Precipitation-induced biomass enhancement and differential allocation in Inner Mongolia's herbaceous and shrub communities.

Changes in precipitation patterns induced by global climate change have profound implications for the structure and function of grassland ecosystems. However, the relationship between plant diversity and ecosystem function across different grassland types, particularly those with varying plant compositions and dominant species, remains inadequately understood. To address this knowledge gap, a five-year experimental manipulation of precipitation was conducted within herbaceous and shrub communities in the desert grasslands of Inner Mongolia. We found that increased precipitation significantly enhances aboveground biomass (AGB), belowground biomass (BGB), and community total biomass (CTB) in both herbaceous and shrub communities. In herbaceous communities, increased precipitation led to a disproportionate increase in both aboveground and belowground biomass, supporting the optimal allocation hypothesis. Structural equation modeling (SEM) further elucidated that precipitation regulates AGB and CTB through species richness and functional traits in herbaceous communities. In shrub communities, precipitation influences AGB, BGB, and CTB by affecting species richness and soil water content. This study highlights the critical role of precipitation in shaping biomass dynamics and allocation strategies within herbaceous and shrub communities in desert steppe of Inner Mongolia. These findings provide essential insights into the potential responses of desert grassland ecosystems to ongoing climate change.

Patterns and driving factors of functional traits of desert species with different elevational distributions in the Tibetan Plateau and adjacent areas.

Variations in functional traits serve as measures of plants' ability to adapt to environment. Exploring the patterns of functional traits of desert plants along elevational gradients is helpful to understand the responses and adaptation strategies of species to changing environments. However, it is unknown whether the relationship between functional traits and elevation is affected by differences in the species' elevational distributions (elevation preference and species' range). Importantly, most researches have concerned with differences in mean trait values and ignored intraspecific trait variation. Here, we measured functional traits of desert plants along a wide elevational gradient in the Tibetan Plateau and adjacent areas and explored functional trait patterns over elevation in species with different elevational distributions. We decomposed trait variation and further investigated characterizations of intraspecific variation. Ultimately, the main drivers of trait variation were identified using redundancy analysis. We found that species' elevational distributions significantly influenced the relationship of functional traits such as plant height, leaf dry matter content, leaf thickness, leaf nitrogen and carbon content with elevation. Species with a lower elevational preference showed greater trait variation than species with a higher elevational preference, suggesting that species that prefer high elevation are more conservative facing environmental changes. We provide evidence that interspecific trait variation in leaf thickness and leaf carbon content decreased with increasing species' range, indicating that increased variations in resistance traits within species make greater responsiveness to environmental changes, enabling species a wider range. Elevation, temperature and precipitation were the main drivers of trait variation in species with a low elevational preference, while the effect of precipitation on trait variation in species with a high elevational preference was not significant. This study sheds new insights on how plants with different elevational distributions regulate their ecological strategies to cope with changing environments.

The effect of nitrogen input on N2O emission depends on precipitation in a temperate desert steppe.

Nitrous oxide (N2O) is the third most important greenhouse gas, and can damage the atmospheric ozone layer, with associated threats to terrestrial ecosystems. However, to date it is unclear how extreme precipitation and nitrogen (N) input will affect N2O emissions in temperate desert steppe ecosystems. Therefore, we conducted an in-situ in a temperate desert steppe in the northwest of Inner Mongolia, China between 2018 and 2021, in which N inputs were combined with natural extreme precipitation events, with the aim of better understanding the mechanism of any interactive effects on N2O emission. The study result showed that N2O emission in this desert steppe was relatively small and did not show significant seasonal change. The annual N2O emission increased in a non-linear trend with increasing N input, with a much greater effect of N input in a wet year (2019) than in a dry year (2021). This was mainly due to the fact that the boost effect of high N input (on June 17th 2019) on N2O emission was greatly amplified by nearly 17-46 times by an extreme precipitation event on June 24th 2019. In contrast, this greatly promoting effect of high N input on N2O emission was not observed on September 26th 2019 by a similar extreme precipitation event. Further analysis showed that soil NH4+-N content and the abundance of ammonia oxidizing bacteria (amoA (AOB)) were the most critical factors affecting N2O emission. Soil moisture played an important indirect role in regulating N2O emission, mainly by influencing the abundance of amoA (AOB) and de-nitrification functional microorganisms (nosZ gene). In conclusion, the effect of extreme precipitation events on N2O emission was greatly increased by high N input. Furthermore, in this desert steppe, annual N2O flux is co-managed through soil nitrification substrate concentration (NH4+-N), the abundance of soil N transformation functional microorganisms and soil moisture. Overall, it was worth noting that an increase in extreme precipitation coupled with increasing N input may significantly increase future N2O emissions from desert steppes.

