Aridity threshold for alpine soil nitrogen isotope signature and ecosystem nitrogen cycling.

Determination of tipping points in nitrogen (N) isotope (δ15N) natural abundance, especially soil δ15N, with increasing aridity, is critical for estimating N-cycling dynamics and N limitation in terrestrial ecosystems. However, whether there are linear or nonlinear responses of soil δ15N to increases in aridity and if these responses correspond well with soil N cycling remains largely unknown. In this study, we investigated soil δ15N and soil N-cycling characteristics in both topsoil and subsoil layers along a drought gradient across a 3000-km transect of drylands on the Qinghai-Tibetan Plateau. We found that the effect of increasing aridity on soil δ15N values shifted from negative to positive with thresholds at aridity index (AI) = 0.27 and 0.29 for the topsoil and subsoil, respectively, although soil N pools and N transformation rates linearly decreased with increasing aridity in both soil layers. Furthermore, we identified markedly different correlations between soil δ15N and soil N-cycling traits above and below the AI thresholds (0.27 and 0.29 for topsoil and subsoil, respectively). Specifically, in wetter regions, soil δ15N positively correlated with most soil N-cycling traits, suggesting that high soil δ15N may result from the "openness" of soil N cycling. Conversely, in drier regions, soil δ15N showed insignificant relationships with soil N-cycling traits and correlated well with factors, such as soil-available phosphorus and foliage δ15N, demonstrating that pathways other than typical soil N cycling may dominate soil δ15N under drier conditions. Overall, these results highlight that different ecosystem N-cycling processes may drive soil δ15N along the aridity gradient, broadening our understanding of N cycling as indicated by soil δ15N under changing drought regimes. The aridity threshold of soil δ15N should be considered in terrestrial N-cycling models when incorporating 15N isotope signals to predict N cycling and availability under climatic dryness.

Plant height as an indicator for alpine carbon sequestration and ecosystem response to warming.

Growing evidence indicates that plant community structure and traits have changed under climate warming, especially in cold or high-elevation regions. However, the impact of these warming-induced changes on ecosystem carbon sequestration remains unclear. Using a warming experiment on the high-elevation Qinghai-Tibetan Plateau, we found that warming not only increased plant species height but also altered species composition, collectively resulting in a taller plant community associated with increased net ecosystem productivity (NEP). Along a 1,500 km transect on the Plateau, taller plant community promoted NEP and soil carbon through associated chlorophyll content and other photosynthetic traits at the community level. Overall, plant community height as a dominant trait is associated with species composition and regulates ecosystem C sequestration in the high-elevation biome. This trait-based association provides new insights into predicting the direction, magnitude and sensitivity of ecosystem C fluxes in response to climate warming.

Response of C:N:P stoichiometry to long-term drainage of peatlands: Evidence from plant, soil, and enzyme.

Drought induced by climate warming and human activities regulates carbon (C) cycling of peatlands by changing plant community composition and soil properties. Estimating the responses of peatlands C cycling to environmental changes requires further study of C: nitrogen (N): phosphorus (P) stoichiometric ratios of soil, plants, and enzyme activities. However, systematic studies on the stoichiometry of above-ground and below-ground ecosystems of peatlands post drainage remain scarce. This study compared stoichimetric ratios of plant and soil and stoichimetric ratios of enzyme activities with different functions in two different parts of a minerotrophic peatland, a natural undisturbed part and a part that had been drained for almost 50 years, in Northern China. For the shrub plants, the average C:N:P ratios of leaf in natural and drained peatland were 448:17:1 and 393:15:1, respectively. This indicated that the growth rate of shrub plants is higher in the drained peatland than in the natural peatland, which makes P element more concentrated in the photosynthetic site. However, from the perspective of the dominant plant, the mean C:N:P ratio of Carex leaf was 650:25:1 in the natural peatland, but was 1028:50:1 for Dasiphora fruticosa in drained peatland. This indicated that the intensification of P-limitation of plant growth after drainage. Soil C:N:P ratios of above water table depth (AWT) were 238:15:1 and 277:12:1, but were 383:17:1 and 404:19:1 for below water table depth (BWT) in the natural and the drained peatland, respectively. Soil C:P ratios were greater than the threshold elemental ratio of C:P (174:1), but the soil C:N ratios were less than the threshold elemental ratio of C:N (23:1), which suggested that P was the most limiting nutrient of soil. The soil microbial activities were co-limited by C&P in Baijianghe peatlands. However, the microbial metabolic P limitation was intensified, but the C limitation was weakened for the above water table depth soil after long-term drainage. There are connection between plant-microbe P limitation in peatlands. The P limitation of microbial metabolism was significant positively correlated with soil C:N but negatively with soil moisture. The increase in the lignocelluloses index suggested considerable decomposition of soil organic matter after peatland drainage. These results of stoichiometric ratios from above- to below ground could provide scientific base for the C cycling of peatland undergone climate change.

