Responses of C:N:P stoichiometric correlations among plants, soils and microorganisms to warming: A meta-analysis.

Plants, soils and microorganisms play important roles in maintaining stable terrestrial stoichiometry. Studying how nutrient balances of these biotic and abiotic players vary across temperature gradients is important when predicting ecosystem changes on a warming planet. The respective responses of plant, soil and microbial stoichiometric ratios to warming have been observed, however, whether and how the stoichiometric correlations among the three components shift under warming has not been clearly understood and identified. In the present study, we have performed a meta-analysis based on 600 case studies from 74 sites or locations to clarify whether and how warming affects plant, soil and microbial stoichiometry, respectively, and their correlations. Our results indicated that: (1) globally, plants had higher C:N and C:P values compared to soil and microbial pools, but their N:P distributions were similar; (2) warming did not significantly alter plant, soil and microbial C:N and C:P values, but had a noticeable effect on plant N:P ratios. When ecosystem types, duration and magnitude of warming were taken into account, there was an inconsistent and even inverse warming response in terms of the direction and magnitude of changes in the C:N:P ratios occurring among plants, soils and microorganisms; (3) despite various warming responses of the stoichiometric ratios detected separately for plants, soils and microorganisms, the stoichiometric correlations among all three parts remained constant even under different warming scenarios. Our study highlighted the complexity of the effect of warming on the C:N:P stoichiometry, as well as the absence and importance of simultaneous measurements of stoichiometric ratios across different components of terrestrial ecosystems, which should be urgently strengthened in future studies.

Carbon, nitrogen, and phosphorus dynamics in China's lakes: climatic and geographic influences.

Given the differences in geomorphology, climate, hydrology, and human activities in various regions, lake chemometrics may also vary. However, the spatial distribution of lake chemistry and the factors affecting such pattern are still unclear. Here, we collected data for carbon, nitrogen, and phosphorus from published literature and databases in 224 lakes and calculated the trophic status index to represent the nutrient classification state of lakes. We found that lakes with high carbon concentrations were located in the Tibet-Qinghai Limnetic Region of western China, whereas lakes with high nitrogen and phosphorus concentrations were located in the Inner Mongolia-Xinjiang Limnetic Region and Northeast Limnetic Region of northern China. Areas with larger cropland and urban residential land (such as the junction of the three lake regions, i.e., the Northeast Limnetic Region, East Limnetic Region, and Inner Mongolia-Xinjiang Limnetic Region) tended to have lakes with high nitrogen and phosphorus concentrations. Our analysis suggested that spatial distribution of carbon, nitrogen, and phosphorus concentrations reflect the effect of climate, geomorphology, and land use in each lake region and nationwide.

Aridity modifies the responses of plant stoichiometry to global warming and nitrogen deposition in semi-arid steppes.

Global warming and nitrogen (N) deposition are known to unbalance the stoichiometry of carbon (C), N, and phosphorus (P) in terrestrial plants, but it is unclear how water availability regulates their effects along a natural aridity gradient. Here, we conducted manipulative experiments to determine the effects of experimental warming (WT) and N addition (NT) on plant stoichiometry in desert, typical, and meadow steppes with decreasing aridity. WT elevated air temperatures by 1.2-2.9 °C using open-top chambers. WT increased forb C:N ratio and thus its N use efficiency and competitiveness in desert steppes, whereas WT reduced forb C:N and C:P ratios in typical and meadow steppes. Plant N:P ratio, which reflects nutrient limitation, was reduced by WT in desert steppes but not for typical or meadow steppes. NT reduced plant C:N ratios and increased N:P ratios in all three steppes. NT reduced forb C:P ratios in desert and typical steppes, but it enhanced grass C:P ratio in meadow steppes, indicating an enhancement of P use efficiency and competitiveness of grasses in wet steppes. WT and NT had synergetic effects on grass C:N and C:P ratios in all three steppes, which helps to increase grasses' productivity. Under WT or NT, the changes in community C:N ratio were positively correlated with increasing aridity, indicating that aridity increases plants' N use efficiency. However, aridity negatively affected the changes in N:P ratios under NT but not WT, which suggests that aridity mitigates P limitation induced by N deposition. Our results imply that warming could shift the dominant functional group into forbs in dry steppes due to altered stoichiometry, whereas grasses become dominated plants in wet steppes under increasing N deposition. We suggest that global changes might break the stoichiometric balance of plants and water availability could strongly modify such processes in semi-arid steppes.

