, they may not properly quantify SOC changes due to using unconserved reference systems. The EMMV-t0 method provides an approach to address the two problems and is potentially useful since it enables SOC comparability and integrating SOC datasets.

An increase in precipitation exacerbates negative effects of nitrogen deposition on soil cations and soil microbial communities in a temperate forest.

World soils are subjected to a number of anthropogenic global change factors. Although many previous studies contributed to understand how single global change factors affect soil properties, there have been few studies aimed at understanding how two naturally co-occurring global change drivers, nitrogen (N) deposition and increased precipitation, affect critical soil properties. In addition, most atmospheric N deposition and precipitation increase studies have been simulated by directly adding N solution or water to the forest floor, and thus largely neglect some key canopy processes in natural conditions. These previous studies, therefore, may not realistically simulate natural atmospheric N deposition and precipitation increase in forest ecosystems. In a field experiment, we used novel canopy applications to investigate the effects of N deposition, increased precipitation, and their combination on soil chemical properties and the microbial community in a temperate deciduous forest. We found that both soil chemistry and microorganisms were sensitive to these global change factors, especially when they were simultaneously applied. These effects were evident within 2 years of treatment initiation. Canopy N deposition immediately accelerated soil acidification, base cation depletion, and toxic metal accumulation. Although increased precipitation only promoted base cation leaching, this exacerbated the effects of N deposition. Increased precipitation decreased soil fungal biomass, possible due to wetting/re-drying stress or to the depletion of Na. When N deposition and increased precipitation occurred together, soil gram-negative bacteria decreased significantly, and the community structure of soil bacteria was altered. The reduction of gram-negative bacterial biomass was closely linked to the accumulation of the toxic metals Al and Fe. These results suggested that short-term responses in soil cations following N deposition and increased precipitation could change microbial biomass and community structure.

Plant-facilitated effects of exotic earthworm Pontoscolex corethrurus on the soil carbon and nitrogen dynamics and soil microbial community in a subtropical field ecosystem.

Earthworms and plants greatly affect belowground properties; however, their combined effects are more attractive based on the ecosystem scale in the field condition. To address this point, we manipulated earthworms (exotic endogeic species Pontoscolex corethrurus) and plants (living plants [native tree species Evodia lepta] and artificial plants) to investigate their combined effects on soil microorganisms, soil nutrients, and soil respiration in a subtropical forest. The manipulation of artificial plants aimed to simulate the physical effects of plants (e.g., shading and interception of water) such that the biological effects of plants could be evaluated separately. We found that relative to the controls, living plants but not artificial plants significantly increased the ratio of fungal to bacterial phospholipid fatty acids (PLFAs) and fungal PLFAs. Furthermore, earthworms plus living plants significantly increased the soil respiration and decreased the soil NH 4+-N, which indicates that the earthworm effects on the associated carbon, and nitrogen processes were greatly affected by living plants. The permutational multivariate analysis of variance results also indicated that living plants but not earthworms or artificial plants significantly changed the soil microbial community. Our results suggest that the effects of plants on soil microbes and associated soil properties in this study were largely explained by their biological rather than their physical effects.

Invariant community structure of soil bacteria in subtropical coniferous and broadleaved forests.

Soil bacteria may be influenced by vegetation and play important roles in global carbon efflux and nutrient cycling under global changes. Coniferous and broadleaved forests are two phyletically distinct vegetation types. Soil microbial communities in these forests have been extensively investigated but few studies have presented comparable data regarding the characteristics of bacterial communities in subtropical forests. We investigated soil bacterial biomass and community composition in three pairs of coniferous and broadleaved forests across a subtropical climatic gradient. We found that bacterial biomass differed between the coniferous and broadleaved forests across the subtropical climate gradient; however, this difference disappeared at some individual sites. In contrast, the same 90 bacterial genera were found in both forest types, and their relative abundances didn't differ between the forest types, with the exception of one genus that was more abundant in broadleaved forests. Soil nitrogen or moisture was associated with bacterial groups in the coniferous and broadleaved forests, respectively. Thus, we inferred that these forests can respond differently to future changes in nitrogen deposition or precipitation. This study highlights soil bacterial invariant community composition in contrasting subtropical forests and provides a new perspective on the potential response and feedback of forests to global changes.

