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Phosphorus (P) limitation is a widespread problem of primary production in dryland submitted to persistent nitrogen (N) deposition. The legume alfalfa (Medicago sativa L.), which can fix N

The B7 family, and their receptors, the CD28 family, are major immune checkpoints that regulate T-cell activation and function. In the present study, we explore the role of two B7 immune-checkpoints: HERV-H LTR-Associating Protein 2 (HHLA2) and B7 Family Member, H4 (B7x), in the progression of gastrointestinal and pancreatic neuroendocrine tumors (GINETs and PNETs). We demonstrated that both HHLA2 and B7x were expressed to a high degree in human GINETs and PNETs. We determined that the expression of B7x and HHLA2 correlates with higher grade and higher incidence of nodal and distant spread. Furthermore, we confirmed that HIF-1α overexpression is associated with the upregulation of B7x both in our in vivo (animal model) and in vitro (cell culture) models. When grown in vitro, islet tumor β-cells lack B7x expression, unless cultured under hypoxic conditions, which results in both hypoxia-inducible factor 1 subunit alpha (HIF-1α) and B7x upregulation. In vivo, we demonstrated that Men1/B7x double knockout (KO) mice (with loss of B7x expression) exhibited decreased islet β-cell proliferation and tumor transformation accompanied by increased T-cell infiltration compared with Men1 single knockout mice. We have also shown that systemic administration of a B7x mAb to our Men1 KO mice with PNETs promotes an antitumor response mediated by increased T-cell infiltration. These findings suggest that B7x may be a critical mediator of tumor immunity in the tumor microenvironment of NETs. Therefore, targeting B7x offers an attractive strategy for the immunotherapy of patients suffering from NETs.

Anthropogenic activities, such as mining, influence soil bacterial community composition and microbial distributions. In the current study, the patterns in microbial distribution and the environmental drivers shaping the soil bacterial community composition in the alpine mining area of the Tianshan Mountain region, China, were investigated, and the bacterial communities were analyzed using 16S rDNA pyrosequencing. The environmental factors and their relationships with the microbial community composition, structure, and diversity were also assessed. The soil organic carbon (SOC) concentration increased along the elevation gradient, with the highest concentration in the mining area, which increased microbial abundance and species richness. Some metals, like Ca, Cu, Pb, and Zn, accumulated significantly in the tailing area and were negatively correlated with the microbial community structure. Proteobacteria, Acidobacteria, Actinobacteria, and Verrucomicrobia were the dominant phyla; these dominant phyla were more abundant in the areas without mining than in the areas with mining at the same altitude. The relative abundance of Proteobacteria and Verrucomicrobia significantly increased along the elevation gradient, while that of Actinobacteria in the mining camp area was more than twice those in the other areas due to higher soil pH. Soil biomass was the highest in the valley. Collectively, these results elucidate the influence of anthropogenic mining activities on soil microbial communities in alpine mining soils and provide a basis for the future management of heavy metal-contaminated areas using the indigenous dominant bacterial phyla.

Knowledge of the effects of pasture degradation on soil organic carbon (SOC) and nitrogen (N) fractions in permafrost soils on the Tibetan Plateau is limited. The aims of this study were (1) to evaluate the changes in SOC and N contents in density fractions under Kobresia pasture due to degradation and (2) to explore the contributions of the changes of SOC and N in density fractions to the changes of SOC and N in whole soil. The impact of Kobresia pasture degradation on SOC and N fractions was investigated in the permafrost region of the Tibetan Plateau. A continuously degraded pasture was identified and classified into three categories of vegetation cover according to their degrees of degradation (i.e., vegetation cover decline from 90% ± 6.6% to 70% ± 8.3% and 45% ± 8.7%). The SOC and N fractions were separated by using the density separation method. The Kobresia pasture degradation significantly decreases SOC and N contents and stocks in soils. The SOC and N contents in the whole soil were positively correlated with the SOC and N contents in the light and heavy fractions (p 0.05). When pasture degraded from vegetation covers 90% to 45%, SOC stock at 0–40-cm soil layer decreased by 28.7% and N stock decreased by 39.2% in the whole soil; 56.6% and 47.6%, respectively, in the light fractions and 14.3% and 40.6%, respectively, in the heavy fractions. The depletion rates of N were higher than those of SOC in the heavy fractions and whole soil. At all sites, more than 80% of the SOC and N stocks were protected in heavy fractions. These results indicate that a decoupling depletion of SOC and soil N appeared with the Kobresia pasture degradation in the permafrost region of the Tibetan Plateau. The Kobresia pasture degradation affects the SOC and N fractions differently and thus regulates soil carbon and N cycling in the permafrost soils on the Tibetan Plateau.

