Effects of N and P enrichment on plant photosynthetic traits in alpine steppe of the Qinghai-Tibetan Plateau.

BACKGROUND: N (nitrogen) and P (phosphorus) play important roles in plant growth and fitness, and both are the most important limiting factors that affect grassland structure and function. However, we still know little about plant physiological responses to N and P enrichment in alpine grassland of the Qinghai-Tibetan Plateau. In our experiment, five dominant common herbaceous species were selected and their photosynthetic parameters, leaf N content, and aboveground biomass were measured. RESULTS: We found that species-specific responses to N and P enrichment were obvious at individual level. N addition (72 kg Nha-1 yr-1), P addition (36 kg Pha-1 yr-1) and NP addition (72 kg Nha-1 yr-1and 36 kg P ha-1 yr-1, simultaneously) significantly promoted net photosynthetic rate of Leymus secalinus. Differential responses also existed in the same functional groups. Responses of forb species to the nutrients addition varied, Aconitum carmichaeli was more sensitive to nutrients addition including N addition (72 kg Nha-1 yr-1), P addition (36 kg Pha-1 yr-1) and NP addition (72 kg Nha-1 yr-1and 36 kg P ha-1 yr-1). Responses of plant community photosynthetic traits were not so sensitive as those of plant individuals under N and P enrichment. CONCLUSIONS: Our findings highlighted that photosynthetic responses of alpine plants to N and P enrichment were species-specific. Grass species Leymus secalinus had a higher competitive advantage compared with other species under nutrient enrichment. Additionally, soil pH variation and nutrients imbalance induced by N and P enrichment is the main cause that affect photosynthetic traits of plant in alpine steppe of the Qinghai-Tibetan Plateau.

Short-term warming and N deposition alter the photosynthetic pigments trade-off in leaves of Leymus secalinus growing in different alpine grassland habitats on Qinghai-Tibetan plateau.

Warming and N (nitrogen) deposition are the two main driving factors of global change. We examined the effects of increased N deposition (8 kg ha-1 year-1) and warming, as well as their combined effect on the leaf photosynthetic pigments of Leymus secalinus, which is one of the key alpine plants growing in different grassland habitats on Qinghai-Tibetan plateau. In 2014, the experiments were established in 12 plots (2×5m) of three types of habitats including alpine meadow (AM), alpine steppe (AS), and cultivated grassland (CG) with the following treatments: CK (control treatment), N (only N deposition), W (only warming), and W&N (warming combined with N deposition). Results showed that the effects of warming and N deposition on photosynthetic pigments of Leymus secalinus varied with different grassland habitat types. In three grassland types, warming led to no significant effects on the total chlorophyll content of L. secalinus, while N deposition alone only significantly enhanced total chlorophyll content in alpine meadow and cultivated grassland. N deposition combined with warming only significantly enhanced total chlorophyll content of L. secalinus in alpine steppe and cultivated grassland. Chla content plays an important role in determining the variation of total chlorophyll content. Chla/Chlb ratio of L. secalinus was more stable in alpine meadow compared with that of L. secalinus in the other two grassland types. Car/Chl ratio of L. secalinus was not prone to be affected by warming and N deposition in all grassland types. Leaf N content was obviously positively correlated with photosynthetic pigments, especially Chla content. Warming and N deposition all affected photosynthetic pigment dynamics and tended to increase Chla by enhancing its weight. Our results highlighted that both warming and N deposition as well as their combination can alter the trade-off of photosynthetic pigments through enhancing the Chla ratio in L. secalinus. In addition, growing habitats should be within consideration when studying alpine plants adaptation mechanism to global change in the future.

Nitrogen Deposition Shifts Grassland Communities Through Directly Increasing Dominance of Graminoids: A 3-Year Case Study From the Qinghai-Tibetan Plateau.

Nitrogen (N) deposition has been increasing for decades and has profoundly influenced the structure and function of grassland ecosystems in many regions of the world. However, the impact of N deposition on alpine grasslands is less well documented. We conducted a 3-year field experiment to determine the effects of N deposition on plant species richness, composition, and community productivity in an alpine meadow of the Qinghai-Tibetan Plateau of China. We found that 3 years of N deposition had a profound effect on these plant community parameters. Increasing N rates increased the dominance of graminoids and reduced the presence of non-graminoids. Species richness was inversely associated with aboveground biomass. The shift in plant species and functional group composition was largely responsible for the increase in productivity associated with N deposition. Climatic factors also interacted with N addition to influence productivity. Our findings suggest that short-term N deposition could increase the productivity of alpine meadows through shifts in composition toward a graminoid-dominated community. Longer-term studies are needed to determine if shifts in composition and increased productivity will be maintained. Future work must also evaluate whether decreasing plant diversity will impair the long-term stability and function of sensitive alpine grasslands.