Threshold responses of soil gross nitrogen transformation rates to aridity gradient.

The responses of soil nitrogen (N) transformations to climate change are crucial for biome productivity prediction under global change. However, little is known about the responses of soil gross N transformation rates to drought gradient. Along an aridity gradient across the 2700 km transect of drylands on the Qinghai-Tibetan Plateau, this study measured three main soil gross N transformation rates in both topsoil (0-10 cm) and subsoil (20-30 cm) using the laboratorial 15 N labeling. The relevant soil abiotic and biotic variables were also determined. The results showed that gross N mineralization and nitrification rates steeply decreased with increasing aridity when aridity was less than 0.5 but just slightly decreased with increasing aridity when aridity was larger than 0.5 at both soil layers. In topsoil, the decreases of the two gross rates were accompanied by the similar decreased patterns of soil total N content and microbial biomass carbon with increasing aridity (p < .05). In subsoil, although the decreased pattern of soil total N with increasing aridity was still similar to the decreases of the two gross rates (p < .05), microbial biomass carbon did not change (p > .05). Instead, bacteria and ammonia oxidizing archaea abundances decreased with increasing aridity when aridity was larger than 0.5 (p < .05). With an aridity threshold of 0.6, gross N immobilization rate increased with increasing aridity in wetter region (aridity < 0.6) accompanied with an increased bacteria/fungi ratio, but decreased with increasing aridity in drier region (aridity > 0.6) where mineral N and microbial biomass N also decreased at both soil layers (p < .05). This study provided new insight to understand the differential responses of soil N transformation to drought gradient. The threshold responses of the gross N transformation rates to aridity gradient should be noted in biogeochemical models to better predict N cycling and manage land in the context of global change.

Depth-dependent drivers of soil aggregate carbon across Tibetan alpine grasslands.

Elucidating the effects underlying soil organic carbon (SOC) variation is imperative for ascertaining the potential drivers of mitigating climate change. However, the drivers of variations in various SOC fractions (e.g., macroaggregate C, microaggregate C, and silt and clay C) at different soil depths remain poorly understood. Here, we investigated the effects and relative contributions of climatic, plant, edaphic, and microbial factors on soil aggregate C between the topsoil (0-10 cm) and subsoil (20-30 cm) across alpine grasslands on the Tibetan Plateau. Results showed that the C content of macroaggregates, microaggregates, and silt and clay fractions in the topsoil was 128.6 %, 49.6 %, and 242.4 % higher than that in the subsoil, respectively. Overall, plant properties were the most determinants controlling soil macroaggregate, microaggregate, and silt + clay associated C for both two soil depths, accounting for 32.2 %, 37.4 %, and 38.8 % of the variation, respectively, followed by edaphic, microbial, and climatic factors. The aggregate C of both soil depths was significantly related with the climatic, plant, edaphic, and microbial factors, but the relative importance of these determinants was soil-depth dependent. Specifically, the effects of plant root biomass and microbial (e.g., microbial biomass carbon and fungal diversity index) factors on each aggregate C weakened with soil depth, but the importance of edaphic factors (e.g., clay content, pH, and bulk density) strengthened with soil depth, except for the weakened effect of bulk density on the microaggregate C. And the effects of climatic factor (e.g., mean annual precipitation) on macroaggregate and microaggregate C increased with soil depth. Our results highlight differential drivers and their impacts on soil aggregate C between the topsoil and subsoil, which benefits biogeochemical models for more accurately forecasting soil C dynamics and its feedbacks to environmental changes.