Phosphorus (P) bioavailability is an important factor in alpine meadows and plays an important role in the response to climate change and the maintenance of ecosystem functioning. However, little is known about how environmental factors, such as elevation and slope aspect, affect soil P bioavailability. We explored the effects of elevational gradient and slope aspect on different forms of P and P availability in the alpine meadows on the southern slope of the Tian Shan Mountain range. Total P was found to be 851.9–1556.7 mg·kg−1 at different elevational gradients and 437.5–1547.0 mg·kg−1 at different slope aspects, and highest at 3337 and 3652 m.a.s.l., but little differences between slope aspects. Olsen P and Labile P linearly increased with the elevational gradient. The valley and the base of the shady slope had higher contents of H2O-Po, NaHCO3-Pi, and NaHCO3-Po, and high-active organic P (NaHCO3-Po, NaOH-Po, and H2O-Po) was positively correlated with soil total carbon (TC), total nitrogen (TN), soil organic carbon (SOC), and aboveground biomass (AGB), but was negatively correlated with pH, aluminum (Al), and calcium (Ca) at different elevational gradients. High-active bioavailable P (H2O-Pi, H2O-Po, NaHCO3-Pi, and NaHCO3-Po) was positively correlated with soil SOC and AGB and was negatively correlated with pH at different slope aspects. Our results suggest that soil P availability in alpine meadows is significantly controlled by topographical factors and the valleys and base of shady slopes are reservoirs of high-active bioavailable P.

The soil hydrological process plays an important role in transporting and distributing water in soils. However, the correlation between water flow behaviors, soil physical properties, infiltration patterns of soil exhibiting different soil layers, and the variation patterns in water flow behavior in rubber-based agroforestry systems remain poorly understood. Here, we mainly used dye-tracer infiltration to classify water flow behavior and HYDRUS-3D to interpret variation patterns of water flow behaviors in a rubber-based agroforestry system in southwest China. The results showed that (1) the soil of forestry patches under Hevea brasiliensis (HB) and Citrus reticulata (CR) exhibited better soil physical properties than those with no vegetation (NV). Compared with these in the NV, the saturated soil water content and field capacity increased by 15% and 21% in CR, and by 2% and 12% in HB, respectively; while the saturated hydraulic conductivity enhanced by 584% (p HB > NV. Finally, (3) the infiltration flow in soil under HB and CR exhibited a faster downward wetting front and an obvious lateral flow. These infiltration patterns compensated for the insufficiency of soil water in NV and finally caused water redistribution to be more homogenized across the whole soil profile. Therefore, our findings indicated that the rubber-based agroforestry system is a successful approach for conserving water and reducing runoff in tropical regions.

Alpine meadow degradation is a major environmental problem in the Qinghai-Tibetan Plateau of China. While it is expanding at a rapid pace, quantitative information on the effect of grassland degradation on soil properties remains largely unavailable. This study aimed to evaluate the effect of vegetation cover reduction on soil organic carbon (SOC) and nutrient stocks. A degraded alpine meadow with vegetation cover gradients of 90% ± 6.6% to 70% ± 8.3% and 45% ± 8.7% was selected, and soil samples were collected at two soil depths (0–20 and 20–40 cm depths). The reduction in vegetation cover decreased soil sand, SOC, total nitrogen (N), available phosphorus (P), inorganic N, microbial biomass carbon and N, soil moisture, and the ratio of SOC to available P, but increased soil pH, bulk density, the ratio of SOC to total N, and clay and silt contents. The decline in vegetation cover from 90% to 45% significantly affected SOC, total N, total P, and available P stocks at 0–40 cm soil depths (p 9% at the COV90 site and was close to 5% at the COV45 site for 0–20 cm soil depth. These results confirmed that vegetation cover reduction is a key factor that influences the soil nutrient stocks and structural stability of the alpine meadow. The losses in SOC and nutrients by vegetation cover reduction on the alpine meadow can have a remarkable influence on a large scale. Meanwhile, the alpine meadow may have huge carbon and N sequestration potentials duo to the potential increase of vegetation productivity in this area.