N Addition Overwhelmed the Effects of P Addition on the Soil C, N, and P Cycling Genes in Alpine Meadow of the Qinghai-Tibetan Plateau.

Although human activities have greatly increased nitrogen (N) and phosphorus (P) inputs to the alpine grassland ecosystems, how soil microbial functional genes involved in nutrient cycling respond to N and P input remains unknown. Based on a fertilization experiment established in an alpine meadow of the Qinghai-Tibetan Plateau, we investigated the response of the abundance of soil carbon (C), N, and P cycling genes to N and P addition and evaluated soil and plant factors related to the observed effects. Our results indicated that the abundance of C, N, and P cycling genes were hardly affected by N addition, while P addition significantly increased most of them, suggesting that the availability of P plays a more important role for soil microorganisms than N in this alpine meadow ecosystem. Meanwhile, when N and P were added together, the abundance of C, N, and P cycling genes did not change significantly, indicating that the promoting effects of P addition on microbial functional genes abundances were overwhelmed by N addition. The Mantel analysis and the variation partitioning analysis revealed the major role of shoot P concentration in regulating the abundance of C, N, and P cycling genes. These results suggest that soil P availability and plant traits are key in governing C, N, and P cycling genes at the functional gene level in the alpine grassland ecosystem.

Impacts of Short-Term Grazing Intensity on the Plant Diversity and Ecosystem Function of Alpine Steppe on the Qinghai-Tibetan Plateau.

Livestock grazing is the primary land use of grasslands worldwide. Grazing has been asserted to alter grassland ecosystem functions, such as productivity, nutrient cycling, and biodiversity conservation. However, few studies have focused on the impact of grazing intensity on the ecosystem multifunctionality (EMF) of alpine grasslands. We conducted a field experiment of manipulating sheep grazing intensity effects on alpine steppe by surveying plant community characteristics and ecosystem functions. Our results showed that plant community composition was altered with increasing grazing intensity, and the dominant species shifted from grasses and sedges to forbs. EMF was the highest under no grazing (CK) and the lowest under heavy grazing (HG), but there was insignificant difference between CK and HG. HG significantly decreased some indicators that reflected nutrient cycling functions, such as soil available nitrogen, plant leaf nitrogen (PN) and phosphorus content (PP). Furthermore, plant diversity had strong correlations with SOC, total nitrogen (TN), and PN. The results could provide scientific bases for biodiversity conservation and sustainable grazing management of alpine steppe.

Effects of 5-Year Nitrogen Addition on Species Composition and Diversity of an Alpine Steppe Plant Community on Qinghai-Tibetan Plateau.

The N deposition rate is notably increased in China, especially in the Qinghai-Tibetan Plateau (QTP). How plants respond to the projected N deposition on the alpine steppe is still in debate. In this study, to investigate the effects of N deposition on the plant community of the alpine steppe, we simulated N deposition at six different N addition rate levels (0, 8, 24, 40, 56, 72 kg N ha-1 y-1) from 2015 to 2019. Species composition and diversity were investigated as the assessment indices. The results showed that the importance value of grasses significantly increased with the increase of the N addition rate, while that of forbs significantly decreased. A high N addition rate (72 kg N ha-1 y-1) induced species composition change, making Leymus secalinus become the most dominant species within the entire plant community. Compared with the control (without N addition), species richness, Shannon-Weiner diversity, Simpson dominance and Pielou Evenness were significantly reduced under a high N addition rate. The changes of plant diversity in the alpine steppe were closely correlated with dynamics of soil nutrients, especially total carbon (TC), total phosphorus (TP) and ammonia nitrogen (NH4-N). Our findings suggested that a high N deposition rate (72 kg N ha-1 y-1) could significantly change plant composition and reduce the diversity of the alpine steppe, though they were less affected by low N deposition rates at present. With the increase of the N deposition rate, plant composition and diversity of the alpine steppe may be negatively affected in the future. In addition, Leymus secalinus is more competitive than other species with an N deposition rate increase. Soil C, soil P and soil NH4-N variation induced by N deposition might play a key role in regulating changes in plant composition and diversity in the alpine steppe. In addition, longer term field investigation needs to be carried out to testify to this phenomenon with the increase of N deposition in the future.

Eco-physiological processes are more sensitive to simulated N deposition in leguminous forbs than non-leguminous forbs in an alpine meadow of the Qinghai-Tibetan Plateau.