Aridity stimulates responses of root production and turnover to warming but suppresses the responses to nitrogen addition in temperate grasslands of northern China.

Global warming and nitrogen (N) deposition are known to affect root dynamics in grasslands. However, previous studies were based only on a single ecosystem type, so it is unclear how warming and N addition affect root traits (root biomass, root-shoot ratio, root production and turnover) along the aridity gradient. In this study, we conducted an experiment to determine the effects of warming and N addition on root traits in desert, typical, and meadow grasslands in northern China, where the aridity gradually decreases from west to east across the region. Warming increased root-shoot ratio in dry year due to decline in soil water, but had a downward trend in root production and turnover in all three grasslands. N addition decreased root-shoot ratio in humid year due to increase in soil N, whereas did not significantly affect root production in any grasslands and increased root turnover in desert and meadow grasslands rather than typical grassland. Warming combined with N addition had negatively additive effects on root turnover in typical and meadow grasslands rather than desert grassland. N addition-induced changes in root biomass and root-shoot ratio were negatively affected by aridity in dry year. Aridity positively affected responses of root production and turnover to warming but negatively affected those responses to N addition. However, root-shoot ratio, root production and turnover under warming combined with N addition were not affected by aridity. Our results suggest that warming suppresses root carbon (C) input but N addition may exacerbate it in temperate grasslands, and warming combined with N addition suppresses it only in wet grasslands. Aridity promotes root C input under warming but suppresses it under N addition. However, aridity may little affect soil C and nutrient dynamics under global warming combined with N deposition in temperate grasslands in the future.

Increasing aridity, temperature and soil pH induce soil C-N-P imbalance in grasslands.

Due to the different degrees of controls exerted by biological and geochemical processes, climate changes are suggested to uncouple biogeochemical C, N and P cycles, influencing biomass accumulation, decomposition and storage in terrestrial ecosystems. However, the possible extent of such disruption in grassland ecosystems remains unclear, especially in China's steppes which have undergone rapid climate changes with increasing drought and warming predicted moving forward in these dryland ecosystems. Here, we assess how soil C-N-P stoichiometry is affected by climatic change along a 3500-km temperate climate transect in Inner Mongolia, China. Our results reveal that the soil from more arid and warmer sites are associated with lower soil organic C, total N and P. The ratios of both soil C:P and N:P decrease, but soil C:N increases with increasing aridity and temperature, indicating the predicted decreases in precipitation and warming for most of the temperate grassland region could lead to a soil C-N-P decoupling that may reduce plant growth and production in arid ecosystems. Soil pH, mainly reflecting long-term climate change in our sites, also contributes to the changing soil C-N-P stoichiometry, indicating the collective influences of climate and soil type on the shape of soil C-N-P balance.

[Latitudinal Changes in Plant Stoichiometric and Soil C, N, P Stoichiometry in Loess Plateau].