Hydro-meteorological variations are fundamental environmental factors affecting site-level ecosystem carbon balance variations; however, how hydro-meteorological variations influence continuous CO2 exchange over alpine meadows during the growing season in permafrost regions on the Tibetan Plateau is poorly understood. Net ecosystem CO2 exchange (NEE) measured using the eddy covariance method was partitioned into its components of ecosystem respiration (ER) and gross ecosystem CO2 exchange (GPP). ER and GPP had a significant relationship with air temperature, vapor pressure deficit, and net radiation, during the growing season of an alpine meadow. There was a positive correlation between GPP and active layer soil temperatures at 5–40 cm except soil temperature at 80 cm, located in the vicinity of the bottom of the active layer. The microbial activity of underground rhizospheric microorganism, mainly distributed in in shallow soil layers at 0–20 cm in alpine permafrost regions on the Tibetan Plateau, can significantly promote plant photosynthesis. Thus, the correlation between GPP and soil temperatures at 5–20 cm was stronger than that between GPP and air temperature. The correlation between ER (GPP) and active layer soil moisture at 5–80 cm was complicated, especially for soils at 5–20 cm. With the increase of active layer soil moisture, ER (GPP) increased first and then decreased. This was related to the thawing process of the active layer, increase of rainfalls, the frozen layer acting as a weak permeable layer, and the replenishment of super-permafrost water. Our results suggested that long-term observations on how hydro-meteorological variations affect the continuous CO2 exchange of alpine meadows with different active layer thicknesses should be carried out in alpine permafrost regions.

Land use patterns and vegetation coverage in semi-arid areas of the Loess Plateau have undergone great changes due to the implementation of the “Grain for Green” project. The introduction of legume pasture species, such as alfalfa (Medicago sativa L.) and sweet clover (Melilotus officinalis L.), is one of the most efficient methods of vegetation restoration and reconstruction in this region. However, there is a need for an effective assessment of the root system distribution and its interaction with soil after long-term introduction. An experiment involving the introduction of alfalfa and sweet clover on abandoned farmlands was initiated in 2003 to assess the long-term effects. After 17 years, root and soil samples at depths of 0–20 and 20–60 cm were collected to characterize the root biomass, root carbon (C), nitrogen (N), and phosphorus (P), soil microbial biomass carbon (MBC) and nitrogen (MBN), soil organic carbon (SOC), and soil N and P. The results showed that the root biomass density of alfalfa in the 0–20 and 20–60 cm layers (63.72 and 12.27 kg m–3, respectively) were significantly higher than for sweet clover (37.43 and 8.97 kg m–3, respectively) and under natural abandonment (38.92 and 9.73 kg m–3, respectively). The SOC, total nitrogen (TN), total phosphorus (TP), available phosphorus (AP), nitrate-nitrogen (NO3–-N), MBC and MBN in the 0–20 and 20–60 cm layers were higher after alfalfa introduction compared with sweet clover introduction and natural abandonment, although the ammonia-nitrogen (NH4+-N) concentration in the 0–20 cm layer was lower. There were significantly positive correlations between root biomass density and both soil nutrients and microbial biomass, while there was a negative correlation between the soil NH4+-N and root biomass density. These results indicate that alfalfa root growth improved soil organic matter accumulation and nutrient mineralization. The accumulation and mineralization of soil nutrients also guaranteed root and microorganism growth. Therefore, it was concluded that alfalfa introduction will promote soil nutrients immobilization and mineralization and may enable sustainable land use in the semi-arid region of the Loess Plateau, China.

Bald patches (BPs) are known to accelerate and simultaneously mitigate the process of desertification. However, the mechanisms of these two synchronous actions are little studied in high desert and cold systems; and the incidence of BPs on alpine meadows degradation in Qinghai-Tibetan Plateau (QTP) of China is still unavailable. This study first aims to investigate the soil properties and the erodibility of the system BPs-VPs at the Beiluhe basin in QTP. Then, we adopted dye tracer and HYDRUS-2/3D methods to interpret the water infiltration patterns from point scale to slope scale. The results show that the mattic epipedon layer on the top soil (0–20 cm) of VPs directly reduced the impact of raindrops on alpine meadow; and the adhesion of root system in VPs prevented the soil particles from stripping and washing away by runoff. The soil particles in BPs were easily eroded by rainfall, lowering the ground level of BPs relative to the ground level of VPs. The two patches therefore alternated to form an erosion interface where marginal meadow was likely detached by raindrops, and washed away through runoff. The saturated hydraulic conductivity (Ks) of surface soil (0–10 cm) was 124% higher in BPs than the VPs. Thereby, BPs caused a high spatial variation of infiltration and runoff in QTP. Moreover, this difference in Ks between the two patches conducted to a lateral flow from BPs to VPs, and to soil layers with different water contents. These findings highlight that the water flow features can potentially disturb the processes of freezing-thawing, frost heaves, and thaw slump; and accelerate the alpine meadow degradation. Therefore, land cover such as crop and vegetation should be applied over the bare soil surface to prevent the degradation of alpine meadow.

