Rhizome Severing Increases Root Lifespan of Leymus chinensis in a Typical Steppe of Inner Mongolia

Background

Root lifespan is an important trait that determines plants'' ability to acquire and conserve soil resources. There have been several studies investigating characteristics of root lifespan of both woody and herbaceous species. However, most of the studies have focused on non-clonal plants, and there have been little data on root lifespan for clonal plants that occur widely in temperate grasslands.

Methodology/Principal Findings

We investigated the effects of rhizome severing on overall root lifespan of Leymus chinensis, a clonal, dominant grass species in the temperate steppe in northern China, in a 2-year field study using modified rhizotron technique. More specifically, we investigated the effects of rhizome severing on root lifespan of roots born in different seasons and distributed at different soil depths. Rhizome severing led to an increase in the overall root lifespan from 81 to 103 days. The increase in root lifespan exhibited spatial and temporal characteristics such that it increased lifespan for roots distributed in the top two soil layers and for roots born in summer and spring, but it had no effect on lifespan of roots in the deep soil layer and born in autumn. We also examined the effect of rhizome severing on carbohydrate and N contents in roots, and found that root carbohydrate and N contents were not affected by rhizome severing. Further, we found that root lifespan of Stipa krylovii and Artemisia frigida, two dominant, non-clonal species in the temperate steppe, was significantly longer (118 d) than that of L. chinensis (81 d), and this value became comparable to that of L. chinensis under rhizome severing (103 d).

Conclusions/Significance

We found that root lifespan in dominant, clonal L. chinensis was shorter than for the dominant, non-clonal species of S. krylovii and A. frigida. There was a substantial increase in the root lifespan of L. chinensis in response to severing their rhizomes, and this increase in root lifespan exhibited temporal and spatial characteristics. These findings suggest that the presence of rhizomes is likely to account for the observed short lifespan of clonal plant species in the temperate steppe.     

<Emphasis Type="Italic">Artemisia frigida</Emphasis> and <Emphasis Type="Italic">Stipa krylovii</Emphasis>, two dominant species in Inner Mongolia steppe,differed in their responses to elevated atmospheric CO<Subscript>2</Subscript> concentration

Aims

Despite extensive studies on effects of elevated CO₂ concentration ([CO₂]_e) on plant growth, few studies have investigated the responses of native grassland plant species to [CO₂]_e in terms of nutrient acquisition.

Methods

The effects of [CO₂]_e (769 ± 23 ppm) on Artemisia frigida and Stipa krylovii, two dominant species in Inner Mongolia steppe were investigated by growing them for 7 weeks in Open-Top Chambers (OTC).

Results

Exposure to [CO₂]_e enhanced shoot and root growth of A. frigida and S. krylovii. Elevated [CO₂] increased photosynthetic rates (Pn) by 34 % in A. frigida but decreased Pn by 52 % in S. krylovii. Moreover, root-secreted acid phosphatase activity in A. frigida was stimulated by [CO₂]_e, while exudation of malate from roots of S. krylovii was suppressed by [CO₂]_e. Exposure to [CO₂]_e led to a decrease in P concentration in shoots and roots of A. frigida and S. krylovii, but total amount of P accumulated in shoots and roots of both species was increased by [CO₂]_e.

Conclusions

The two dominant species in temperate steppes differed in their responses to [CO₂]_e, such that A. frigida was more adapted to [CO₂]_e than S. krylovii under low availability of soil P.

     

Responses of litter decomposition and nutrient release rate to water and nitrogen addition differed among three plant species dominated in a semi-arid grassland

Background and aims

Precipitation and nitrogen (N) deposition are predicted to increase in northern China. The present paper aimed to better understand how different dominant species in semi-arid grasslands in this region vary in their litter decomposition and nutrient release responses to increases in precipitation and N deposition.

Methods

Above-ground litter of three dominant species (two grasses, Agropyron cristatum and Stipa krylovii, and one forb, Artemisia frigida) was collected from areas without experimental treatments in a semi-arid grassland in Inner Mongolia. Litter decomposition was studied over three years to determine the effects of water and N addition on litter decomposition rate and nutrient dynamics.

