Zhuwen Xu

Effects of Water and Nitrogen Addition on Species Turnover in Temperate Grasslands in Northern China

Global nitrogen (N) deposition and climate change have been identified as two of the most important causes of current plant diversity loss. However, temporal patterns of species turnover underlying diversity changes in response to changing precipitation regimes and atmospheric N deposition have received inadequate attention. We carried out a manipulation experiment in a steppe and an old-field in North China from 2005 to 2009, to test the hypothesis that water addition enhances plant species richness through increase in the rate of species gain and decrease in the rate of species loss, while N addition has opposite effects on species changes. Our results showed that water addition increased the rate of species gain in both the steppe and the old field but decreased the rates of species loss and turnover in the old field. In contrast, N addition increased the rates of species loss and turnover in the steppe but decreased the rate of species gain in the old field. The rate of species change was greater in the old field than in the steppe. Water interacted with N to affect species richness and species turnover, indicating that the impacts of N on semi-arid grasslands were largely mediated by water availability. The temporal stability of communities was negatively correlated with rates of species loss and turnover, suggesting that water addition might enhance, but N addition would reduce the compositional stability of grasslands. Experimental results support our initial hypothesis and demonstrate that water and N availabilities differed in the effects on rate of species change in the temperate grasslands, and these effects also depend on grassland types and/or land-use history. Species gain and loss together contribute to the dynamic change of species richness in semi-arid grasslands under future climate change.

Patterns of Plant Biomass Allocation in Temperate Grasslands across a 2500-km Transect in Northern China

Plant biomass allocation between below- and above-ground parts can actively adapt to the ambient growth conditions and is a key parameter for estimating terrestrial ecosystem carbon (C) stocks. To investigate how climatic variations affect patterns of plant biomass allocation, we sampled 548 plants belonging to four dominant genera (Stipa spp., Cleistogenes spp., Agropyron spp., and Leymus spp.) along a large-scale (2500 km) climatic gradient across the temperate grasslands from west to east in northern China. Our results showed that Leymus spp. had the lowest root/shoot ratios among the each genus. Root/shoot ratios of each genera were positively correlated with mean annual temperature (MAT), and negatively correlated with mean annual precipitation (MAP) across the transect. Temperature contributed more to the variation of root/shoot ratios than precipitation for Cleistogenes spp. (C4 plants), whereas precipitation exerted a stronger influence than temperature on their variations for the other three genera (C3 plants). From east to west, investment of C into the belowground parts increased as precipitation decreased while temperature increased. Such changes in biomass allocation patterns in response to climatic factors may alter the competition regimes among co-existing plants, resulting in changes in community composition, structure and ecosystem functions. Our results suggested that future climate change would have great impact on C allocation and storage, as well as C turnover in the grassland ecosystems in northern China.

Nitrogen and water addition reduce leaf longevity of steppe species

BACKGROUND AND AIMS Changes in supplies of resources will modify plant functional traits. However, few experimental studies have addressed the effects of nitrogen and water variations, either singly or in combination, on functional traits. METHODS A 2-year field experiment was conducted to test the effects of nitrogen and water addition on leaf longevity and other functional traits of the two dominant (Agropyron cristatum and Stipa krylovii) and three most common species (Cleistogenes squarrosa, Melilotoides ruthenica and Potentilla tanacetifolia) in a temperate steppe in northern China. KEY RESULTS Additional nitrogen and water increased leaf nitrogen content and net photosynthetic rate, and changed other measured functional traits. Leaf longevity decreased significantly with both nitrogen addition (-6 days in 2007 and -5·4 days in 2008; both P < 0·001) and watering (-13 days in 2007 and -9·9 days in 2008; both P < 0·001), and significant differences in leaf longevity were also found among species. Nitrogen and water interacted to affect leaf longevity and other functional traits. Soil water content explained approx. 70 % of the shifts in leaf longevity. Biomass at both species and community level increased under water and nitrogen addition because of the increase in leaf biomass production per individual plant. CONCLUSIONS The results suggest that additional nitrogen and water supplies reduce plant leaf longevity. Soil water availability might play a fundamental role in determining leaf longevity and other leaf functional traits, and its effects can be modified by soil nitrogen availability in semi-arid areas. The different responses of species to resource alterations may cause different global change ramifications under future climate change scenarios.