Plant functional traits mediate the response magnitude of plant-litter-soil microbial C: N: P stoichiometry to nitrogen addition in a desert steppe.

Global nitrogen deposition is significantly altering the carbon (C), nitrogen (N) and phosphorus (P) stoichiometry in terrestrial ecosystems, yet how N deposition simultaneously affects plant-litter-soil-soil microbial stoichiometry in arid grassland is still unclear. In a five-year experimental study conducted in a desert steppe in Northern China, we investigated the effects of N addition on the C:N:P stoichiometry of plants, litter, soil, and soil microbes. We also used structural equation modelling (SEM) exploring the direct or indirect effects of N addition, plant species diversity, functional traits and diversity, soil microbial diversity, soil pH, soil electrical conductivity (EC) and moisture on the stoichiometry in plant-soil system. The results showed that N addition increased the N, P concentrations and N:P in plants, the N concentration and N:P in litter, and the C, N concentrations, C:P and N:P in microbes. Conversely, it decreased the C:N and C:P in plants, and litter C:N. Functional traits, functional dispersion (FDis), soil pH and EC accounted for a substantial proportion of the observed variations in elemental concentrations (from 42 % to 69 %) and stoichiometry (from 9 % to 73 %) across different components. SEM results showed that N addition decreased C:N and C:P in plants and litter by increasing FDis and leaf N content, while increased plant and litter N:P by decreasing leaf C content and increasing specific leaf area, respectively. Furthermore, N addition increased microbial C:P by increasing leaf thickness. We also found the mediating effects of soil pH and EC on C:N, C:P of litter and microbial N:P. Overall, our research suggests that plant functional traits as key predictors of nutrient cycling responses in desert steppes under N addition. This study extends the application of plant functional traits, enhances our understanding of C and nutrient cycling and facilitates predicting the response of desert steppes to N deposition.

Growing-season drought and nitrogen addition interactively impair grassland ecosystem stability by reducing species diversity, asynchrony, and stability.

Aboveground net primary productivity controls the amount of energy available to sustain all living organisms, and its sustainable provision relies on the stability of grassland ecosystems. Human activities leading to global changes, such as increased nitrogen (N) deposition and the more frequent occurrence of extreme precipitation events, with N addition increasing the sensitivity of ecosystem production stability to changes in the precipitation regime. However, whether N addition, in combination with seasonal precipitation increases or severe drought, affects ecosystem stability remains unclear. In this study, we conducted a six-year environmental change monitoring experiment in a semiarid grassland in northern China to test the effects of N addition, seasonal drought, and precipitation increases on the temporal stability of ecosystem productivity. Our study revealed that an interaction between drought and N addition reduced species diversity, species asynchrony, species stability, and thus ecosystem stability. These environmental change drivers (except for precipitation increase) induced a positive relationship between species asynchrony and diversity, whereas N addition interactively with drought and precipitation increase led to a negative relationship between diversity and species stability. Only N addition interactively with drought induced a positive species diversity-ecosystem stability relationship because lower species stability was overcome by increased species asynchrony. Our study is great importance to illustrate that production temporal stability tends to be inhibited with drought, though interactively with nutrient N addition. These findings highlight the primary role of asynchronous dynamics among species in modulating the effects of environmental change on diversity-stability relationships.