Results

Litter mass loss and nutrient release were faster for the forb species than for the two grasses during decomposition. Both water and N addition increased litter mass loss of the grass A. cristatum, while the treatments showed no impacts on that of the forb A. frigida. Supplemental N had time-dependent, positive effects on litter mass loss of the grass S. krylovii. During the three-year decomposition study, the release of N from litter was inhibited by N addition for the three species, and it was promoted by water addition for the two grasses. Across all treatments, N and potassium (K) were released from the litter of all three species, whereas calcium (Ca) was accumulated. Phosphorus (P) and magnesium (Mg) were released from the forb litter but accumulated in the grass litter after three years of decomposition.

Conclusions

Our findings revealed that the litter decomposition response to water and N supplementation differed among dominant plant species in a semi-arid grassland, indicating that changes in dominant plant species induced by projected increases in precipitation and N deposition are likely to affect litter decomposition, nutrient cycling, and further biogeochemical cycles in this grassland. The asynchronous nutrient release of different species’ litter found in the present study highlights the complexity of nutrient replenishment from litter decomposition in the temperate steppe under scenarios of enhancing precipitation and N deposition.

     

Allelopathic potential of Artemisia frigida and successional changes of plant communities in the northern China steppe

In the northern China steppe, overgrazing has decreased the abundance of many species that were originally dominant, but increased the abundance of Artemisia frigida. We aimed to determine whether the adaptive and competitive abilities of A. frigida are associated with allelopathy. Soil nutrient characteristics could not explain the poor growth of the originally dominant species. Volatile compounds released from A. frigida leaves and aqueous extracts (0.025, 0.05, 0.075, 0.10, and 0.15 g ml^?1) from A. frigida leaves and roots and from soil under A. frigida inhibited seed germination and seedling growth of three dominant species (Leymus chinensis, Stipa krylovii, and Cleistogenes squarrosa). Allelopathic activity varied according to extract concentration, test species, and extract source. Germination was most strongly inhibited in S. krylovii, followed by L. chinensis and then C. squarrosa. Seedling growth was most strongly inhibited in L. chinensis, followed by S. krylovii and then C. squarrosa. Gas chromatography–mass spectrometry analyses of the leaf volatiles identified 27 compounds, primarily monoterpene or sesquiterpene compounds and their oxygen-containing derivatives, such as eucalyptol, beta.-myrcene, 1,6-octadien-3-ol,3,7-dimethyl, 3-carene, bicyclo[2.2.1]heptan-2-one,1,7,7-trimethyl-,(1R), cis-sabinenehydrate, camphene, and alpha-Pinene. These findings suggest that allelochemicals from A. frigida can modify the surrounding micro-habitat. The responses of target plants to allelopathy of A. frigida may be one reason for changes in plant community succession in the northern China steppe.     

Linking ethylene to nitrogen-dependent leaf longevity of grass species in a temperate steppe

Background and Aims

Leaf longevity is an important plant functional trait that often varies with soil nitrogen supply. Ethylene is a classical plant hormone involved in the control of senescence and abscission, but its role in nitrogen-dependent leaf longevity is largely unknown.

Methods

Pot and field experiments were performed to examine the effects of nitrogen addition on leaf longevity and ethylene production in two dominant plant species, Agropyron cristatum and Stipa krylovii, in a temperate steppe in northern China.

Key Results

Nitrogen addition increased leaf ethylene production and nitrogen concentration but shortened leaf longevity; the addition of cobalt chloride, an ethylene biosynthesis inhibitor, reduced leaf nitrogen concentration and increased leaf longevity. Path analysis indicated that nitrogen addition reduced leaf longevity mainly through altering leaf ethylene production.

Conclusions

These findings provide the first experimental evidence in support of the involvement of ethylene in nitrogen-induced decrease in leaf longevity.     

Effects of mowing on methane uptake in a semiarid grassland in northern China

Background

Mowing is a widely adopted management practice for the semiarid steppe in China and affects CH₄ exchange. However, the magnitude and the underlying mechanisms for CH₄ uptake in response to mowing remain uncertain.

