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Plant root variations and their relations with soil moisture and nutrient supply have been well documented for many species, while effects of drought, combined with extreme poor soil nutrients, on plant roots remain unclear.Herein, we addressed root vertical distributions of two typical xerophyte semishrub species, Artemisia sphaerocephala and A. intramongolica, and their relations with soil moisture, total soil nitrogen and carbon contents in arid Hunshandake desert, China. The two species experienced similar light regimes and precipitation, but differed in soil moisture and soil nutrients.Root vertical distribution patterns (e.g., coarse root diameter, root depth and root biomass) differed considerable for the two species due to high heterogeneity of soil environments. Coarse and fine root biomasses for A. intramongolica, distributed in relatively moist fixed dunes, mainly focused on surface layers (94%); but those for A. sphaerocephala dropped gradually from the surface to 140 cm depth. Relations between root traits (e.g., diameter, root biomass) and soil moisture were positive for A. intramongolica, but those for A. sphaerocephala were negative.In general, the root traits for both species positively correlated with total soil nitrogen and carbon contents. These findings suggest that both soil moisture and poor soil nutrients were the limiting resources for growth and settlement of these two species., The root traits for Artemisia sphaerocephala and A. intramongolica were positively correlated with total soil nitrogen and carbon contents. These findings indicate that both soil moisture and poor soil nutrients were the limiting resources for growth in these two species.

Nitrogen (N) is a major limiting element for productivity in temperate grasslands, particularly during early spring when soil N availability is low and the vegetative demand for it is high. Therefore, knowing whether and how plant species adopt different N acquisition strategies during the winter-spring transitional period is essential for understanding ecosystem functioning in temperate grasslands. In this study, parallel experiments with 15N tracer were conducted to examine plant N acquisition strategies during winter-spring transition in a meadow and a typical steppe in northern China. We found that soil microbes immobilized ∼20% of the 15N tracer during the spring thawing period at both sites, and then released half of it back to the soil before late spring, confirming that soil microbes competed effectively with the plant roots for mineral N in early spring. Perennial bunch grasses adopted an active N acquisition strategy at the beginning of the spring thawing period. In contrast, perennial forbs and rhizome grasses began to take up N in the middle of the spring thawing period, and they acquired more N than the bunch grasses. However, sagebrushes and legumes accounted for little 15N recovery, indicating their dependence on internal N accumulation or N fixation. At the ecosystem level, no significant difference in the magnitude of plant 15N uptake was observed between the meadow steppe and typical steppe, although the plant biomass N in the meadow steppe was twice that of the typical steppe during the thawing period. This was attributed to the higher soil inorganic N and faster net N mineralization rate in the meadow steppe than in the typical steppe. Our results suggest that temporal niche differentiation in N acquisition during early spring may facilitate species coexistence in temperate grasslands despite strong plant-microbe or plant-plant competition for N. The divergent N acquisition strategies at the species level and convergent N acquisition strategies at the ecosystem level should be considered for model development to better simulate vegetation growth particularly under spring N stress.

In nitrogen (N)-limited temperate regions, winter is an important period of N accumulation. The accumulated N is released during snowmelt and thawing, and the availability peaks in early spring. However, the early-spring dynamics of specific N forms (i.e., ammonium NH4+ and nitrate NO3−) in temperate grasslands are still not fully understood. Here, we added 15NH4+ and 15NO3− (equivalent to 150 mg 15N m−2) to the soils of a meadow steppe and a typical steppe in northern China immediately after snowmelt, then quantified the retention dynamics of 15NH4+ and 15NO3− in soils, microbes, and plants over the subsequent growing season. Approximately 70% of the added 15N tracers were initially retained within the soil−microbe−plant systems in both temperate grasslands. In early spring, much 15N was immobilized in soils and microbes, while little had been taken up by plants. During the subsequent growing season, approximately 45% of the 15N was rapidly lost by the soils and microbes, but plant 15N acquisition gradually increased. Although soils and plants retained similar levels of 15NH4+ and 15NO3− during the growing season, soil microbes retained more 15NH4+ than 15NO3−. Different plant taxa had distinct 15N acquisition capacities: perennial grasses and forbs accumulated the 15N tracers rapidly, while annuals did not. Perennial grasses were effective immobilizers of the 15NH4+, whereas forbs were effective immobilizers of the 15NO3−. These findings provided evidence of the substantial retention of early-spring N over the following growing season in temperate grasslands, regardless of the vegetation type and N form. However, it was clear that the dynamics of early-spring 15NH4+ and 15NO3− differed within the soil−microbe−plant systems.