Soil microbial communities regulate soil carbon feedbacks to climate warming through microbial respiration (i.e., metabolic rate). A thorough understanding of the responses of composition, biomass, and metabolic rate of soil microbial community to warming is crucial to predict soil carbon stocks in a future warmer climate. Therefore, we conducted a field manipulative experiment in a semiarid grassland on the Loess Plateau of China to evaluate the responses of the soil microbial community to increased temperature from April 2015 to December 2017. Soil temperature was 2.0°C higher relative to the ambient when open‐top chambers (OTCs) were used. Warming did not affect microbial biomass or the composition of microbial functional groups. However, warming significantly decreased microbial respiration, directly resulting from soil pH decrease driven by the comediation of aboveground biomass increase, inorganic nitrogen increase, and moisture decrease. These findings highlight that the soil microbial community structure of semiarid grasslands resisted the short‐term warming by 2°C, although its metabolic rate declined., This study showed that there were no warming effects on soil microbial community composition and on soil microbial biomass, warming decreased soil microbial metabolic rate, and soil microbial community structure resisted to short‐term warming by 2°C.

Soil physical properties, infiltrability, and water flow behavior are closely associated with root morphology. In this study, dye tracer experiments were conducted in the field to assess the effects of roots with various morphometric features on soil water flow behavior and soil infiltrability. Preferential flow is the dominant type of water flow in the three study sites (Minqin, Dongtai, and Mengla). The presence of roots caused noncapillary porosity and saturated hydraulic conductivity to increase by 26% and 252% in the Minqin, 92% and 93% in the Dongtai, and 234% and 96% in the Mengla, respectively, relative to that in in soils without roots. The various morphometric features of roots affect water flow behavior in soil. The fibrous roots of maize induced water to flow and distribute evenly throughout the plough layer of the study site in Minqin. Furrows and ridges in this site exhibited different soil physical and hydrological properties. Furrows could store higher amounts of water than ridges, thereby increasing the likelihood of water absorption by plants. In Dongtai, ponding and surface runoff occurred when water infiltration was inhibited by the mud layer, which exhibited a high soil bulk density value of 1.43 g cm−3 and a low saturated hydraulic conductivity value of 1.67 × 10−5 cm s−1. These phenomena were not observed in plots the smooth roots of Spartina alterniflora penetrated the mud layer. In this site, preferential flow and lateral flow, which is triggered by the sandy loam layer, are important for water discharge from the beach to the ocean. In Mengla, water flowed evenly on hard soil, which exhibited a high bulk density value of 1.43 g cm−3. The fine roots of rubber could guide water infiltration into deep soil layers (73.54 cm), thereby redistributing water to the root zones of rubber trees. Therefore, our findings indicated that water infiltration behavior, which is crucial for water distribution, is affected by the various morphometric features of roots.

Evaluating the responses of soil respiration and its components to global environmental change is crucial for predicting future terrestrial carbon cycle. However, the effects of warming and nitrogen (N) addition on them remain unclear. A field manipulative experiment was conducted in a semi-arid grassland on the Loess Plateau to evaluate the responses of soil total respiration (Rt), autotrophic respiration (Ra) and heterotrophic respiration (Rh) to warming and N addition from April 2015 to December 2016. Open-top chambers were used to elevate temperature and N was added as NH4NO3 at a rate of 4.42 g N m−2 yr−1. Warming significantly decreased Rt and Rh by 7.4% and 9.5%, respectively, but had no significant effect on Ra. N addition significantly stimulated Ra by 34%, whereas it inhibited Rh by 11% and had no significant effect on Rt. The maximum N-inducing stimulation of Rt and Ra were observed in the month with the most rainfall events in 2015. N addition significantly increased the contribution of Ra to Rt by 10%. Warming decreased the Q10 values of Rt and Rh but had no significant effect on Q10 values of Ra. N addition significantly increased the Q10 values of Rt and Rh, whereas it decreased the Q10 values of Ra. The combination of warming and N addition had a synergistic effect on the cumulant of Rh, whereas it had an antagonistic effect on Ra. No interactive effect between warming and N addition was observed on Rt. Our results emphasized that Ra and Rh responded differently to warming and/or N addition, and the extreme rainfall frequency influenced the responses of Rt and its components to N addition. Our findings suggested that Rt has the potential to resist climate warming and increasing N deposition by differentiating the responses of its inherent components.