Methodology/Principal Findings

In two consecutive growing seasons, we measured the effect of mowing on CH₄ uptake in a steppe community. Vegetation was mowed to 2 cm (M2), 5 cm (M5), 10 cm (M10), 15 cm (M15) above soil surface, respectively, and control was set as non-mowing (NM). Compared with control, CH₄ uptake was substantially enhanced at almost all the mowing treatments except for M15 plots of 2009. CH₄ uptake was significantly correlated with soil microbial biomass carbon, microbial biomass nitrogen, and soil moisture. Mowing affects CH₄ uptake primarily through its effect on some biotic factors, such as net primary productivity, soil microbial C\N supply and soil microbial activities, while soil temperature and moisture were less important.

Conclusions/Significance

This study found that mowing affects the fluxes of CH₄ in the semiarid temperate steppe of north China.     

Water- and Plant-Mediated Responses of Ecosystem Carbon Fluxes to Warming and Nitrogen Addition on the Songnen Grassland in Northeast China

Background

Understanding how grasslands are affected by a long-term increase in temperature is crucial to predict the future impact of global climate change on terrestrial ecosystems. Additionally, it is not clear how the effects of global warming on grassland productivity are going to be altered by increased N deposition and N addition.

Methodology/Principal Findings

In-situ canopy CO₂ exchange rates were measured in a meadow steppe subjected to 4-year warming and nitrogen addition treatments. Warming treatment reduced net ecosystem CO₂ exchange (NEE) and increased ecosystem respiration (ER); but had no significant impacts on gross ecosystem productivity (GEP). N addition increased NEE, ER and GEP. However, there were no significant interactions between N addition and warming. The variation of NEE during the four experimental years was correlated with soil water content, particularly during early spring, suggesting that water availability is a primary driver of carbon fluxes in the studied semi-arid grassland.

Conclusion/Significance

Ecosystem carbon fluxes in grassland ecosystems are sensitive to warming and N addition. In the studied water-limited grassland, both warming and N addition influence ecosystem carbon fluxes by affecting water availability, which is the primary driver in many arid and semiarid ecosystems. It remains unknown to what extent the long-term N addition would affect the turn-over of soil organic matter and the C sink size of this grassland.     

Changes in nitrogen resorption traits of six temperate grassland species along a multi-level N addition gradient

Nitrogen (N) resorption from senescing leaves is an important mechanism of N conservation for terrestrial plant species, but changes in N-resorption traits over wide-range and multi-level N addition gradients have not been well characterized. Here, a 3-year N addition experiment was conducted to determine the effects of N addition on N resorption of six temperate grassland species belonging to three different life-forms: Stipa krylovii Roshev. (grass), Cleistogenes squarrosa (T.) Keng (grass), Artemisia frigida Willd. (semishrub), Melissitus ruthenica C.W.Wang (semishrub and N-fixer), Potentilla acaulis L. (forb) and Allium bidentatum Fisch.ex Prokh. (forb). Generally, N concentrations in green leaves increased asymptotically for all species. N concentrations in senescent leaves for most species (5/6) also increased asymptotically, except that the N concentration in senescent leaves of A. bidentatum was independent of N addition. N-resorption efficiency decreased with increasing N addition level only for S. krylovii and A. frigida, while no clear responses were found for other species. These results suggest that long-term N fertilization increased N uptake and decreased N-resorption proficiency, but the effects on N-resorption efficiency were species-specific for different temperate grassland species in northern China. These inter-specific differences in N resorption may influence the positive feedback between species dominance and N availability and thus soil N cycling in the grassland ecosystem in this region.     

Nitrogen, phosphorus and potassium nutritional status of semiarid steppe grassland in Inner Mongolia

In grazed semiarid steppe ecosystems, much attention has been paid to aspects of growth limitation by water. So far, potential limitation of primary production by plant nutrients was rarely considered. This knowledge is essential for identification of sustainable land-use practices in these large and important ecosystems on the background of over-exploitation and climate change. In the present study plant nutrient concentrations and ratios were investigated with factorial additions of water and N fertilizer at two sites with contrasting soil nutrient availability. Combined analysis of nutrient concentrations, contents, biomass production, and plant N:P ratios consistently confirmed primary growth limitation by water and a strong N limitation when sufficient amounts of water were supplied. P limitation only occurred at the site with low P availability when in addition to the natural supply, water and N fertilizer were given. According to reported thresholds of N:K and K:P ratios, K was not limiting in any plot. The observed nutritional patterns in the plant community were related to the dynamics of species composition and their specific nutrient status. Stipa grandis had the highest N:P ratio whereas Artemisia frigida showed lowest N:P. These nutrient characteristics were related to growth strategies of dominant species. Accordingly, the relative biomass contribution of S. grandis and A. frigida strongly affected the nutrient status of the plant community. Plant N:P ratios indicate the relative limitation by N or P in the semiarid grasslands under sufficient water supply, but other methods of nutritional diagnosis should be used when plant N:P ratios remain below critical values.     

