Qingmin Pan

PRIMARY PRODUCTION AND RAIN USE EFFICIENCY ACROSS A PRECIPITATION GRADIENT ON THE MONGOLIA PLATEAU

Understanding how the aboveground net primary production (ANPP) of arid and semiarid ecosystems of the world responds to variations in precipitation is crucial for assessing the impacts of climate change on terrestrial ecosystems. Rain-use efficiency (RUE) is an important measure for acquiring this understanding. However, little is known about the response pattern of RUE for the largest contiguous natural grassland region of the world, the Eurasian Steppe. Here we investigated the spatial and temporal patterns of ANPP and RUE and their key driving factors based on a long-term data set from 21 natural arid and semiarid ecosystem sites across the Inner Mongolia steppe region in northern China. Our results showed that, with increasing mean annual precipitation (MAP), (1) ANPP increased while the interannual variability of ANPP declined, (2) plant species richness increased and the relative abundance of key functional groups shifted predictably, and (3) RUE increased in space across different ecosystems but decreased with increasing annual precipitation within a given ecosystem. These results clearly indicate that the patterns of both ANPP and RUE are scale dependent, and the seemingly conflicting patterns of RUE in space vs. time suggest distinctive underlying mechanisms, involving interactions among precipitation, soil N, and biotic factors. Also, while our results supported the existence of a common maximum RUE, they also indicated that its value could be substantially increased by altering resource availability, such as adding nitrogen. Our findings have important implications for understanding and predicting ecological impacts of global climate change and for management practices in arid and semiarid ecosystems in the Inner Mongolia steppe region and beyond.

Grazing intensity impacts soil carbon and nitrogen storage of continental steppe

Recent studies have underscored the importance of grasslands as potential carbon (C) sinks. We performed a grazing experiment with seven stocking rates (SR0, SR1.5, SR3.0, SR4.5, SR6.0, SR7.5, and SR9.0 for 0, 1.5, 3.0, 4.5, 6.0, 7.5, and 9.0 sheep ha−1, respectively) to investigate the effect of increasing grazing pressure on soil C and nitrogen (N) storage in the temperate grasslands of northern China. The results revealed that C and N storage in both 0–10 cm and 10–30 cm soil layers decreased linearly with increasing stocking rates. Carbon storage in the 0–10 cm soil layer was significantly higher in lightly grazed grasslands than in heavily grazed grasslands after a 5-yr grazing treatment. Our findings suggest an underlying transformation from soil C sequestration under light grazing to C loss under heavy grazing, and that the threshold for this transformation is 4.5 sheep ha−1 (grazing period from June to September). Results confirmed that grasslands used for grazing in northern China have the capacity to sequester C in the soil under appropriate grazing pressure, but that they lose C under heavy grazing. Therefore, appropriate grazer densities will promote soil C sequestration in the grasslands of northern China.

Hierarchical Plant Responses and Diversity Loss after Nitrogen Addition: Testing Three Functionally-Based Hypotheses in the Inner Mongolia Grassland

Background Numerous studies have shown that nitrogen (N) deposition decreases biodiversity in terrestrial ecosystems. To explain the N-induced species loss, three functionally based hypotheses have been proposed: the aboveground competition hypothesis, the belowground competition hypothesis, and the total competition hypothesis. However, none of them is supported sufficiently by field experiments. A main challenge to testing these hypotheses is to ascertain the role of shoot and root competition in controlling plant responses to N enrichment. Simultaneously examining both aboveground and belowground responses in natural ecosystems is logistically complex, and has rarely been done. Methodology/Principal Findings In a two-year N addition experiment conducted in a natural grassland ecosystem, we investigated both above- and belowground responses of plants at the individual, species, and community levels. Plants differed significantly in their responses to N addition across the different organizational levels. The community-level species loss was mainly due to the loss of perennial grasses and forbs, while the relative abundance of plant species was dependent mainly on individual-level responses. Plasticity in biomass allocation was much smaller within a species than between species, providing a biological basis for explaining the functionally based species loss. All species increased biomass allocation to aboveground parts, but species with high belowground allocations were replaced by those with high aboveground allocations, indicating that the increased aboveground competition was the key process responsible for the observed diversity loss after N addition in this grassland ecosystem. Conclusions/Significance Our findings shed new light on the validity of the three competing hypotheses concerning species loss in response to N enrichment. They also have important implications for predicting the future impacts of N deposition on the structure and functioning of terrestrial ecosystems. In addition, we have developed a new technique for ascertaining the roles of aboveground and belowground competition in determining plant responses to N fertilization.