Dryness weakens the positive effects of plant and fungal ß diversities on above- and belowground biomass.

Plant and microbial diversity are key to determine ecosystem functioning. Despite the well-known role of local-scale α diversity in affecting vegetation biomass, the effects of community heterogeneity (ß diversity) of plants and soil microbes on above- and belowground biomass (AGB and BGB) across contrasting environments still remain unclear. Here, we conducted a dryness-gradient transect survey over 3000 km across grasslands on the Tibetan Plateau. We found that plant ß diversity was more dominant than α diversity in maintaining higher levels of AGB, while soil fungal ß diversity was the key driver in enhancing BGB. However, these positive effects of plant and microbial ß diversity on AGB and BGB were strongly weakened by increasing climatic dryness, mainly because higher soil available phosphorus caused by increasing dryness reduced both plant and soil fungal ß diversities. Overall, these new findings highlight the critical role of above- and belowground ß diversity in sustaining grassland biomass, raising our awareness to the ecological risks of large-scale biotic homogenization under future climate change.

Plant Evolution History Overwhelms Current Environment Gradients in Affecting Leaf Chlorophyll Across the Tibetan Plateau.

Aims: Leaf chlorophyll (Chl) is a fundamental component and good proxy for plant photosynthesis. However, we know little about the large-scale patterns of leaf Chl and the relative roles of current environment changes vs. plant evolution in driving leaf Chl variations. Locations: The east to west grassland transect of the Tibetan Plateau. Methods: We performed a grassland transect over 1,600 km across the Tibetan Plateau, measuring leaf Chl among 677 site-species. Results: Leaf Chl showed a significantly spatial pattern across the grasslands in the Tibetan Plateau, decreasing with latitude but increasing with longitude. Along with environmental gradient, leaf Chl decreased with photosynthetically active radiation (PAR), but increased with water availability and soil nitrogen availability. Furthermore, leaf Chl also showed significant differences among functional groups (C4 > C3 species; legumes < non-legume species), but no difference between annual and perennial species. However, we surprisingly found that plant evolution played a dominant role in shaping leaf Chl variations when comparing the sum and individual effects of all the environmental factors above. Moreover, we revealed that leaf Chl non-linearly decreased with plant evolutionary divergence time. This well-matches the non-linearly increasing trend in PAR or decreasing trend in temperature during the geological time-scale uplift of the Tibetan Plateau. Main Conclusion: This study highlights the dominant role of plant evolution in determining leaf Chl variations across the Tibetan Plateau. Given the fundamental role of Chl for photosynthesis, these results provide new insights into reconsidering photosynthesis capacity in alpine plants and the carbon cycle in an evolutionary view.

Environmental factors, bacterial interactions and plant traits jointly regulate epiphytic bacterial community composition of two alpine grassland species.

Epiphytic microbes on the surfaces of leaves and roots can bring substantial benefits or damages to their plant hosts. Although various factors have been proposed for shaping the epiphytic microbial composition, the contributions of environment factors, endogenous microbial taxa interactions, host plant traits, and their interactive effects are poorly understood. Here, we conducted a field investigation along a precipitation gradient and collected leaf and root surface microbes of two alpine plant species for 16S rRNA sequencing. We found that epiphytic bacterial community composition significantly changed along the precipitation gradient through ordination analyses and permutational multivariate analysis of variance. Beneficial bacterial taxa from Caulobacteraceae, Sphingomonadaceae, Comamonadaceae and Rhizobiales were enriched in the high precipitation zones. The stress-tolerant Hymenobacteraceae, Micrococcaceae, and Geodermatophilaceae occurred more frequently in the phyllosphere, and the Thermoleophilia, Thermomicrobiales and Bacillales were enriched in the rhizosphere at the drier sites. Mean annual precipitation was the most important factor regulating the epiphytic bacterial community composition. The direct effect of climate on bacterial community composition was higher in the phyllosphere than in the rhizosphere where joint effects of climate, plant traits and soil properties predominated. Distinct leaf trichome cover and plant height clearly explained the host effect on the phyllosphere bacterial community composition while belowground traits did not explain the host effect well on the rhizosphere bacterial community composition. We detected a significant role of bacterial taxa interactions in shaping microbial communities, where greater negative taxa interactions led to lesser composition changes. Structural equation modeling showed that environmental factors and bacterial interactions substantially contributed to the variation in epiphytic community composition, followed by host plant traits. This study advances our understanding of complex factors affecting alpine epiphytic community assembly and further confirms the role of biotic interactions.