Increased nitrogen (N) deposition can affect ecosystem processes and thus influence plant eco-physiological processes in grasslands. However, how N deposition affects eco-physiological processes of leguminous and non-leguminous forbs in alpine grasslands is understudied. A long-term field experiment using a range of simulated N deposition rates (0, 8, 24, 40, 56, and 72 kg N ha-1 year-1) was established to examine the effects of N deposition on various eco-physiological parameters in leguminous and non-leguminous forbs in an alpine meadow of the Qinghai-Tibetan Plateau. We found that the responses of leguminous and non-leguminous forbs to simulated N deposition varied. Net photosynthetic rate of leguminous and non-leguminous forbs exhibited different response patterns, but chronic increases in simulated N deposition rates may lead to negative effects in both functional groups. Neither functional group responded differently in aboveground biomass under the highest N addition level (72 kg N ha-1 year-1) compared to the control. Differences in aboveground biomass of leguminous forbs were observed at intermediate N levels. Short-term simulated N deposition significantly promoted N uptake of both functional groups. In leguminous forbs, simulated N deposition affected net photosynthetic rates (PN) and aboveground biomass (AGB) mainly via stomatal conductance (gs), water use efficiency (WUE), and plant N uptake. In non-leguminous forbs, simulated N deposition affected PN and AGB mainly through WUE and plant N uptake. Our findings suggest that leguminous and non-leguminous forbs have differential response mechanisms to N deposition, and compared with non-leguminous forbs, leguminous forbs are more sensitive to continuing increased N deposition. The obvious decline trend in photosynthetic capacity in leguminous forbs is likely to exacerbate the already divergent ecological processes between leguminous and non-leguminous forbs. More importantly, these changes are likely to alter the future composition, function, and stability of alpine meadow ecosystems.

Effects of simulated N deposition on photosynthesis and productivity of key plants from different functional groups of alpine meadow on Qinghai-Tibetan plateau.

Nitrogen (N) deposition may alter physiological process of plants in grassland ecosystem. However, little is known about the response mechanism of individual plants in alpine regions to N deposition. We conducted a field experiment, and three treatments including 0 kg Nha-1year-1 (CK), 8 kgNha-1year-1 (Low N), and 72 kg N ha-1 year-1 (High N) were established to simulate N deposition in alpine meadow of Qinghai-Tibetan plateau. Our objectives were to determine the influence of N deposition on photosynthesis of different functional types of herbage species in alpine meadow, and finally characterize the links of plant productivity and photosynthesis with soil nutrients. The results showed that responses of alpine plants were species-specific under N deposition. Compared with grass species Agropyron cristatum and forb species Thalictrum aquilegifolium, the sedge species Carex melanantha was much more sensitive to N deposition; a lower N load (8 kgNha-1year-1) can cause a negative effect on its photosynthesis and productivity. Additionally, N deposition can promote plant N uptake and significantly decreased the C (carbon)/N (nitrogen) ratio. Compared with CK and low N deposition, high N deposition inhibited the photosynthesis and growth of the forb species Thalictrum aquilegifolium and sedge species Carex melanantha. In all three functional types of herbage species, the grass species A. cristatum tended to show a much higher photosynthetic capacity and better growth potential; thus, suggesting that grass species A. cristatum will be a more adaptative alpine plants under N deposition. Our findings suggested that plant photosynthetic responses to N deposition were species-specific, low N deposition was not beneficial for all the herbage species, and N deposition may change plant composition by the differential photosynthetic responses among species in alpine grassland. Plant composition shift to grass-dorminant in alpine regions might be attributed to a much higher photosynthetic potential and N use efficiency of grass species.

Effects of Warming and N Deposition on the Physiological Performances of Leymus secalinus in Alpine Meadow of Qinghai-Tibetan Plateau.

Warming and Nitrogen (N) deposition are key global changes that may affect eco-physiological process of territorial plants. In this paper, we examined the effects of warming, N deposition, and their combination effect on the physiological performances of Leymus secalinus. Four treatments were established in an alpine meadow of Qinghai-Tibetan plateau: control (CK), warming (W), N deposition (N), and warming plus N deposition (NW). Warming significantly decreased the photosynthetic rate (Anet ), stomatal conductance (gs ), intercellular CO2 concentration (Ci ), and transpiration rate (Tr ), while N deposition and warming plus N deposition significantly increased those parameters of L. secalinus. Warming significantly increased the VPD and Ls , while N deposition and warming plus N deposition had a significant positive effect. Warming negatively reduced the leaf N content, Chla, Chlb, and total Chl content, while N deposition significantly promoted these traits. Warming, N deposition, and their combination significantly increased the activity of SOD, POD, and CAT. Besides, warming and warming plus N deposition significantly increased the MDA content, while N deposition significantly decreased the MDA content. N deposition and warming plus N deposition significantly increased the Rubisco activity, while warming showed no significant effect on Rubisco activity. N deposition and warming plus N deposition significantly increased the Fv/Fm, ΦPSII, qP, and decreased NPQ, while warming significantly decreased the Fv/Fm, ΦPSII, qP, and increased NPQ. N deposition strengthened the relations between gs , Chl, Chla, Chlb, Rubisco activity, and Anet . Under warming, only gs showed a significantly positive relation with Anet . Our findings suggested that warming could impair the photosynthetic potential of L. secalinus enhanced by N deposition. Additionally, the combination of warming and N deposition still tend to lead positive effects on L. secalinus.