Plant responses following grazing removal at different stocking rates in an Inner Mongolia grassland ecosystem

Grazing removal is widely used in grassland management. Plant responses following grazing removal at different organizational levels, however, are not well understood. We examined plant responses at different stocking rates in an Inner Mongolia grassland ecosystem dominated by Leymus chinensis and Stipa grandis. Our results indicated that plant response patterns differed significantly among stocking rates, at different levels of organization, and between wet and dry years. Community aboveground net primary production (ANPP) recovered more quickly at low and moderate stocking rates than those at high stocking rates. Response of aboveground net primary production (RANPP) was significantly positively correlated with both individual biomass and density responses of L. chinensis. Overcompensation of L. chinensis after grazing removal contributed greatly to positive RANPP at the community level. Significant compensatory effects were found between the two dominant species and between dominant species and the remaining non-dominant species. Variation in precipitation significantly affected community ANPP, relationships between community and species responses, and compensatory effects between species. Our study suggests that periodic grazing removal is likely to be a useful method for grassland management and that a combination of species with compensatory effects can be advantageous for reseeding practices in grassland restoration.

Nonlinear responses of ecosystem carbon fluxes and water-use efficiency to nitrogen addition in Inner Mongolia grassland

Summary Nitrogen (N) deposition is a continuous process and likely to affect ecosystem carbon (C) and water fluxes in a nonlinear way. However, experimental evidence is still lacking because most previous studies on these impacts usually used two discrete levels of N treatment. By a 12-year, 6-level N addition experiment in Inner Mongolia grassland, we found that the responses of C fluxes, including net ecosystem carbon exchange (NEE), gross ecosystem productivity (GEP), ecosystem respiration (ER), all exhibited nonlinear patterns with increasing N addition rate while that of evapotranspiration did not significantly change. As a result, the response of ecosystem water-use efficiency (EWUE) followed a similar pattern with NEE. These N-induced changes in C fluxes were greatly affected by the distribution of precipitation among different stages of growing seasons and mainly driven by the alterations in biomass production rather than soil temperature and soil moisture. This study highlights the importance of the nonlinearity of N addition impacts on ecosystem C fluxes, which should be incorporated into the global-C-cycling models for better predicting future C balance. Our results also have implications for the use of fertilization in restoring the degraded grasslands given that N addition can promote biomass production and ecosystem C uptake without additional water evapotranspiration but also change ecosystem composition.

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.

Hierarchical Reproductive Allocation and Allometry within a Perennial Bunchgrass after 11 Years of Nutrient Addition

Bunchgrasses are one of the most important plant functional groups in grassland ecosystems. Reproductive allocation (RA) for a bunchgrass is a hierarchical process; however, how bunchgrasses adjust their RAs along hierarchical levels in response to nutrient addition has never been addressed. Here, utilizing an 11-year nutrient addition experiment, we examined the patterns and variations in RA of Agropyron cristatum at the individual, tiller and spike levels. We evaluated the reproductive allometric relationship at each level by type II regression analysis to determine size-dependent and size-independent effects on plant RA variations. Our results indicate that the proportion of reproductive individuals in A. cristatum increased significantly after 11 years of nutrient addition. Adjustments in RA in A. cristatum were mainly occurred at the individual and tiller levels but not at the spike level. A size-dependent effect was a dominant mechanism underlying the changes in plant RA at both individual and tiller levels. Likewise, the distribution of plant size was markedly changed with large individuals increasing after nutrient addition. Tiller-level RA may be a limiting factor for the adjustment of RA in A. cristatum. To the best of our knowledge, this study is the first to examine plant responses in terms of reproductive allocation and allometry to nutrient enrichment within a bunchgrass population from a hierarchical view. Our findings have important implications for understanding the mechanisms underlying bunchgrass responses in RA to future eutrophication due to human activities. In addition, we developed a hierarchical analysis method for disentangling the mechanisms that lead to variation in RA for perennial bunchgrasses.

