Dima Chen

Evidence that acidification‐induced declines in plant diversity and productivity are mediated by changes in below‐ground communities and soil properties in a semi‐arid steppe

Summary 1. Anthropogenic acid deposition–induced soil acidification is one of the major threats to biodiversity, ecosystem functioning and services. Few studies, however, have explored in detail how above-ground changes in plant species richness and productivity resulting from soil acidification are mediated by effects on below-ground biota and soil properties. 2. To increase our understanding of this linkage, we collected data on below- and above-ground communities and soil properties in a 3-year field experiment with seven levels of acid addition rate to build-up broad intensities of soil acidification in the semi-arid Inner Mongolian grassland. 3. Acid addition directly elevated concentrations of soil Al 3+ ions, decreased the base cations Ca 2+ , Mg 2+ and Na + , and increased soil moisture and available phosphorus. Acid addition also appears to have altered the soil microbial community via changes in H + and Al 3+ ions and altered the nematode community via changes in H + ions and soil moisture. 4. The observed changes in soil N availability (i.e. net N mineralization, NO � -N and NH þ -N) could be explained by mediating changes in the H + and Al 3+ ions, microbial community (i.e. community structure, bacteria and fungi/bacteria as indicated by phospholipid fatty acids analysis) and the nematode community (i.e. total abundance, taxa richness and maturity index). 5. Declines in plant species richness and productivity were greater at high intensities of soil acidification in the second sampling year than in the first sampling year. The changes in plant community observed were mostly explained by soil nutrient pathways (e.g. N availability or base mineral cations), which were in turn regulated by the soil microbial or nematode communities as well as by the direct effects of the increase in H + or Al 3+ ions. 6. Synthesis. Our results suggest that the below-ground microbial and nematode communities are more sensitive to soil acidification than the plant communities are, and further that soil acidification–induced changes in plants are mediated by changes in below-ground communities and soil nutrients. These findings improve our understanding of the links between below- and above-ground communities in the Inner Mongolia grassland, especially in the context of anthropogenic acid enrichment.

Soil acidification exerts a greater control on soil respiration than soil nitrogen availability in grasslands subjected to long‐term nitrogen enrichment

Summary Terrestrial ecosystems worldwide are receiving increasing amounts of biologically reactive nitrogen (N) as a consequence of anthropogenic activities. This intended or unintended fertilization can have a wide-range of impacts on biotic communities and hence on soil respiration. Reduction in below-ground carbon (C) allocation induced by high N availability has been assumed to be a major mechanism determining the effects of N enrichment on soil respiration. In addition to increasing available N, however, N enrichment causes soil acidification, which may also affect root and microbial activities. The relative importance of increased N availability vs. soil acidification on soil respiration in natural ecosystems experiencing N enrichment is unclear. We conducted a 12-year N enrichment experiment and a 4-year complementary acid addition experiment in a semi-arid Inner Mongolian grassland. We found that N enrichment had contrasting effects on root and microbial respiration. N enrichment significantly increased root biomass, root N content and specific root respiration, thereby promoting root respiration. In contrast, N enrichment significantly suppressed microbial respiration likely by reducing total microbial biomass and changing the microbial community composition. The effect on root activities was due to both soil acidity and increased available N, while the effect on microbes primarily stemmed from soil acidity, which was further confirmed by results from the acid addition experiment. Our results indicate that soil acidification exerts a greater control than soil N availability on soil respiration in grasslands experiencing long-term N enrichment. These findings suggest that N-induced soil acidification should be included in predicting terrestrial ecosystem C balance under future N deposition scenarios.

Vertebrate herbivore‐induced changes in plants and soils: linkages to ecosystem functioning in a semi‐arid steppe

Summary 1. Large grazing herbivores have been reported to determine the structure and function of grassland ecosystems. However, the ecological linkages between structure and functioning components have yet been thoroughly explored. 2. Here, we test the hypothesis of the impact of grazing on soil nematode community (e.g. structure and composition) and linkages to ecosystem functioning (e.g. soil N mineralization and ANPP) via changes in pathways of plant community, soil nutrients and soil environment using a field experiment maintained for 5 years with seven levels of grazing intensity in the Inner Mongolian grassland. 3. A structural equation model (SEM) with nematode abundances as response variables showed that plant-feeding and fungal-feeding nematodes were driven by changes in the plant community, and bacterial-feeding nematodes were affected by soil abiotic nutrients and environment, while omnivorous + carnivorous nematodes were altered by soil environment and bacterial-feeding nematodes. This indicates that the top-down control by grazing leads to bottom-up control in the soil food web. 4. We found that grazing affected the ecosystem functioning via different pathways. Grazing effects soil N mineralization by changing plant community, soil nutrients, soil environment and nematodes community structure, while it affects ANPP by altering soil N mineralization and soil environment. 5. Our findings could provide a better understanding of the responses of plants and soils to grazing and the linkages between structure and functioning of above-ground and belowground in the semi-arid steppe.