Increased annual methane uptake driven by warmer winters in an alpine meadow.

Pronounced nongrowing season warming and changes in soil freeze-thaw (F-T) cycles can dramatically alter net methane (CH4 ) exchange rates between soils and the atmosphere. However, the magnitudes and drivers of warming impacts on CH4 uptake in different stages of the F-T cycle are poorly understood in cold alpine ecosystems, which have been found to be a net sink of atmospheric CH4 . Here, we reported a year-round ecosystem daily CH4 uptake in an alpine meadow on the Qinghai-Tibetan Plateau after a 5-year warming experiment that included a control, a low-level warming treatment (+2.4℃ at 5 cm soil depth), and a high-level warming treatment (+4.5℃ at 5 cm soil depth). We found that warming shortened the F-T cycle under the low-level warming and soils did not freeze under the high-level warming. Although both warming treatments increased the mean CH4 uptake rate, only the high-level warming significantly increased annual CH4 uptake compared to the control. The warming-induced stimulation of CH4 uptake mainly occurred in the cold season, which was mostly during spring thaw under low-level warming and during the frozen winter under high-level warming due to a longer period with thawed soil. We also found that warming significantly stimulated daily CH4 uptake mainly by reducing near-surface soil water content in the warm season, whereas both soil water content and temperature controlled daily CH4 uptake in different ways during the autumn freeze, frozen winter, and spring thaw periods of the control. Our study revealed a strong warming effect on CH4 uptake during the entire F-T cycle in the alpine meadow, especially the unfrozen winter. Our results also suggested the important roles of soil pH, available phosphorus, and methanotroph abundance in regulating annual CH4 uptake in response to warming, which should be incorporated into biogeochemical models for accurately forecasting CH4  fluxes under future climate scenarios.

[Effects of Balance Acupuncture Combined with Motor Relearning for Lower Limb Motor Function of Stroke Patients with Hemiplegia].

OBJECTIVE: To investigate the effect of balance acupuncture combined with motor relearning training on lower limb motor function in stroke patients with hemiplegia. METHODS: Eighty stroke patients were randomly assigned to motor relearning training group and balance acupuncture plus motor relearning group (n＝40 cases in each). The motor relearning training program consisting of upper-limb functional training, lying supine, bedside sitting, sitting-balancing, standing up and down, standing-balancing, walking, orofacial functional training, etc. was given to patients of the two groups. Balance acupuncture stimulating was applied to Piantan-， Jiantong-， Xitong-， Tuntong- and Huaitong-points for 30 min, once daily, 5 times a week for 8 weeks. The lower limb motor performance ability was assessed by using Fugl-Meyer Assessment (FMA-L) scale, the balance function assessed by using Berg balance scale (BBS), the motor ability evaluated by using Rivermead motor index (RMI), and the gait (walking speed, cadence, and step length) evaluated by using Brunnstrom hemiplegia gait evaluation scale. RESULTS: After the treatment, the scores of FMA-L, BBS, RMI and hemiplegia gait were significantly increased in the two groups compared with their own pre-treatment (P<0.05), and were significantly higher in the balance acupuncture plus motor relearning group than in the motor relearning training group (P<0.05). CONCLUSION: The balance acupuncture combined with motor relearning can improve lower limb motor function and balance function, and has a better effect than simple motor relearning training.

[Improvement of Upper Limb and Hand Functions of Stroke Patients by Balancing Acupuncture Combined with Motor Relearning Training].