Abundance and community structure of ammonia-oxidizing Archaea and Bacteria in response to fertilization and mowing in a temperate steppe in Inner Mongolia

Based on a 6-year field trial in a temperate steppe in Inner Mongolia, we investigated the effects of nitrogen (N) and phosphorus (P) fertilization and mowing on the abundance and community compositions of ammonia-oxidizing Bacteria (AOB) and Archaea (AOA) upon early (May) and peak (August) plant growth using quantitative PCR (qPCR), terminal-restriction fragment length polymorphism (T-RFLP), cloning and sequencing. The results showed that N fertilization changed AOB community composition and increased AOB abundance in both May and August, but significantly decreased AOA abundance in May. By contrast, P fertilization significantly influenced AOB abundance only in August. Mowing significantly decreased AOA abundance and had little effect on AOA community compositions in May, while significantly influencing AOB abundance in both May and August, Moreover, AOA and AOB community structures showed obvious seasonal variations between May and August. Phylogenetic analysis showed that all AOA sequences fell into the Nitrososphaera cluster, and the AOB community was dominated by Nitrosospira Cluster 3. The results suggest that fertilization and mowing play important roles in affecting the abundance and community compositions of AOA and AOB.

Environmental changes drive the temporal stability of semi-arid natural grasslands through altering species asynchrony

Summary 1. Stability is an important property of ecological systems, many of which are experiencing increasing levels of anthropogenic environmental changes. However, how these environmental changes influence ecosystem stability remains poorly understood. 2. We conducted an 8-year field experiment in a semi-arid natural grassland to explore the effects of two common environmental changes, precipitation and nitrogen enrichment, on the temporal stability of plant above-ground biomass. A split-plot design, with precipitation as the main plot factor and nitrogen as the subplot factor, was used. Temporal stability was related to potential explanatory abiotic and biotic variables using regressions and structural equation modelling. 3. Increase in growing season precipitation enhanced plant species richness and promoted temporal stability of plant above-ground biomass. Nitrogen fertilization, however, reduced both plant species richness and temporal stability of plant above-ground biomass. Contrary to expectations, species richness was not an important driver of stability. Instead, community temporal stability was mainly driven by water and nitrogen availability that modulated the degree of species asynchrony and, to a lesser extent, by the stability of dominant plant species. 4. Synthesis. Our results highlight the importance of limiting resources for regulating community biomass stability and suggest that the projected increase in growing season precipitation may potentially offset negative effects of increased atmospheric nitrogen deposition on species diversity and community stability in semi-arid grasslands.

Plant nutrients do not covary with soil nutrients under changing climatic conditions

Nitrogen (N) and phosphorus (P) play vital roles in plant growth and development. Yet how climate regimes and soil fertility influence plant N and P stoichiometry is not well understood, especially in the belowground plant parts. Here we investigated plant aboveground and belowground N and P concentrations ([N] and [P]) and their stoichiometry in three dominant genera along a 2200 km long climatic gradient in northern China. Results showed that temperature explained more variation of [N] and [P] in C4 plants, whereas precipitation exerted a stronger influence on [N] and [P] in C3 plants. Both plant aboveground and belowground [N] and [P] increased with decreasing precipitation, and increasing temperatures yet were negatively correlated with soil [N] and [P]. Plant N:P ratios were unrelated with all climate and soil variables. Plant aboveground and belowground [N] followed an allometric scaling relationship, but the allocation of [P] was isometric. These results imply that internal processes stabilize plant N:P ratios and hence tissue N:P ratios may not be an effective parameter for predicting plant nutrient limitation. Our results also imply that past positive relationships between plant and nutrient stocks may be challenged under changing climatic conditions. While any modeling would need to be able to replicate currently observed relationships, it is conceivable that some relationships, such as those between temperature or rainfall and carbon:nutrient ratios, should be different under changing climatic conditions.

Effects of experimentally-enhanced precipitation and nitrogen on resistance, recovery and resilience of a semi-arid grassland after drought

Resistance, recovery and resilience are three important properties of ecological stability, but they have rarely been studied in semi-arid grasslands under global change. We analyzed data from a field experiment conducted in a native grassland in northern China to explore the effects of experimentally enhanced precipitation and N deposition on both absolute and relative measures of community resistance, recovery and resilience—calculated in terms of community cover—after a natural drought. For both absolute and relative measures, communities with precipitation enhancement showed higher resistance and lower recovery, but no change in resilience compared to communities with ambient precipitation in the semi-arid grassland. The manipulated increase in N deposition had little effect on these community stability metrics except for decreased community resistance. The response patterns of these stability metrics to alterations in precipitation and N are generally consistent at community, functional group and species levels. Contrary to our expectations, structural equation modeling revealed that water-driven community resistance and recovery result mainly from changes in community species asynchrony rather than species diversity in the semi-arid grassland. These findings suggest that changes in precipitation regimes may have significant impacts on the response of water-limited ecosystems to drought stress under global change scenarios.