Field investigations and sampling were conducted in Loess Plateau, including Fu County, Ganquan County, Ansai County, Jingbian County and Hengshan County and Yuyang District. Our objective was to examine changes of leaf and soil stoichiometry characteristics along latitudinal gradient in Loess Plateau, and to provide references for the prediction of soil nutrient status of the ecosystem and constraints of plant nutrition elements in Loess Plateau. The results showed that (1) Across the 35.95 degrees-38.36 degrees N latitude gradient, leaf C, N and P stoichiometry were ranging from 336.95 to 477.38 mg x g(-1) for C, from 18.09 to 33.173 mg x g(-1) for N and from 1.07 to 1.73 mg x g(-1) for P, the arithmetic means were 442.9 mg x g(-1), 25.79 mg x g(-1) and 1.37 mg x g(-1), separately, the variation coefficients were 11.9%, 17.4% and 13.3%. There were obvious correlation between leaf C, N, P and latitude, leaf C, C : N ratio and C: P ratio significantly decreased with the increasing latitude, while leaf N and P significantly increased with the increasing latitude. The relationship between N: P ratio and latitude was not significant. (2) The content of soil organic C and soil total N decreased with increasing latitude and soil layer. In contrast, with the increase of latitude, soil P increased and then decreased. In the 0-10 cm, 10-20 cm soil layers, soil C: N ratio did not change significantly with latitude, while in the 20-40 cm layer, C: N ratio decreased obviously, but soil C: P and N: P ratios decreased with the increasing latitude in all soil layers. (3) Leaf C, C: N and C: P ratios were correlated to soil organic C, soil total N and soil total P in all soil layers, leaf N and P were correlated to soil organic C and soil total N, while leaf N: P ratio was not correlated to soil organic C, soil total N and soil total P. There was a certain correlation between the leaf C, N, P and latitude, however, the correlations between leaf and soil C, N, P were inconsistent. These results demonstrate that the plants were under P limitation in Loess Plateau.

Effects of thinning intensities on soil infiltration and water storage capacity in a Chinese pine-oak mixed forest.

Thinning is a crucial practice in the forest ecosystem management. The soil infiltration rate and water storage capacity of pine-oak mixed forest under three different thinning intensity treatments (15%, 30%, and 60%) were studied in Qinling Mountains of China. The thinning operations had a significant influence on soil infiltration rate and water storage capacity. The soil infiltration rate and water storage capacity in different thinning treatments followed the order of control (nonthinning): <60%, <15%, and <30%. It demonstrated that thinning operation with 30% intensity can substantially improve soil infiltration rate and water storage capacity of pine-oak mixed forest in Qinling Mountains. The soil initial infiltration rate, stable infiltration rate, and average infiltration rate in thinning 30% treatment were significantly increased by 21.1%, 104.6%, and 60.9%, compared with the control. The soil maximal water storage capacity and noncapillary water storage capacity in thinning 30% treatment were significantly improved by 20.1% and 34.3% in contrast to the control. The soil infiltration rate and water storage capacity were significantly higher in the surface layer (0~20 cm) than in the deep layers (20~40 cm and 40~60 cm). We found that the soil property was closely related to soil infiltration rate and water storage capacity.

[Fine root biomass and production of four vegetation types in Loess Plateau, China].

Fine roots (≤ 2 mm) play a major role in biogeochemical cycling in ecosystems. By the methods of soil cores and ingrowth soil cores, we studied the biomass and annual production of fine roots in 0-40 cm soil layers of four main vegetation types, i. e. , Robinia pseudoacacia plantation, deciduous shrubs, abandoned grassland, and Artemisia desertorum community in Loess Plateau, China. The spatial patterns of fine root biomass and production were negatively associated with latitudes. The fine root biomass in the 0-40 cm soil layer was in the order of deciduous shrubs (220 g · m(-2)), R. pseudoacacia plantation (163 g · m(-2)), abandoned grassland (162 g · m(-2)) and A. desertorum community (79 g · m(-2)). The proportion of ≤ 1 mm fine root biomass (74.1%) in the 0-40 cm soil layer of abandoned grassland was significantly higher than those in the other three vegetation types. The fine root biomass of the four vegetation types was mainly distributed in the 0-10 cm soil layer and decreased with soil depth. The proportion of fine root biomass (44.1%) in the 0-10 cm soil layer of abandoned grassland was significantly higher than those in other three vegetation types. The fine root productions of four vegetation types were in the order of abandoned grassland (315 g · m(-2) · a(-1)) > deciduous shrubs (249 g · m(-2) a(-1)) > R. pseudoacacia plantation (219 g · m(-2) · a(-1)) > A. desertorum community (115 g · m(-2) · a(-1)), and mainly concentrated in the 0-10 cm top soil layer and decreased with the soil depth. The proportion of the annual production (40.4%) in the 0-10 cm soil layer was the highest in abandoned grassland. Fine roots of abandoned grassland turned over faster than those from the other three vegetation types.