Grassland sensitivity to drought is related to functional composition across East Asia and North America.

Plant traits can be helpful for understanding grassland ecosystem responses to climate extremes, such as severe drought. However, intercontinental comparisons of how drought affects plant functional traits and ecosystem functioning are rare. The Extreme Drought in Grasslands experiment (EDGE) was established across the major grassland types in East Asia and North America (six sites on each continent) to measure variability in grassland ecosystem sensitivity to extreme, prolonged drought. At all sites, we quantified community-weighted mean functional composition and functional diversity of two leaf economic traits, specific leaf area and leaf nitrogen content, in response to drought. We found that experimental drought significantly increased community-weighted means of specific leaf area and leaf nitrogen content at all North American sites and at the wetter East Asian sites, but drought decreased community-weighted means of these traits at moderate to dry East Asian sites. Drought significantly decreased functional richness but increased functional evenness and dispersion at most East Asian and North American sites. Ecosystem drought sensitivity (percentage reduction in aboveground net primary productivity) positively correlated with community-weighted means of specific leaf area and leaf nitrogen content and negatively correlated with functional diversity (i.e., richness) on an intercontinental scale, but results differed within regions. These findings highlight both broad generalities but also unique responses to drought of community-weighted trait means as well as their functional diversity across grassland ecosystems.

Effects of Grazing, Extreme Drought, Extreme Rainfall and Nitrogen Addition on Vegetation Characteristics and Productivity of Semiarid Grassland.

Grassland use patterns, water and nutrients are the main determinants of ecosystem structure and function in semiarid grasslands. However, few studies have reported how the interactive effects of rainfall changes and nitrogen deposition influence the recovery of semiarid grasslands degraded by grazing. In this study, a simulated grazing, increasing and decreasing rainfall, nitrogen deposition test platform was constructed, and the regulation mechanism of vegetation characteristics and productivity were studied. We found that grazing decreased plant community height (CWMheight) and litter and increased plant density. Increasing rainfall by 60% from May to August (+60%) increased CWMheight; decreasing rainfall by 60% from May to August (-60%) and by 100% from May to June (-60 d) decreased CWMheight and coverage; -60 d, +60% and increasing rainfall by 100% from May to June (+60 d) increased plant density; -60% increased the Simpson dominance index (D index) but decreased the Shannon-Wiener diversity index (H index); -60 d decreased the aboveground biomass (ABG), and -60% increased the underground biomass (BGB) in the 10-60 cm layer. Nitrogen addition decreased species richness and the D index and increased the H index and AGB. Rainfall and soil nitrogen directly affect AGB; grazing and rainfall can also indirectly affect AGB by inducing changes in CWMheight; grazing indirectly affects BGB by affecting plant density and soil nitrogen. The results of this study showed that in the semiarid grassland of Inner Mongolia, grazing in the nongrowing season and grazing prohibition in the growing season can promote grassland recovery, continuous drought in the early growing season will have dramatic impacts on productivity, nitrogen addition has a certain impact on the species composition of vegetation, and the impact on productivity will not appear in the short term.

Extreme drought does not alter the stability of aboveground net primary productivity but decreases the stability of belowground net primary productivity in a desert steppe of northern China.