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% and 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.

Effects of plant functional group loss on soil biota and net ecosystem exchange: a plant removal experiment in the Mongolian grassland

Summary The rapid loss of global biodiversity can greatly affect the functioning of above-ground components of ecosystems. However, how such biodiversity losses affect below-ground communities and linkages to soil carbon (C) sequestration is unclear. Here, we describe how losses in plant functional groups (PFGs) affect soil microbial and nematode communities and net ecosystem exchange (NEE) in a 4-year removal experiment conducted on the Mongolian plateau, the worlds largest remaining natural grassland. Our results demonstrated that the biomasses or abundances of most components of the two below-ground communities (microbes and nematodes) were negatively affected by PFG loss and were positively related to above-ground plant biomass. The removal of dominant PFGs (perennial bunchgrasses and perennial rhizomatous grasses) reduced the biomass or abundance of below-ground community components while removal of less dominant PFGs (perennial forbs and annuals/biennials) did not change or increased the biomass or abundance of below-ground community components. The biomass-based ratio of fungal to bacterial microbes and the number-based ratio of fungal-feeding to bacterial-feeding nematodes decreased with increasing PFG losses. Variation partitioning analyses showed that the identity of PFGs together with above-ground plant biomass explained most of the total variation in soil microbes and that the identity of PFGs and above-ground plant biomass together with nematode food resources explained most of the total variation in soil nematodes. The increase in NEE with PFG loss was mainly explained by decreases in above-ground plant biomass and the ratio of fungi to bacteria. Synthesis. The shift of below-ground communities from a fungal-based to a bacterial-based energy channel as PFG richness decreases indicates that less diverse grassland ecosystems will have lower nutrient retention and hence be more sensitive to land-use or climate change. The dominant effects of above-ground plant biomass and below-ground communities on NEE indicate that PFG loss resulting from land-use or climate change has the potential to reduce C sequestration in semi-arid grassland soils. These findings suggest that predictive models may need to consider the composition of above-ground and below-ground communities in order to accurately simulate the dynamics of CO2 fluxes in terrestrial ecosystems.

Functional correlations between specific leaf area and specific root length along a regional environmental gradient in Inner Mongolia grasslands

Summary 1. Among above- and below-ground traits, specific leaf area (SLA, cm 2 g � 1 ) and specific root length (SRL, m g � 1 ) are the two key traits reflecting species resource acquisition strategies. However, patterns of variation in SLA and SRL have rarely been examined simultaneously across evolutionary history and environmental gradients, and the SLA–SRL relationship is still controversial on several grounds. 2. We examined the inter- and intraspecific variations in SLA and SRL of different root branching orders and the SLA–SRL relationship across 55 species and 21 plant communities of four vegetation types along a 2000-km transect in the Inner Mongolia grassland. 3. With increasing root branching order, the interspecific variation in SRL increased, but the intraspecific variation in SRL decreased considerably, and the form of SLA–SRL relationship shifted from positive to negative. This indicates that acquisition of soil resources (e.g. water and nutrients) is a fundamental strategy for plant investment to root length. When inter- and intraspecific variations in SLA and SRLs were partitioned into alpha (within-community) and beta (among-community) components, the alpha component exhibited substantially greater inter- and intraspecific variations than the beta component. 4. Across the transect, the evolutionarily late diverged species in phylogenetic tree evolved towards low SLA and SRL-1 (SRL for first-order roots) and tended to distribute in resourcepoor conditions along the environmental gradient. The early diverged species, in contrast, had high SLA and SRL-1 and mostly distributed in resource-rich conditions. 5. Our findings suggest that patterns of inter- and intraspecific variations in SLA and SRL of different root branching orders and the form of SLA–SRL relationship could be well explained by within- and among-community filtering processes and species divergence time. Coordination and trade-offs between leaves and roots do not mutually exclude but operate simultaneously at different scales and among different root branching orders in arid and semi-arid grasslands.