Understanding the factors contributing to spatial variation in the functional structure of plant is crucial for revegetation and ecosystem management of the Loess Plateau, China. As one of the measures in the “Grain for Green” project, lucerne ( Medicago sativa L.) has been widely planted in this seriously-eroded region to improve revegetation and conserve soil and water. However, information on how environmental factors influence the long-term development of lucerne-rich vegetation is scarce. This study aimed to identify the main environmental variables controlling the lucerne-rich vegetation through multivariate analyses. Vegetation and soil surveys were performed in 28 fields containing 11-year-old lucerne. Vegetation variables were total aboveground biomass, aboveground biomass of lucerne, total cover, lucerne cover, total abundance, species richness, Shannon–Wiener’s diversity index, Simpson’s predominance index, and evenness index. Topographic variables were slope, slope position, and slope aspect. Soil variables were soil total nitrogen (N) and phosphorus (P), available P, mineral N, organic carbon (SOC), moisture content (SMC), microbial biomass C and N. Analyses verified that vegetation variables, soil variables, and topographic variables were significantly correlated. One of the most important factors that influence lucerne revegetation process was SOC. SMC, total P, slope, and mineral N were also key factors that influenced vegetation variables. The obtained values of total aboveground biomass and cover were still high after 11 years planting. The fields that in the north– and south–facing slopes did not significantly differ in terms of total cover, and total and lucerne aboveground biomass. The results suggest that soil properties (such as SOC, TN, and mineral N, etc.) and topographic variables (i.e., slope) interacted with each other and acted on plants simultaneously in the lucerne-rich vegetation. To revegetation the degraded ecosystems by lucerne planting, we should consider the effects of the changes of soil properties, such as SOC, SMC, and soil P, on vegetation development.

Different rubber-based agroforestry systems are adopted to control and mitigate land degradation. However, the soil physical conditions and soil hydrological processes of different agroforestry systems are still unclear. Thus, in this study, rubber (Hevea brasiliensis) monoculture, rubber and Clerodendranthus spicatus agroforestry system (RCS) and rubber and Amomum villosum agroforestry system (RAV) were developed from a degraded land that had similar backgrounds of terrain and management measures for 50 years. Conventional methods were applied, and dye tracer experiments were conducted to measure the soil physical conditions and determine the water movement in soil. After 5 years' growth, both RCS and RAV could effectively promote the soil physical conditions and optimize soil structure by improving the proportion of the three soil phases. Favourable soil properties, multiple-layered canopies and ground cover in agroforestry systems could promote the formation of three-dimensional hydraulic redistribution in soil profile. The infiltration of rainfall into the soil was enhanced; meanwhile, surface runoff and soil erosion were mitigated, and then more water was transported, redistributed and stored into the different soil layers by the more dominant preferential flow, water exchange and lateral flow in soil profiles. These water supply mechanisms could allow planting intercrops with rubber trees to uptake water from different water sources and coexist in an agroforestry system. Our results highlighted that rubber-based agroforestry systems are a useful management practice to maximize the utilization of land and water resources. Copyright © 2017 John Wiley & Sons, Ltd.

Lucerne (Medicago sativa L.) is widely planted on the semi-arid Loess Plateau in China to control soil erosion, yet the extent to which topography affects the productivity of lucerne still remains poorly understood. This study aimed to evaluate how topographic position influences soil properties and aboveground biomass in lucerne-rich vegetation. A total of 112 quadrats were established in 28 fields covered with 11-year-old lucerne vegetation. In each quadrat, the slope, slope position, slope aspect, altitude, soil properties (soil moisture content, organic carbon, total nitrogen [N] and phosphorus [P], available P and inorganic N), and aboveground biomass were measured. Redundancy and multiple regression analyses were performed to determine the relationships among topographic factors, soil properties and aboveground biomass. A modest proportion of variation in soil variables was explained by topographic variables. The altitude, slope and slope position, rather than the slope aspect, were the key factors that influencing soil variables. Soil organic carbon, total N, inorganic N, the ratio of organic carbon to available P (C/P), and the ratio of total N to available P were positively correlated with altitude, whereas available P was negatively correlated with altitude. The soil moisture content was primarily affected by the slope and slope position. The topographic factors did not directly affect the total and lucerne aboveground biomass. The total and lucerne aboveground biomass were positively correlated to soil moisture content, inorganic N and C/P. Therefore, the aboveground biomass of lucerne-rich vegetation could be indirectly regulated by the slope and slope position through the effect on the soil moisture content and by altitude through the effect on soil inorganic N and C/P. This work highlighted the importance of soil properties such as soil moisture and P dynamics in the revegetation process of lucerne in this semi-arid region. As soil properties can be directly governed by topography, considering topography could enhance the quality of vegetation restoration in the large hilly region of the Loess Plateau.