OBJECTIVE: To observe the therapeutic effect of balance acupuncture combined with motor relearning training for upper limb and hand functions of stroke patients. METHODS: Sixty-two stroke patients were randomly divided into balance acupuncture group (n＝31) and routine acupuncture group (n＝31). For patients of the balance acupuncture group, Piantan, Jiantong and Wantong points on the healthy side were used. When Jiantong point taken, the acupuncture needle was removed after the patient experienced an electric shock-like spreading needling sensation. When Wantong point employed, the needle was removed after the patient experienced a local, intensified or spreading needling sensation. When Piantan point used, the needle was retained after the patient experienced an electric shock-like needling sensation, then, the motor relearning training was conducted, and the needle was removed immediately after the training. For patients of the routine acupuncture group, Jianyu(LI 15), Jianzhen (SI 9), Quchi (LI 11), etc. were needled with the needles retained for 30 min after getting needling sensations. The motor relearning training was also carried out after removal of the needle. The treatment in both groups was performed once daily, 6 days a week, and lasted for 8 weeks. The Fugl-Meyer score and motor function scale (MAS) of the upper limb, and the fine performance score and motor function score of the hand were assessed before and after the treatment. RESULTS: Following treatment, the Fugl-Meyer score and MAS of the upper limbs, and the motor function score and fine performance score of the hand were significantly increased in both groups compared with pre-treatment in each group (P<0.05 ), suggesting a functional improvement of both upper limb and hand. The therapeutic effect of the balance acupuncture was obviously superior to that of routine acupuncture in improving functions of both the upper limb and hand (P<0.05)．. CONCLUSION: Balance acupuncture combined with motor relearning training is helpful to improve the comprehensive function of the upper limb and hand in stroke patients.

A high plant density reduces the ability of maize to use soil nitrogen.

Understanding the physiological changes associated with high grain yield and high N use efficiency (NUE) is important when increasing the plant density and N rate to develop optimal agronomic management. We tested the hypothesis that high plant densities resulting in crowding stress reduce the ability of plants to use the N supply post-silking, thus decreasing the grain yield and NUE. In 2013 and 2014, a field experiment, with five N-application rates and three plant densities (6.0, 7.5, and 9.0 plants m-2), was conducted in the North China Plain (NCP). The calculated maximum grain yield and agronomic use efficiency (AEN) at a density of 7.5 plants m-2 were 12.4 Mg ha-1 and 39.3 kg kg-1, respectively, which were significantly higher than the values obtained at densities of 6.0 (11.3 Mg ha-1 and 30.2 kg kg-1) and 9.0 plant m-2 (11.7 Mg ha-1 and 27.8 kg kg-1). A high plant density of 9.0 plants m-2 decreased the post-silking N accumulation, leaf N concentration and net photosynthesis, which reduced the post-silking dry matter production, resulting in a low yield and NUE. Although a relatively low grain yield was observed at a density of 9.0 plants m-2, the optimal N rate increased from 150 to 186 kg N ha-1 at a density of 7.5 plants m-2. These results indicate that high plant densities with crowding stress reduce the ability of plants to use soil N during the post-silking period, and high rate of N fertilizer was needed to increase grain yield. We conclude that selecting the appropriate plant density combined with optimal N management could increase grain yields and the NUE in the NCP.

Designing a new cropping system for high productivity and sustainable water usage under climate change.

The food supply is being increasingly challenged by climate change and water scarcity. However, incremental changes in traditional cropping systems have achieved only limited success in meeting these multiple challenges. In this study, we applied a systematic approach, using model simulation and data from two groups of field studies conducted in the North China Plain, to develop a new cropping system that improves yield and uses water in a sustainable manner. Due to significant warming, we identified a double-maize (M-M; Zea mays L.) cropping system that replaced the traditional winter wheat (Triticum aestivum L.) -summer maize system. The M-M system improved yield by 14-31% compared with the conventionally managed wheat-maize system, and achieved similar yield compared with the incrementally adapted wheat-maize system with the optimized cultivars, planting dates, planting density and water management. More importantly, water usage was lower in the M-M system than in the wheat-maize system, and the rate of water usage was sustainable (net groundwater usage was ≤150 mm yr-1). Our study indicated that systematic assessment of adaptation and cropping system scale have great potential to address the multiple food supply challenges under changing climatic conditions.