Biomass and its allocation in relation to temperature, precipitation, and soil nutrients in Inner Mongolia grasslands, China.

AIM: Understanding and predicting ecosystem functioning such as biomass accumulation requires an accurate assessment of large-scale patterns of biomass distribution and partitioning in relation to climatic and soil environments. METHODS: We sampled above- and belowground biomass from 26 sites spanning 1500 km in Inner Mongolian grasslands, compared the difference in aboveground, belowground biomass and below-aboveground biomass ratio (AGB, BGB, and B/A, respectively) among meadow steppe, typical steppe, and desert steppe types. The relationships between AGB, BGB, B/A and climatic and soil environments were then examined. RESULTS: We found that AGB and BGB differed significantly among three types of grasslands while B/A did not differ. Structural equation model analyses indicated that mean annual precipitation was the strongest positive driver for AGB and BGB. AGB was also positively associated with soil organic carbon, whereas B/A was positively associated with total soil nitrogen. CONCLUSIONS: These results indicated that precipitation positively influence plant production in Inner Mongolian grasslands. Contrary to the prediction from the optimal partitioning hypothesis, biomass allocation to belowground increased with soil total nitrogen, suggesting that more productive sites may increase belowground allocation as an adaptive strategy to potentially high fire frequencies.

Water supply changes N and P conservation in a perennial grass Leymus chinensis.

Changes in precipitation can influence soil water and nutrient availability, and thus affect plant nutrient conservation strategies. Better understanding of how nutrient conservation changes with variations in water availability is crucial for predicting the potential influence of global climate change on plant nutrient-use strategy. Here, green-leaf nitrogen (N) and phosphorus (P) concentrations, N- and P-resorption proficiency (the terminal N and P concentration in senescent leaves, NRP and PRP, respectively), and N- and P-resorption efficiency (the proportional N and P withdrawn from senescent leaves prior to abscission, NRE and PRE, respectively) of Leymus chinensis (Trin.) Tzvel., a typical perennial grass species in northern China, were examined along a water supply gradient to explore how plant nutrient conservation responds to water change. Increasing water supply at low levels (< 9000 mL/year) increased NRP, PRP and PRE, but decreased green-leaf N concentration. It did not significantly affect green-leaf P concentration or NRE. By contrast, all N and P conservation indicators were not significantly influenced at high water supply levels (> 9000 mL/year). These results indicated that changes in water availability at low levels could affect leaf-level nutrient characteristics, especially for the species in semiarid ecosystems. Therefore, global changes in precipitation may pose effects on plant nutrient economy, and thus on nutrient cycling in the plant-soil systems.

Plant nitrogen dynamics and nitrogen-use strategies under altered nitrogen seasonality and competition.

BACKGROUND AND AIMS: Numerous studies have examined the effects of climatic factors on the distribution of C(3) and C(4) grasses in various regions throughout the world, but the role of seasonal fluctuations in temperature, precipitation and soil N availability in regulating growth and competition of these two functional types is still not well understood. This report is about the effects of seasonality of soil N availability and competition on plant N dynamics and N-use strategies of one C(3) (Leymus chinensis) and one C(4) (Chloris virgata) grass species. METHODS: Leymus chinensis and C. virgata, two grass species native to the temperate steppe in northern China, were planted in a monoculture and a mixture under three different N seasonal availabilities: an average model (AM) with N evenly distributed over the growing season; a one-peak model (OM) with more N in summer than in spring and autumn; and a two-peak model (TM) with more N in spring and autumn than in summer. KEY RESULTS: The results showed that the altered N seasonality changed plant N concentration, with the highest value of L. chinensis under the OM treatment and C. virgata under the TM treatment, respectively. N seasonality also affected plant N content, N productivity and N-resorption efficiency and proficiency in both the C(3) and C(4) species. Interspecific competition influenced N-use and resorption efficiency in both the C(3) and C(4) species, with higher N-use and resorption efficiency in the mixture than in monoculture. The C(4) grass had higher N-use efficiency than the C(3) grass due to its higher N productivity, irrespective of the N treatment or competition. CONCLUSIONS: The observations suggest that N-use strategies in the C(3) and C(4) species used in the study were closely related to seasonal dynamics of N supply and competition. N seasonality might be involved in the growth and temporal niche separation between C(3) and C(4) species observed in the natural ecosystems.