Increased precipitation induces a positive plant-soil feedback in a semi-arid grassland

Background and AimsGiven that plant growth is often water limited in drylands, it has been proposed that water seems to influence productivity by altering physiological/metabolic responses and nutrient availability in short term. It is unclear, however, whether water mediates a positive plant-soil feedback and whether the feedback drives variations in plant productivity.MethodsA 4-year field experiment was performed to examine the effects of water and nitrogen (N) addition on nutrient concentrations in soil and plant, nutrient resorption and potential return, in a temperate grassland in northern China.ResultsWater addition enhanced plant N and phosphorus (P) concentrations but reduced plant N and P resorption efficiency, leading to the increased potential N and P return to soil via litterfall. Enhanced nutrient potential return likely contributed to an increase of plant productivity in the following year. These “fertilization effects” caused by water addition were similar to those by N addition.ConclusionsOur study suggests that the positive plant-soil feedback induced by increased precipitation may have a role in water-induced increases in productivity, and highlights the “fertilization effect” of water addition in a semiarid grassland in short term.

Responses of Soil Bacterial Communities to Nitrogen Deposition and Precipitation Increment Are Closely Linked with Aboveground Community Variation

It has been predicted that precipitation and atmospheric nitrogen (N) deposition will increase in northern China; yet, ecosystem responses to the interactive effects of water and N remain largely unknown. In particular, responses of belowground microbial community to projected global change and their potential linkages to aboveground macro-organisms are rarely studied. In this study, we examined the responses of soil bacterial diversity and community composition to increased precipitation and multi-level N deposition in a temperate steppe in Inner Mongolia, China, and explored the diversity linkages between aboveground and belowground communities. It was observed that N addition caused the significant decrease in bacterial alpha-diversity and dramatic changes in community composition. In addition, we documented strong correlations of alpha- and beta-diversity between plant and bacterial communities in response to N addition. It was found that N enriched the so-called copiotrophic bacteria, but reduced the oligotrophic groups, primarily by increasing the soil inorganic N content and carbon availability and decreasing soil pH. We still highlighted that increased precipitation tended to alleviate the effects of N on bacterial diversity and dampen the plant-microbe connections induced by N. The counteractive effects of N addition and increased precipitation imply that even though the ecosystem diversity and function are predicted to be negatively affected by N deposition in the coming decades; the combination with increased precipitation may partially offset this detrimental effect.

Soil exchangeable base cations along a chronosequence of Caragana microphylla plantation in a semi-arid sandy land, China

As a pioneer leguminous shrub species for vegetation re-establishment, Caragana microphylla is widely distributed in the semi-fixed and fixed sandy lands of the Horqin region, North China. C. microphylla plantations modify organic carbon (SOC), nitrogen (N) and phosphorus dynamics, bulk density and water-holding capacity, and biological activities in soils, but little is known with regard to soil exchange properties. Variation in soil exchangeable base cations was examined under C. microphylla plantations with an age sequence of 0, 5, 10, and 22 years in the Horqin Sandy Land, and at the depth of 0–10, 10–20, and 20–30 cm, respectively. C. microphylla has been planted on the non-vegetated sand dunes with similar physical-chemical soil properties. The results showed that exchangeable calcium (Ca), magnesium (Mg), and potassium (K), and cation exchange capacity (CEC) were significantly increased, and Ca saturation tended to decrease, while Mg and K saturations were increased with the plantation years. No difference was observed for exchangeable sodium (Na) neither with plantation years nor at soil depths. Of all the base cations and soil layers, exchangeable K at the depth of 0–10 cm accumulated most quickly, and it increased by 1.76, 3.16, and 4.25 times, respectively after C. microphylla was planted for 5, 10, and 22 years. Exchangeable Ca, Mg, and K, and CEC were significantly (P<0.001) and positively correlated with SOC, total N, pH, and electrical conductivity (EC). Soil pH and SOC are regarded as the main factors influencing the variation in exchangeable cations, and the preferential absorption of cations by plants and different leaching rates of base cations that modify cation saturations under C. microphylla plantation. It is concluded that as a nitrogen-fixation species, C. microphylla plantation is beneficial to increasing exchangeable base cations and CEC in soils, and therefore can improve soil fertility and create favorable microenvironments for plants and creatures in the semi-arid sandy land ecosystems.