The accrual of soil organic carbon (SOC) and nitrogen (N) in grassland is an important management option to improve the ecosystem functions of grassland. However, how abiotic (such as grazing exclusion (GE)) and biotic factors influence SOC and N and their different fractions in Tibetan alpine meadows remains unclear. In this study, we evaluated the relative importance of abiotic and biotic factors that drive SOC and N contents in soil density fractions by performing redundancy analysis based on three long-term (10 years) fenced alpine meadows maintained in the permafrost region of the Tibetan Plateau in China. Biotic factors comprise plant aboveground biomass, cover and diversity, whereas abiotic factors include soil properties (i.e. soil moisture, pH, clay, silt, sand, total phosphorus, available phosphorus, microbial biomass carbon and N, available N, C:N ratio and C:P ratio) and GE. Site rather than GE has significant effects on the SOC and N contents. GE caused no increase in the SOC and N contents in the whole soil and fractions. Redundancy analysis showed that 96.7% of the variations in SOC and N fractions can be explained by the selected explanatory variables. Aboveground biomass, cover, soil moisture and clay contents were key factors that affected the SOC and N fractions. The SOC and N fractions were mainly explained by the interaction between vegetation and soil, followed by soil, vegetation and GE. The study highlighted the importance of considering the covariation of vegetation and soil for evaluating the SOC and N dynamics in alpine meadows. The effect of GE (such as 10 years) on the SOC and N contents in alpine meadows can be weak in the permafrost region of the Tibetan Plateau.

Revegetation of degraded arable land with legume pasture has been proposed as an effective approach to improve vegetation conditions and control soil erosion on the Loess Plateau, China. This paper investigated the effects of revegetation on soil carbon pool and its fractions. Three treatments (natural revegetation, Medicago sativa L. introduction and Melilotus suaveolens L. introduction) were applied over 11 years from 2003 to 2013 on the semi-arid Loess Plateau; both aboveground biomass and soil organic carbon (SOC) stocks as well as its fractions (i.e., the light-fraction organic carbon (LFOC) and heavy-fraction organic carbon (HFOC)) were measured. We found that plant aboveground biomass was greater under Melilotus treatment than under Medicago treatment and natural revegetation in the first two years. Afterward, the aboveground biomass was greatest under Medicago treatment. Medicago treatment yielded higher SOC and HFOC stocks than the other two treatments during the experimental period. After revegetation for 10 years, the SOC stocks at the 0–1.5 m soil layer under Medicago treatment were 52.8 and 44.8 Mg ha−1 higher than those under natural revegetation and Melilotus treatments, respectively. The mean rates of SOC and HFOC sequestration in the top 20 cm of the soil layer under Medicago treatment were 0.27 and 0.23 Mg ha−1 year−1 in year 11, which were greater than the other two treatments (0.22, 0.15 and −0.07, −0.03 under the two treatments, respectively). Revegetation through introduction of Medicago to degraded arable land can potentially promote SOC accumulation by enhancing the transformation from LFOC to HFOC on the semi-arid Loess Plateau.

Revegetation facilitated by legume species introduction has been used for soil erosion control on the Loess Plateau, China. However, it is still unclear how vegetation and soil resources develop during this restoration process, especially over the longer term. In this study, we investigated the changes of plant aboveground biomass, vegetation cover, species richness and density of all individuals, and soil total nitrogen, mineral nitrogen, total phosphorus and available phosphorus over 11 years from 2003 to 2013 in three treatments (natural revegetation, Medicago sativa L. introduction and Melilotus suaveolens L. introduction) on the semi-arid Loess Plateau. Medicago significantly increased aboveground biomass and vegetation cover, and soil total nitrogen and mineral nitrogen contents. The Medicago treatment had lower species richness and density of all individuals, lower soil moisture in the deep soil (i.e., 1.4-5m), and lower soil available phosphorus. Melilotus introduction significantly increased aboveground biomass in only the first two years, and it was not an effective approach to improve vegetation biomass and cover, and soil nutrients, especially in later stages of revegetation. Overall, our study suggests that M. sativa can be the preferred plant species for revegetation of degraded ecosystems on the Loess Plateau, although phosphorus fertilizer should be applied for the sustainability of the revegetation.