Effects of functional diversity loss on ecosystem functions are influenced by compensation

Understanding the impacts of biodiversity loss on ecosystem functioning and services has been a central issue in ecology. Experiments in synthetic communities suggest that biodiversity loss may erode a set of ecosystem functions, but studies in natural communities indicate that the effects of biodiversity loss are usually weak and that multiple functions can be sustained by relatively few species. Yet, the mechanisms by which natural ecosystems are able to maintain multiple functions in the face of diversity loss remain poorly understood. With a long-term and large-scale removal experiment in the Inner Mongolian grassland, here we showed that losses of plant functional groups (PFGs) can reduce multiple ecosystem functions, including biomass production, soil NO3 -N use, net ecosystem carbon exchange, gross ecosystem productivity, and ecosystem respiration, but the magnitudes of these effects depended largely on which PFGs were removed. Removing the two dominant PFGs (perennial rhizomatous grasses and perennial bunchgrasses) simultaneously resulted in dramatic declines in all examined functions, but such declines were circumvented when either dominant PFG was present. We identify the major mechanism for this as a compensation effect by which each dominant PFG can mitigate the losses of others. This study provides evidence that compensation ensuing from PFG losses can mitigate their negative consequence, and thus natural communities may be more resilient to biodiversity loss than currently thought if the remaining PFGs have strong compensation capabilities. On the other hand, ecosystems without well-developed compensatory functional diversity may be much more vulnerable to biodiversity loss.

Ammonia emissions from soil under sheep grazing in Inner Mongolian grasslands of China

Ammonia (NH3) emission and redeposition play a major role in terrestrial nitrogen (N) cycles and can also cause environmental problems, such as changes in biodiversity, soil acidity, and eutrophication. Previous field grazing experiments showed inconsistent (positive, neutral, and negative) NH3 volatilization from soils in response to varying grazing intensities. However, it remains unclear whether, or to what extent, NH3 emissions from soil are affected by increasing grazing intensities in Inner Mongolian grasslands. Using a 5-year grazing experiment, we investigated the relationship between NH3 volatilization from soil and grazing pressure (0.0, 3.0, 6.0, and 9.0 sheep/hm2) from June to September of 2009 and 2010 via the vented-chamber method. The results show that soil NH3 volatilization was not significantly different at different grazing intensities in 2009, although it was higher at the highest stocking rate during 2010. There was no significant linear relationship between soil NH3 volatilization rates and soil NH4+-N, but soil NH3 volatilization rates were significantly related to soil water content and air temperature. Grazing intensities had no significant influence on soil NH3 volatilization. Soil NH3 emissions from June to September (grazing period), averaged over all grazing intensities, were 9.6±0.2 and 19.0±0.2 kg N/hm2 in 2009 and 2010, respectively. Moreover, linear equations describing monthly air temperature and precipitation showed a good fit to changes in soil NH3 emissions (r=0.506, P=0.014). Overall, grazing intensities had less influence than that of climatic factors on soil NH3 emissions. Our findings provide new insights into the effects of grazing on NH3 volatilization from soil in Inner Mongolian grasslands, and have important implications for understanding N cycles in grassland ecosystems and for estimating soil NH3 emissions on a regional scale.

Scale dependence of the diversity–stability relationship in a temperate grassland

A positive relationship between biodiversity and ecosystem stability has been reported in many ecosystems; however, it has yet to be determined whether and how spatial scale affects this relationship. Here, for the first time, we assessed the effects of alpha, beta and gamma diversity on ecosystem stability and the scale dependence of the slope of the diversity-stability relationship.By employing a long-term (33 years) dataset from a temperate grassland, northern China, we calculated the all possible spatial scales with the complete combination from the basic 1-m2 plots.Species richness was positively associated with ecosystem stability through species asynchrony and overyielding at all spatial scales (1, 2, 3, 4 and 5 m2). Both alpha and beta diversity were positively associated with gamma stability.Moreover, the slope of the diversity-area relationship was significantly higher than that of the stability-area relationship, resulting in a decline of the slope of the diversity-stability relationship with increasing area.Synthesis. With the positive species diversity effect on ecosystem stability from small to large spatial scales, our findings demonstrate the need to maintain a high biodiversity and biotic heterogeneity as insurance against the risks incurred by ecosystems in the face of global environmental changes.