Plant diversity enhances productivity and soil carbon storage

Significance Soil carbon sequestration plays an important role in mitigating anthropogenic increases in atmospheric CO2 concentrations. Recent studies have shown that biodiversity increases soil organic carbon (SOC) storage in experimental grasslands. However, the effects of species diversity on SOC storage in natural ecosystems have rarely been studied, and the potential mechanisms are yet to be understood. The results presented here show that favorable climate conditions, particularly high precipitation, tend to increase both species richness and belowground biomass, which had a consistent positive effect on SOC storage in forests, shrublands, and grasslands. Nitrogen deposition and soil pH generally have a direct negative effect on SOC storage. Ecosystem management that maintains high levels of plant diversity can enhance SOC storage and other ecosystem services that depend on plant diversity. Despite evidence from experimental grasslands that plant diversity increases biomass production and soil organic carbon (SOC) storage, it remains unclear whether this is true in natural ecosystems, especially under climatic variations and human disturbances. Based on field observations from 6,098 forest, shrubland, and grassland sites across China and predictions from an integrative model combining multiple theories, we systematically examined the direct effects of climate, soils, and human impacts on SOC storage versus the indirect effects mediated by species richness (SR), aboveground net primary productivity (ANPP), and belowground biomass (BB). We found that favorable climates (high temperature and precipitation) had a consistent negative effect on SOC storage in forests and shrublands, but not in grasslands. Climate favorability, particularly high precipitation, was associated with both higher SR and higher BB, which had consistent positive effects on SOC storage, thus offsetting the direct negative effect of favorable climate on SOC. The indirect effects of climate on SOC storage depended on the relationships of SR with ANPP and BB, which were consistently positive in all biome types. In addition, human disturbance and soil pH had both direct and indirect effects on SOC storage, with the indirect effects mediated by changes in SR, ANPP, and BB. High soil pH had a consistently negative effect on SOC storage. Our findings have important implications for improving global carbon cycling models and ecosystem management: Maintaining high levels of diversity can enhance soil carbon sequestration and help sustain the benefits of plant diversity and productivity.

Patterns of plant carbon, nitrogen, and phosphorus concentration in relation to productivity in China’s terrestrial ecosystems

Significance Estimates of nutrient allocation in different plant tissues and the relationships between the nutrient contents and photosynthetic capacity are critical to predicting ecosystem carbon sequestration under global change. Here, we provide an assessment of large-scale patterns of community-level nitrogen and phosphorus concentrations in different plant tissues and then examine how nutrient allocations are coupled with plant productivity. The results show that nutrient concentrations in leaves are less responsive to abiotic environments than those in woody stems and roots (stable leaf nutrient concentration hypothesis); the relationships between vegetation primary productivity and leaf nutrient contents are stronger when less nutrients are allocated to the woody tissues (productivity–nutrient allocation hypothesis) and are stronger in deciduous than in evergreen vegetation (productivity–leaf lifespan hypothesis). Plant nitrogen (N) and phosphorus (P) content regulate productivity and carbon (C) sequestration in terrestrial ecosystems. Estimates of the allocation of N and P content in plant tissues and the relationship between nutrient content and photosynthetic capacity are critical to predicting future ecosystem C sequestration under global change. In this study, by investigating the nutrient concentrations of plant leaves, stems, and roots across China’s terrestrial biomes, we document large-scale patterns of community-level concentrations of C, N, and P. We also examine the possible correlation between nutrient content and plant production as indicated by vegetation gross primary productivity (GPP). The nationally averaged community concentrations of C, N, and P were 436.8, 14.14, and 1.11 mg·g−1 for leaves; 448.3, 3.04 and 0.31 mg·g−1 for stems; and 418.2, 4.85, and 0.47 mg·g−1 for roots, respectively. The nationally averaged leaf N and P productivity was 249.5 g C GPP·g-1 N·y−1 and 3,157.9 g C GPP·g–1 P·y−1, respectively. The N and P concentrations in stems and roots were generally more sensitive to the abiotic environment than those in leaves. There were strong power-law relationships between N (or P) content in different tissues for all biomes, which were closely coupled with vegetation GPP. These findings not only provide key parameters to develop empirical models to scale the responses of plants to global change from a single tissue to the whole community but also offer large-scale evidence of biome-dependent regulation of C sequestration by nutrients.

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.

Reconciling multiple impacts of nitrogen enrichment on soil carbon: plant, microbial and geochemical controls

Impacts of reactive nitrogen (N) inputs on ecosystem carbon (C) dynamics are highly variable, and the underlying mechanisms remain unclear. Here, we proposed a new conceptual framework that integrates plant, microbial and geochemical mechanisms to reconcile diverse and contrasting impacts of N on soil C. This framework was tested using long-term N enrichment and acid addition experiments in a Mongolian steppe grassland. Distinct mechanisms could explain effects of N on particulate and mineral-associated soil C pools, potentially explaining discrepancies among previous N addition studies. While plant production predominated particulate C changes, N-induced soil acidification strongly affected mineral-associated C through decreased microbial growth and pH-sensitive associations between iron and aluminium minerals and C. Our findings suggest that effects of N-induced acidification on microbial respiration and geochemical properties should be included in Earth system models that predict ecosystem C budgets under future N deposition/input scenarios.