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.     

Transgenerational effect alters the interspecific competition between two dominant species in a temperate steppe

One of the key aims of global change studies is to predict more accurately how plant community composition responds to future environmental changes. Although interspecific relationship is one of the most important forces structuring plant communities, it remains a challenge to integrate long‐term consequences at the plant community level. As an increasing number of studies have shown that maternal environment affects offspring phenotypic plasticity as a response to global environment change through transgenerational effects, we speculated that the transgenerational effect would influence offspring competitive relationships. We conducted a 10‐year field experiment and a greenhouse experiment in a temperate grassland in an Inner Mongolian grassland to examine the effects of maternal and immediate nitrogen addition (N) and increased precipitation (Pr) on offspring growth and the interspecific relationship between the two dominant species, Stipa krylovii and Artemisia frigida. According to our results, Stipa kryloii suppressed A. frigida growth and population development when they grew in mixture, although immediate N and Pr stimulated S. kryloii and A. frigida growth simultaneously. Maternal N and Pr declined S. krylovii dominance and decreased A. frigida competitive suppression to some extent. The transgenerational effect should further facilitate the coexistence of the two species under scenarios of increased nitrogen input and precipitation. If we predicted these species'' interspecific relationships based only on immediate environmental effects, we would overestimate S. krylovii''s competitive advantage and population development, and underestimate competitive outcome and population development of A. frigida. In conclusion, our results demonstrated that the transgenerational effect of maternal environment on offspring interspecific competition must be considered when evaluating population dynamics and community composition under the global change scenario.     

Complete Chloroplast Genome Sequences of Mongolia Medicine Artemisia frigida and Phylogenetic Relationships with Other Plants

Background

Artemisia frigida Willd. is an important Mongolian traditional medicinal plant with pharmacological functions of stanch and detumescence. However, there is little sequence and genomic information available for Artemisia frigida, which makes phylogenetic identification, evolutionary studies, and genetic improvement of its value very difficult. We report the complete chloroplast genome sequence of Artemisia frigida based on 454 pyrosequencing.

Methodology/Principal Findings

The complete chloroplast genome of Artemisia frigida is 151,076 bp including a large single copy (LSC) region of 82,740 bp, a small single copy (SSC) region of 18,394 bp and a pair of inverted repeats (IRs) of 24,971 bp. The genome contains 114 unique genes and 18 duplicated genes. The chloroplast genome of Artemisia frigida contains a small 3.4 kb inversion within a large 23 kb inversion in the LSC region, a unique feature in Asteraceae. The gene order in the SSC region of Artemisia frigida is inverted compared with the other 6 Asteraceae species with the chloroplast genomes sequenced. This inversion is likely caused by an intramolecular recombination event only occurred in Artemisia frigida. The existence of rich SSR loci in the Artemisia frigida chloroplast genome provides a rare opportunity to study population genetics of this Mongolian medicinal plant. Phylogenetic analysis demonstrates a sister relationship between Artemisia frigida and four other species in Asteraceae, including Ageratina adenophora, Helianthus annuus, Guizotia abyssinica and Lactuca sativa, based on 61 protein-coding sequences. Furthermore, Artemisia frigida was placed in the tribe Anthemideae in the subfamily Asteroideae (Asteraceae) based on ndhF and trnL-F sequence comparisons.

Conclusion

The chloroplast genome sequence of Artemisia frigida was assembled and analyzed in this study, representing the first plastid genome sequenced in the Anthemideae tribe. This complete chloroplast genome sequence will be useful for molecular ecology and molecular phylogeny studies within Artemisia species and also within the Asteraceae family.     

Scale-Dependent Effects of Grazing on Plant C: N: P Stoichiometry and Linkages to Ecosystem Functioning in the Inner Mongolia Grassland

Background

Livestock grazing is the most prevalent land use of grasslands worldwide. The effects of grazing on plant C, N, P contents and stoichiometry across hierarchical levels, however, have rarely been studied; particularly whether the effects are mediated by resource availability and the underpinning mechanisms remain largely unclear.