Consistent effects of canopy vs. understory nitrogen addition on the soil exchangeable cations and microbial community in two contrasting forests.

Anthropogenic N deposition has been well documented to cause substantial impacts on the chemical and biological properties of forest soils. In most studies, however, atmospheric N deposition has been simulated by directly adding N to the forest floor. Such studies thus ignored the potentially significant effect of some key processes occurring in forest canopy (i.e., nitrogen retention) and may therefore have incorrectly assessed the effects of N deposition on soils. Here, we conducted an experiment that included both understory addition of N (UAN) and canopy addition of N (CAN) in two contrasting forests (temperate deciduous forest vs. subtropical evergreen forest). The goal was to determine whether the effects on soil exchangeable cations and microbial biomass differed between CAN and UAN. We found that N addition reduced pH, BS (base saturation) and exchangeable Ca and increased exchangeable Al significantly only at the temperate JGS site, and reduced the biomass of most soil microbial groups only at the subtropical SMT site. Except for soil exchangeable Mn, however, effects on soil chemical properties and soil microbial community did not significantly differ between CAN and UAN. Although biotic and abiotic soil characteristics differ significantly and the responses of both soil exchangeable cations and microbial biomass were different between the two study sites, we found no significant interactive effects between study site and N treatment approach on almost all soil properties involved in this study. In addition, N addition rate (25 vs. 50 kg N ha(-1) yr(-1)) did not show different effects on soil properties under both N addition approaches. These findings did not support previous prediction which expected that, by bypassing canopy effects (i.e., canopy retention and foliage fertilization), understory addition of N would overestimate the effects of N deposition on forest soil properties, at least for short time scale.

Response of soil respiration and ecosystem carbon budget to vegetation removal in Eucalyptus plantations with contrasting ages.

Reforested plantations have substantial effects on terrestrial carbon cycling due to their large coverage area. Although understory plants are important components of reforested plantations, their effects on ecosystem carbon dynamics remain unclear. This study was designed to investigate the effects of vegetation removal/understory removal and tree girdling on soil respiration and ecosystem carbon dynamics in Eucalyptus plantations of South China with contrasting ages (2 and 24 years old). We conducted a field manipulation experiment from 2008 to 2009. Understory removal reduced soil respiration in both plantations, whereas tree girdling decreased soil respiration only in the 2-year-old plantations. The net ecosystem production was approximately three times greater in the 2-year-old plantations (13.4 t C ha(-1) yr(-1)) than in the 24-year-old plantations (4.2 t C h(-1) yr(-1)). The biomass increase of understory plants was 12.6 t ha(-1) yr(-1) in the 2-year-old plantations and 2.9 t ha(-1) yr(-1) in the 24-year-old plantations, accounting for 33.9% nd 14.1% of the net primary production, respectively. Our findings confirm the ecological importance of understory plants in subtropical plantations based on the 2 years of data. These results also indicate that Eucalyptus plantations in China may be an important carbon sink due to the large plantation area.

Earthworms facilitate carbon sequestration through unequal amplification of carbon stabilization compared with mineralization.

A recent review concluded that earthworm presence increases CO2 emissions by 33% but does not affect soil organic carbon stocks. However, the findings are controversial and raise new questions. Here we hypothesize that neither an increase in CO2 emission nor in stabilized carbon would entirely reflect the earthworms' contribution to net carbon sequestration. We show how two widespread earthworm invaders affect net carbon sequestration through impacts on the balance of carbon mineralization and carbon stabilization. Earthworms accelerate carbon activation and induce unequal amplification of carbon stabilization compared with carbon mineralization, which generates an earthworm-mediated 'carbon trap'. We introduce the new concept of sequestration quotient to quantify the unequal processes. The patterns of CO2 emission and net carbon sequestration are predictable by comparing sequestration quotient values between treatments with and without earthworms. This study clarifies an ecological mechanism by which earthworms may regulate the terrestrial carbon sink.