Grazing exclusion (GE) is widely used in the Tibetan Plateau of China to restore degraded grasslands. However, the effects of GE on vegetation and soil properties in alpine meadows are still not well understood. We compared GE with adjacent continuously grazing alpine meadows at five sites to evaluate the impact of GE (10–14 years) on vegetation and soil properties in the permafrost region of the Tibetan Plateau. We collected soil samples at two soil depths (0–20 and 20–40 cm) from GE and adjacent grazing sites. The results showed that GE and Site had significant effects on aboveground biomass and cover, Site but not GE had significant effects on diversity index. GE significantly increased the aboveground biomass and cover in three of the five sites, particularly where vegetation cover was greatest. At each site, GE had inconsistent effects on plant species diversity (a positive effect at one site, no significant effects at three sites, and a negative effect at the other site). No significant differences were found between GE and adjacent grazing grasslands in soil properties (soil organic carbon [SOC], total nitrogen [N], total phosphorus [P], available P, mineral N, microbial biomass carbon [MBC] and nitrogen [MBN] and pH) for both soil depths at each site. A general linear mixed model confirmed that site and sampling depth, rather than GE, significantly affected soil properties (except for MBC). Both total aboveground biomass and cover were positively correlated with soil total P, MBC and MBN, were negatively correlated with available P and pH, and were not significantly correlated with SOC and total N. The results indicate that long-term GE (10–14 years) in alpine meadow did not significantly impact soil properties, and its impact on plant biomass and vegetation cover depends on the vegetation condition of the site. Our results can provide new insights for alpine meadow management in the permafrost region of the Tibetan Plateau.

The relationship between species richness and productivity has been a central issue in community ecology, and this issue has resulted in much debate in the ecological literature. To evaluate whether species richness is consistently associated with productivity and the underlying mechanisms, a potted experiment with various combinations of three perennial plant species (Elymus nutans, Roegneria nutans and Festuca sinensis) was conducted under three fertilization levels over three years with interannual variation in rainfall in a subalpine meadow on the Tibetan plateau, China. The additive partitioning method was used for measuring net, selection and complementarity effects. The results suggest that species richness, composition and density had significant effects on aboveground biomass, but their effects were much less than abiotic factors (fertility and year). Relative yield total (RYT), net, selection and complementarity effects were not consistently positive or negative across years in the mixed communities. Both the interaction of year and species richness and the interaction of year and composition had significant effects on aboveground biomass, selection and complementarity effects. Fertilization and density had no significant effects on net, selection and complementarity effects. Our results indicate that plant richness is not consistently associated with productivity in these experimental plant communities, considering that selection and complementarity effects can vary with complex environmental conditions, and that these factors influence plant productivity. We suggest that different forms of relationships between species richness and productivity may be exhibited based on biotic or abiotic factors in plant communities.

The Loess Plateau of China has suffered from soil erosion for several decades. As part of the Chinese “Grain for Green” project, legume species have been introduced to restore degraded ecosystems in this region. However, information on how environmental variables influence the recovery of vegetation after legume introduction is scarce. We characterized the composition of plant communities and different environmental variables 11 years after the introduction of the legumes Medicago sativa L. and Melilotus suaveolens L. in abandoned fields of the Loess Plateau. The objectives of this research were to evaluate how environmental factors such as duration of experiment, precipitation, soil moisture, soil nutrition, and topography affect the changes in plant species composition, richness and diversity and to identify the key factors driving plant species succession. Multivariate analyses were used to evaluate the relationships between plant communities and environmental variables. These analyses showed that plant species composition varied through time, with annual species being replaced by perennial herbaceous species gradually. The introduction of Medicago and Melilotus to abandoned fields had different effects on later-successional species and changed the successional trajectory of vegetation in the abandoned fields studied. Time since restoration was the most important factor influencing the composition of vegetation. Slope position, soil moisture content, annual precipitation, and slope/aspect were also key factors driving the composition of the plant community. Our results have implications for studies of secondary succession and the topographic and climatic impacts on vegetation change in restoration ecosystems of the semi-arid Loess Plateau, and emphasize the importance of plant–topography–climate interactions in defining the structure and composition of plant communities.