Methodology/Principal Findings

Using a multi-organization-level approach, we examined the effects of grazing on the C, N, and P contents and stoichiometry in plant tissues (leaves and roots) and linkages to ecosystem functioning across three vegetation types (meadow, meadow steppe, and typical steppe) in the Inner Mongolia grassland, China. Our results showed that the effects of grazing on the C, N, and P contents and stoichiometry in leaves and roots differed substantially among vegetation types and across different hierarchical levels (species, functional group, and vegetation type levels). The magnitude of positive effects of grazing on leaf N and P contents increased progressively along the hierarchy of organizational levels in the meadow, whereas its negative effect on leaf N content decreased considerably along hierarchical levels in both the typical and meadow steppes. Grazing increased N and P allocation to aboveground in the meadow, while greater N and P allocation to belowground was found in the typical and meadow steppes. The differences in soil properties, plant trait-based resource use strategies, tolerance or defense strategies to grazing, and shifts in functional group composition are likely to be the key mechanisms for the observed patterns among vegetation types.

Conclusions/Significance

Our findings suggest that the enhanced vegetation-type-level N contents by grazing and species compensatory feedbacks may be insufficient to prevent widespread declines in primary productivity in the Inner Mongolia grassland. Hence, it is essential to reduce the currently high stocking rates and restore the vast degraded steppes for sustainable development of arid and semiarid grasslands.     

Independent effects of warming and nitrogen addition on plant phenology in the Inner Mongolian steppe

Background and Aims

Phenology is one of most sensitive traits of plants in response to regional climate warming. Better understanding of the interactive effects between warming and other environmental change factors, such as increasing atmosphere nitrogen (N) deposition, is critical for projection of future plant phenology.

Methods

A 4-year field experiment manipulating temperature and N has been conducted in a temperate steppe in northern China. Phenology, including flowering and fruiting date as well as reproductive duration, of eight plant species was monitored and calculated from 2006 to 2009.

Key Results

Across all the species and years, warming significantly advanced flowering and fruiting time by 0·64 and 0·72 d per season, respectively, which were mainly driven by the earliest species (Potentilla acaulis). Although N addition showed no impact on phenological times across the eight species, it significantly delayed flowering time of Heteropappus altaicus and fruiting time of Agropyron cristatum. The responses of flowering and fruiting times to warming or N addition are coupled, leading to no response of reproductive duration to warming or N addition for most species. Warming shortened reproductive duration of Potentilla bifurca but extended that of Allium bidentatum, whereas N addition shortened that of A. bidentatum. No interactive effect between warming and N addition was found on any phenological event. Such additive effects could be ascribed to the species-specific responses of plant phenology to warming and N addition.

Conclusions

The results suggest that the warming response of plant phenology is larger in earlier than later flowering species in temperate grassland systems. The effects of warming and N addition on plant phenology are independent of each other. These findings can help to better understand and predict the response of plant phenology to climate warming concurrent with other global change driving factors.     

Impact of historic grazing on steppe soils on the northern Tibetan Plateau

Aims

Few studies have focused on changes in the physical and chemical properties of soils that are induced by grazing at high altitudes. Our aim was to identify potential responses of soil to grazing pressure on the semiarid steppe of the northern Tibetan Plateau and their probable causes.

Methods

Fractal geometry to describe soil structure, soil dynamics, and physical processes within soil is becoming an increasingly useful tool that allows a better understanding of the performance of soil systems. In this study, we sampled four experimental areas in the northern part of the Tibetan Plateau under different grazing intensities: ungrazed, lightly grazed, moderately grazed and heavily grazed plots. Fractal methods were applied to characterise particle-size distributions and pore patterns of soils under different grazing intensities.

Results

Our results reveal a highly significant decrease in the fractal dimensions of particle size distributions (D ₁ ) and the fractal dimensions of all pores (D ₂ ) with increasing grazing intensity. Soil organic carbon (SOC), total N and total P concentrations increased significantly with decreasing grazing intensity. We did not find differences in soil pH in response to grazing.