Nitrogen response efficiency increased monotonically with decreasing soil resource availability: a case study from a semiarid grassland in northern China.

The concept of nutrient use efficiency is central to understanding ecosystem functioning because it is the step in which plants can influence the return of nutrients to the soil pool and the quality of the litter. Theory suggests that nutrient efficiency increases unimodally with declining soil resources, but this has not been tested empirically for N and water in grassland ecosystems, where plant growth in these ecosystems is generally thought to be limited by soil N and moisture. In this paper, we tested the N uptake and the N use efficiency (NUE) of two Stipa species (S. grandis and S. krylovii) from 20 sites in the Inner Mongolia grassland by measuring the N content of net primary productivity (NPP). NUE is defined as the total net primary production per unit N absorbed. We further distinguished NUE from N response efficiency (NRE; production per unit N available). We found that NPP increased with soil N and water availability. Efficiency of whole-plant N use, uptake, and response increased monotonically with decreasing soil N and water, being higher on infertile (dry) habitats than on fertile (wet) habitats. We further considered NUE as the product of the N productivity (NP the rate of biomass increase per unit N in the plant) and the mean residence time (MRT; the ratio between the average N pool and the annual N uptake or loss). The NP and NUE of S. grandis growing usually in dry and N-poor habitats exceeded those of S. krylovii abundant in wet and N-rich habitats. NUE differed among sites, and was often affected by the evolutionary trade-off between NP and MRT, where plants and communities had adapted in a way to maximize either NP or MRT, but not both concurrently. Soil N availability and moisture influenced the community-level N uptake efficiency and ultimately the NRE, though the response to N was dependent on the plant community examined. These results show that soil N and water had exerted a great impact on the N efficiency in Stipa species. The intraspecific differences in N efficiency within both Stipa species along soil resource availability gradient may explain the differences in plant productivity on various soils, which will be conducive to our general understanding of the N cycling and vegetation dynamics in northern Chinese grasslands.

Photosynthetic responses of C3 and C4 species to seasonal water variability and competition.

This study examined the impacts of seasonal water variability and interspecific competition on the photosynthetic characteristics of a C3 (Leymus chinensis) and a C4 (Chloris virgata) grass species. Plants received the same amount of water but in three seasonal patterns, i.e. the one-peak model (more water in the summer than in the spring and autumn), the two-peak model (more water in the spring and autumn than in the summer), and the average model (water evenly distributed over the growing season). The effects of water variability on the photosynthetic characteristics of the C3 and C4 species were dependent on season. There were significant differences in the photosynthetic characteristics of the C4 species in the summer and the C3 species in the autumn among the three water treatments. Interspecific competition exerted negative impacts on the C3 species in August and September but had no effects on the C4 species in any of the four measuring dates. The relative competitive capability of the two species was not altered by water availability. The assimilation rate, the maximum quantum yield of net CO2 assimilation, and the maximum rate of carboxylation of the C3 species were 13-56%, 5-11%, and 11-48% greater, respectively, in a monoculture than in a mixture in August and September. The results demonstrated that the photosynthetic characteristics of the C3 and C4 species were affected by water availability, but the effects varied considerably with season.