In N-limited temperate regions, atmospheric N deposition remains high over the non-growing season. However, the retention dynamics of non-growing season N input within the ecosystem remain unclear. Using an isotopic approach, we investigated the initial retention and subsequent dynamics of 15N (1.5 g 15N m−2) in the soils, microbes, plants, and litter over three years in grazing-prohibited (PG) and heavily grazed treatments (HG) in northern China. For initial retention (21 days after 15N addition), most 15N was immobilized in soils and microbes, while less was taken up by plants. Soil and microbial 15N immobilization were significantly higher when grazing was prohibited, although plant 15N acquisition was not affected by grazing. After initial retention, rapid 15N loss was observed in microbes and soils, while 15N levels were sustained longer in plants and litter. The 15N residence times were longer when grazing was prohibited. The 15N acquisition capacity varies among plant taxa: perennial grasses and forbs accumulated 15N rapidly, while sagebrush and legumes acted slowly. Although the added 15N had significant contribution to early spring N demands of soil microbes and plants, it did not increase microbial or plant biomass N. Our results showed that non-growing season exogenous N was primarily retained by soil biota in temperate grasslands in the early stage, but N retention is finally sustained in soil and plants. The findings highlighted the importance of soil microbes in sustaining N upon N input, inferring the needs for considering the microbial role for better understanding N cycling in the temperate grasslands.

Soil carbon (C) and nitrogen (N) cycling are sensitive to changes in environmental factors and play critical roles in the responses of terrestrial ecosystems to natural and anthropogenic perturbations. This study was conducted to quantify the effects of belowground particulate litter (BPL) addition, increased precipitation and their interactions on soil C and N mineralization in two adjacent sites where belowground photosynthate allocation was manipulated through vegetation clipping in a temperate steppe of northeastern China from 2010 to 2011. The results show that BPL addition significantly increase soil C mineralization rate (CMR) and net N mineralization rate (NMR). Although increased precipitation-induced enhancement of soil CMR essentially ceased after the first year, stimulation of soil NMR and net nitrification rate continued into the second year. Clipping only marginally decreased soil CMR and NMR during the two years. There were significant synergistic interactions between BPL addition (and increased precipitation) and clipping on soil CMR and NMR, likely to reflect shifts in soil microbial community structure and a decrease in arbuscular mycorrhizal fungi biomass due to the reduction of belowground photosynthate allocation. These results highlight the importance of plants in mediating the responses of soil C and N mineralization to potentially increased BPL and precipitation by controlling belowground photosynthate allocation in the temperate steppe.

Global climate change is predicted to stimulate primary production and consequently increases litter inputs. Changing precipitation regimes together with enhanced litter inputs may affect plant community composition and structure, with consequent influence on diversity and ecosystem functioning. Responses of plant community to increased precipitation and belowground litter addition were examined lasting 5 years in a semiarid temperate grassland of northeastern China. Increased precipitation enhanced community species richness and abundance of annuals by 16.8% and 44%, but litter addition suppressed them by 25% and 54.5% after 5 years, respectively. During the study period, perennial rhizome grasses and forbs had consistent negative relationship under ambient plots, whereas positive relationship between the two functional groups was found under litter addition plots after 5 years. In addition, increased precipitation and litter addition showed significant interaction on community composition, because litter addition significantly increased biomass and abundance of rhizome grasses under increased precipitation plots but had no effect under ambient precipitation levels. Our findings emphasize the importance of water availability in modulating the responses of plants community to potentially enhanced litter inputs in the semiarid temperate grassland.

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.

Background Although it has been widely accepted that global changes will pose the most important constrains to plant survival and distribution, our knowledge of the adaptive mechanism for plant with large-scale environmental changes (e.g. drought and high temperature) remains limited. Methodology/principal findings An experiment was conducted to examine anatomical and physiological plasticity in Leymus chinensis along a large-scale geographical gradient from 115° to 124°E in northeast China. Ten sites selected for plant sampling at the gradient have approximately theoretical radiation, but differ in precipitation and elevation. The significantly increasing in leaf thickness, leaf mass per area, vessel and vascular diameters, and decreasing in stoma density and stoma index exhibited more obvious xerophil-liked traits for the species from the moist meadow grassland sites in contrast to that from the dry steppe and desert sites. Significant increase in proline and soluble sugar accumulation, K(+)/Na(+) for the species with the increasing of stresses along the gradient showed that osmotic adjustment was enhanced. Conclusion/significance Obvious xerophytic anatomical traits and stronger osmotic adjustment in stress conditions suggested that the plants have much more anatomical and physiological flexibilities than those in non-stress habitats along the large-scale gradient.