Conclusions

Grazing induced a significant deterioration of the physical and chemical topsoil properties in the semiarid steppe of the northern Tibetan Plateau. Fractal dimensions can be a useful parameter for quantifying soil degradation due to human activities.     

Soil microbial properties and plant growth responses to carbon and water addition in a temperate steppe: the importance of nutrient availability

Background

Global climatic change is generally expected to stimulate net primary production, and consequently increase soil carbon (C) input. The enhanced C input together with potentially increased precipitation may affect soil microbial processes and plant growth.

Methodology/Principal Findings

To examine the effects of C and water additions on soil microbial properties and plant growth, we conducted an experiment lasting two years in a temperate steppe of northeastern China. We found that soil C and water additions significantly affected microbial properties and stimulated plant growth. Carbon addition significantly increased soil microbial biomass and activity but had a limited effect on microbial community structure. Water addition significantly increased soil microbial activity in the first year but the response to water decreased in the second year. The water-induced changes of microbial activity could be ascribed to decreased soil nitrogen (N) availability and to the shift in soil microbial community structure. However, no water effect on soil microbial activity was visible under C addition during the two years, likely because C addition alleviated nutrient limitation of soil microbes. In addition, C and water additions interacted to affect plant functional group composition. Water addition significantly increased the ratio of grass to forb biomass in C addition plots but showed only minor effects under ambient C levels. Our results suggest that soil microbial activity and plant growth are limited by nutrient (C and N) and water availability, and highlight the importance of nutrient availability in modulating the responses of soil microbes and plants to potentially increased precipitation in the temperate steppe.

Conclusions/Significance

Increased soil C input and precipitation would show significant effects on soil microbial properties and plant growth in the temperate steppe. These findings will improve our understanding of the responses of soil microbes and plants to the indirect and direct climate change effects.     

Tradeoffs between nitrogen- and water-use efficiency in dominant species of the semiarid steppe of Inner Mongolia

In water-limited environments, photosynthetic carbon gain and loss of water by transpiration are in a permanent tradeoff as both are contrarily regulated by stomata conductance. In semiarid steppe grasslands water limitation may covary with nitrogen limitation. Steppe grassland species are capable of optimizing their use of limiting resources, water and nitrogen, but regulation is still poorly understood. In a two-year experiment with addition of water (irrigation simulating a wet year) and nitrogen (0, 25, and 50 kg urea-N?ha^?1) we assessed intrinsic water use efficiency (WUE_i), nitrogen use efficiency (NUE), and related plant functional traits (PFTs) of four dominant C₃ species (Leymus chinensis, Agropyron cristatum, Stipa grandis, and Artemisia frigida). Water and N fertilizer supplementation significantly increased plant primary production, and N effect was more pronounced under irrigated conditions. Parallel with the responses of plant production, a strong tradeoff between WUE_i and NUE was detected: water supply increased NUE but decreased WUE_i, whereas N addition slightly increased WUE_i at the expense of NUE. This tradeoff occurred at the leaf level, and involved the responses of leaf N concentration and specific leaf area. WUE_i of species changed among treatments in a predictable manner by the parameter of leaf N content per area. Dominant plant species commonly achieved a higher utilization efficiency of the more limiting resource via altering PFTs, which was an important mechanism of adaptation to variable resource limitation in semiarid grasslands.     

Plant Functional Diversity and Species Diversity in the Mongolian Steppe

Background

The Mongolian steppe is one of the most important grasslands in the world but suffers from aridization and damage from anthropogenic activities. Understanding structure and function of this community is important for the ecological conservation, but has seldom been investigated.

Methodology/Principal Findings

In this study, a total of 324 quadrats located on the three main types of Mongolian steppes were surveyed. Early-season perennial forbs (37% of total importance value), late-season annual forbs (33%) and late-season perennial forbs (44%) were dominant in meadow, typical and desert steppes, respectively. Species richness, diversity and plant functional type (PFT) richness decreased from the meadow, via typical to desert steppes, but evenness increased; PFT diversity in the desert and meadow steppes was higher than that in typical steppe. However, above-ground net primary productivity (ANPP) was far lower in desert steppe than in the other two steppes. In addition, the slope of the relationship between species richness and PFT richness increased from the meadow, via typical to desert steppes. Similarly, with an increase in species diversity, PFT diversity increased more quickly in both the desert and typical steppes than that in meadow steppe. Random resampling suggested that this coordination was partly due to a sampling effect of diversity.