Extreme droughts strongly impact grassland ecology, both functionally and structurally. However, a comprehensive understanding of the drought impacts on the ecosystem stability is critical for its sustainable development under changing climate. We experimentally report the impact of extreme drought on the temporal stability of aboveground net primary productivity (ANPP) and belowground net primary productivity (BNPP) in a desert steppe of northern China. The relative importance evaluation of extreme drought, soil properties, species asynchrony, taxonomic, functional, and phylogenetic diversity was performed using structural equation modeling (SEM) to measure the temporal ANPP and BNPP stabilities. Our findings suggested that extreme drought decreased BNPP stability but did not affect ANPP stability. Extreme drought reduced taxonomic and phylogenetic diversity, ANPP, and soil water content but did not affect species asynchrony, functional diversity, or BNPP. Species richness, Shannon-Wiener index, and soil water content were positively correlated with BNPP stability. The SEM results showed a drought-mediated indirect weakening of BNPP stability via modification of species richness. Asynchrony of species unrelated to drought, however, directly affected ANPP stability. The mechanisms underlying the response determination of ANPP and BNPP stability to extreme drought in desert steppe varied notably. Depending on the species asynchrony, ANPP reduced by extreme drought could maintain higher stability. However, extreme drought lowered BNPP stability by altering species richness.

Effects of multi-resource addition on grassland plant productivity and biodiversity along a resource gradient.

The change of plant biodiversity caused by resource-enhancing global changes has greatly affected grassland productivity. However, it remains unclear how multi-resource enrichment induces the effects of multifaceted biodiversity on grassland productivity under different site resource constraints. We conducted a multiple resource addition (MRA) experiment of water and nutrients at three sites located along a resource gradient in northern China. This allowed us to assess the response of aboveground net primary productivity (ANPP), species (species richness and plant density), functional (functional richness and community-weighted mean of traits) and phylogenetic (phylogenetic richness) diversity to increasing number of MRA. We used structural equation model (SEM) to examine the direct and indirect effects of MRA and multifaceted biodiversity on ANPP. The combined addition of the four resources increased ANPP at all three sites. But with increasing number of MRA, biodiversity varied at the three sites. At the high resource constraint site, species richness, plant density and leaf nitrogen concentration (LNC) increased. At the medium resource constraint site, plant height and LNC increased, leaf dry matter content (LDMC) decreased. At the low resource constraint site, species, functional and phylogenetic richness decreased, and height increased. The SEM showed that MRA increased ANPP directly at all three sites, and indirectly by increasing plant density at the high constraint site and height at the medium constraint site. Independent of MRA, ANPP was affected by height at the high resource constraint site and LNC at the low resource constraint site. Our results illustrate that multi-resource addition positively affects productivity, while affects biodiversity depending on site resource constraint. The study highlights that site resource constraint conditions need to be taken into consideration to better predict grassland structure and function, particularly under the future multifaceted global change scenarios.

Contrasting relationships between plant-soil microbial diversity are driven by geographic and experimental precipitation changes.

Soil microbe diversity plays a key role in dryland ecosystem function under global climate change, yet little is known about how plant-soil microbe relationships respond to climate change. Altered precipitation patterns strongly shape plant community composition in deserts and steppes, but little research has demonstrated whether plant biodiversity attributes mediate the response of soil microbial diversity to long- and short-term precipitation changes. Here we used a comparative study to explore how altered precipitation along the natural and experimental gradients affected associations of soil bacterial and fungal diversity with plant biodiversity attributes (species, functional and phylogenetic diversity) and soil properties in desert-shrub and steppe-grass communities. We found that along both gradients, increasing precipitation increased soil bacterial and fungal richness in the desert and soil fungal richness in the steppe. Soil bacterial richness in the steppe was also increased by increasing precipitation in the experiment but was decreased along the natural gradient. Plant biodiversity and soil properties explained the variations in soil bacterial and fungal richness from 43 % to 96 % along the natural gradient and from 19 to 46 % in the experiment. Overall, precipitation effects on soil bacterial or fungal richness were mediated by plant biodiversity attributes (species richness and plant height) or soil properties (soil water content) along the natural gradient but were mediated by plant biodiversity attributes (functional or phylogenetic diversity) in the experiment. These results suggest that different mechanisms are responsible for the responses of soil bacterial and fungal diversity to long- and short-term precipitation changes. Long- and short-term precipitation changes may modify plant biodiversity attribute effects on soil microbial diversity in deserts and steppes, highlighting the importance of precipitation changes in shaping relationships between plant and soil microbial diversity in water-limited areas.