Responses of soil respiration (Rs) to increasing nitrogen (N) deposition and warming will have far-reaching influences on global carbon (C) cycling. However, the seasonal (growing and non-growing seasons) difference of Rs responses to warming and N deposition has rarely been investigated. We conducted a field manipulative experiment in a semi-arid alfalfa-pasture of northwest China to evaluate the response of Rs to nitrogen addition and warming from March 2014 to March 2016. Open-top chambers were used to elevate temperature and N was enriched at a rate of 4.42g m-2yr-1 with NH4NO3. Results showed that (1) N addition increased Rs by 14% over the two-year period; and (2) warming stimulated Rs by 15% in the non-growing season, while inhibited it by 5% in the growing season, which can be explained by decreased plant coverage and soil water. The main effect of N addition did not change with time, but that of warming changed with time, with the stronger inhibition observed in the dry year. When N addition and warming were combined, an antagonistic effect was observed in the growing season, whereas a synergism was observed in the non-growing season. Overall, warming and N addition did not affect the Q10 values over the two-year period, but these treatments significantly increased the Q10 values in the growing season compared with the control treatment. In comparison, combined warming and nitrogen addition significantly reduced the Q10 values compared with the single factor treatment. These results suggest that the negative indirect effect of warming-induced water stress overrides the positive direct effect of warming on Rs. Our results also imply the necessity of considering the different Rs responses in the growing and non-growing seasons to climate change to accurately evaluate the carbon cycle in the arid and semi-arid regions.

The repair of injured tendons remains a great challenge because of the poor intrinsic healing capacity of tendons. In this study, we examined the spatiotemporal expression pattern of the bone morphogenetic protein 2 (bmp-2) gene in normal and experimentally injured rat patellar tendons. We also investigated the ability of exogenously applied BMP-2 to promote patellar tendon healing. In situ hybridization with bmp-2 and alk-6 (bmp-2 receptor) antisense riboprobes revealed that both genes were normally expressed at low levels in intact rat tendons. However, bmp-2 expression was significantly upregulated in tenocytes found in the wound site at 7 d and later following tendon injury. In addition, it was found that bmp-2 was expressed in cultured patellar tenocytes. Application of exogenous BMP-2 to the tendon wound site significantly enhanced tendon repair. Moreover, in vitro and in vivo studies further demonstrated that BMP-2 enhanced tenocyte proliferation and migration to the wound site. The detectable amount of BMP-2 in normal tendons suggests that a basal level of bmp-2 expression was likely present to maintain the normal functions of the patellar tendon. Injury to the tendon induced increased bmp-2 expression intrinsically, but the expression level was insufficient for proper tendon repair. Our findings indicate that it is possible to significantly improve tendon healing by applying exogenous BMP-2 to the wound site, inferring that this protein could be developed as a potential therapeutic reagent for the treatment of damaged tendons.

Understanding the relationships between vegetative and environmental variables is important for revegetation and ecosystem management on the Loess Plateau, China. Lucerne (Medicago sativa L.) has been widely used in the region to improve revegetation, soil and water conservation and to enhance livestock production. However, there is little information on how environmental factors influence long-term succession in lucerne-rich vegetation. Our objective was to identify the main environmental variables controlling the succession process in lucerne-rich vegetation such that native species are not suppressed after sowing on the Loess Plateau. Vegetation and soil surveys were performed in 31 lucerne fields (three lucerne fields without any management from 2003–2013 and 28 fields containing 11-year-old lucerne with one cutting each year). Time after planting was the most important factor affecting plant species succession. Cutting significantly affected revegetation characteristics, such as aboveground biomass, plant density and diversity. Soil moisture content, soil organic carbon, soil available phosphorus and slope aspect were key environmental factors affecting plant species composition and aboveground biomass, density and diversity. Long-term cutting can cause self-thinning in lucerne, maintain the stability of lucerne production and slow its degradation. For effective management of lucerne fields, phosphate fertilizer should be applied and cutting performed.

Hereditary nonpolyposis colorectal cancer (HNPCC) is one of the most common autosomal dominant inherited diseases. Mutations in the human mismatch repair (MMR) proteins MLH1, MSH2, MSH6, PMS1, and PMS2 have been found to co-segregate with HNPCC. The MLH1 and MSH2 proteins have been demonstrated to interact with PMS1, PMS2, and MSH6 proteins. A previous study reported that missense mutations in specific regions of MLH1 can lead to defects in protein-protein interactions with PMS2. Here we report that three missense alterations previously identified as single nucleotide polymorphisms (SNPs) in PMS2 (P511K, T597S, and M622I) cause defective protein-protein interactions with MLH1, even though the alterations are not in the previously reported interaction domain. These results suggest that an additional domain in PMS2 affects MLH1-PMS2 interaction. This study also demonstrates that SNPs can result in gene alterations that indeed have a functional effect on protein phenotype. Thus, these three SNPs may ultimately represent variants with an increased risk factor for tumorgenesis in HNPCC. This study is one of the first to use a functional assay to appraise the role of SNPs and suggests that traditional definitions of polymorphisms and mutations are in need of reconsideration.