Conclusions/Significance

These results indicate that desert steppe should be strictly protected because of its limited functional redundancy, which its ecological functioning is sensitive to species loss. In contrast, despite high potential forage production shared by the meadow and typical steppes, management of these two types of steppes should be different: meadow steppe should be preserved due to its higher conservation value characterized by more species redundancy and higher spatial heterogeneity, while typical steppe could be utilized moderately because its dominant grass genus Stipa is resistant to herbivory and drought.     

Plant Trait Diversity Buffers Variability in Denitrification Potential over Changes in Season and Soil Conditions

Background

Denitrification is an important ecosystem service that removes nitrogen (N) from N-polluted watersheds, buffering soil, stream, and river water quality from excess N by returning N to the atmosphere before it reaches lakes or oceans and leads to eutrophication. The denitrification enzyme activity (DEA) assay is widely used for measuring denitrification potential. Because DEA is a function of enzyme levels in soils, most ecologists studying denitrification have assumed that DEA is less sensitive to ambient levels of nitrate (NO₃ ⁻) and soil carbon and thus, less variable over time than field measurements. In addition, plant diversity has been shown to have strong effects on microbial communities and belowground processes and could potentially alter the functional capacity of denitrifiers. Here, we examined three questions: (1) Does DEA vary through the growing season? (2) If so, can we predict DEA variability with environmental variables? (3) Does plant functional diversity affect DEA variability?

Methodology/Principal Findings

The study site is a restored wetland in North Carolina, US with native wetland herbs planted in monocultures or mixes of four or eight species. We found that denitrification potentials for soils collected in July 2006 were significantly greater than for soils collected in May and late August 2006 (p<0.0001). Similarly, microbial biomass standardized DEA rates were significantly greater in July than May and August (p<0.0001). Of the soil variables measured—soil moisture, organic matter, total inorganic nitrogen, and microbial biomass—none consistently explained the pattern observed in DEA through time. There was no significant relationship between DEA and plant species richness or functional diversity. However, the seasonal variance in microbial biomass standardized DEA rates was significantly inversely related to plant species functional diversity (p<0.01).

Conclusions/Significance

These findings suggest that higher plant functional diversity may support a more constant level of DEA through time, buffering the ecosystem from changes in season and soil conditions.     

Litter species traits, but not richness, contribute to carbon and nitrogen dynamics in an alpine meadow on the Tibetan Plateau

Aims

Litter, as afterlife of plants, plays an important role in driving belowground decomposition processes. Here we tested effects of litter species identity and diversity on carbon (C) and nitrogen (N) dynamics during litter decomposition in N-limited alpine meadow soil from the Qinghai–Tibet Plateau.

Methods

We incubated litters of four meadow species, a sedge (“S”, Kobresia humilis), a grass (“G”, Elymus nutans), a herb (“H”, Saussurea superba), and a legume (“L”, Oxytropis falcata), in monoculture and in mixture with meadow soil. CO₂ release was measured 21 times during the incubation, and soil available N and microbial biomass C and N were measured before and after the experiment.

Results

The organic C decay rate did not differ much among soils amended with monocultures or mixtures of litter, except in the H, S, L, and S+H treatments, which had much higher decay rates. Potential decomposable C pools were lowest in the control, highest in the L treatment, and intermediate in the S treatment. Mineralized N was completely immobilized by soil microbes in all treatments except the control, S+L, and S+G+L treatments. Litter mixtures had both additive and non-additive effects on CO₂-C emission (mainly antagonistic effects), net N mineralization (mainly synergistic), and microbial biomass C and N (both). Overall, these parameters were not significantly correlated with litter species richness. Similarly, microbial C or N was not significantly correlated with litter N content or C/N. However, cumulative CO₂-C emission and net N mineralization were positively correlated with litter N content and negatively correlated with litter C/N.

Conclusions

Litter N content and C/N rather than litter species richness drove the release of CO₂-C and net available N in this ecosystem. The antagonistic effects of litter mixtures contributed to a modest release of CO₂-C, but their synergistic effects enhanced net available N. We suggest that in alpine meadow communities, balancing species with high and low N contents will benefit soil carbon sequestration and plant competition for available N with soil microbes.