Nutrient additions in pristine Patagonian Sphagnum bog vegetation: can phosphorus addition alleviate (the effects of) increased nitrogen loads

Sphagnum-bog ecosystems have a limited capability to retain carbon and nutrients when subjected to increased nitrogen (N) deposition. Although it has been proposed that phosphorus (P) can dilute negative effects of nitrogen by increasing biomass production of Sphagnum mosses, it is still unclear whether P-addition can alleviate physiological N-stress in Sphagnum plants. A 3-year fertilisation experiment was conducted in lawns of a pristine Sphagnum magellanicum bog in Patagonia, where competing vascular plants were practically absent. Background wet deposition of nitrogen was low (≈ 0.1-0.2 g · N · m(-2) · year(-1)). Nitrogen (4 g · N · m(-2) · year(-1)) and phosphorus (1 g · P · m(-2) · year(-1)) were applied, separately and in combination, six times during the growing season. P-addition substantially increased biomass production of Sphagnum. Nitrogen and phosphorus changed the morphology of Sphagnum mosses by enhancing height increment, but lowering moss stem density. In contrast to expectations, phosphorus failed to alleviate physiological stress imposed by excess nitrogen (e.g. amino acid accumulation, N-saturation and decline in photosynthetic rates). We conclude that despite improving growth conditions by P-addition, Sphagnum-bog ecosystems remain highly susceptible to nitrogen additions. Increased susceptibility to desiccation by nutrients may even worsen the negative effects of excess nitrogen especially in windy climates like in Patagonia.     

Ecophysiological adjustment of two Sphagnum species in response to anthropogenic nitrogen deposition

Here, it was investigated whether Sphagnum species have adjusted their nitrogen (N) uptake in response to the anthropogenic N deposition that has drastically altered N-limited ecosystems, including peatlands, worldwide. A lawn species, Sphagnum balticum, and a hummock species, Sphagnum fuscum, were collected from three peatlands along a gradient of N deposition (2, 8 and 12 kg N ha(-1) yr(-1)). The mosses were subjected to solutions containing a mixture of four N forms. In each solution one of these N forms was labeled with (15)N (namely (15)NH(+)(4), (15)NO(-)(3) and the amino acids [(15)N]alanine (Ala) and [(15)N]glutamic acid (Glu)). It was found that for both species most of the N taken up was from , followed by Ala, Glu, and very small amounts from NO(-)(3). At the highest N deposition site N uptake was reduced, but this did not prevent N accumulation as free amino acids in the Sphagnum tissues. The reduced N uptake may have been genetically selected for under the relatively short period with elevated N exposure from anthropogenic sources, or may have been the result of plasticity in the Sphagnum physiological response. The negligible Sphagnum NO(-)(3) uptake may make any NO(-)(3) deposited readily available to co-occurring vascular plants.     

松嫩草地80种草本植物叶片氮磷化学计量特征

以松嫩草地常见草本植物为研究对象, 分析了各生活型和功能群叶片氮磷化学计量特征。结果显示: 松嫩草地80种草本植物的叶片氮、磷质量浓度分别为(24.2 ± 0.96) mg·g ^-1和(2.0 ± 0.10) mg·g ^-1, 面积浓度分别为(13.0 ± 0.54) mg·cm ^-2和(1.0 ± 0.05) mg·cm ^-2, 氮磷比为13.0 ± 0.39, 氮磷比与叶片磷质量浓度、叶片氮、磷面积浓度有显著相关关系; 松嫩草地植物生长受到氮限制。一年生植物叶片氮、磷质量浓度和变异系数高于其他生活型, 各生活型之间氮面积浓度和氮磷比差异不显著。豆科植物叶片氮的质量浓度、面积浓度和氮磷比高于其他功能群。在不同生活型或功能群之间, 植物叶片磷的面积浓度差异不显著, 都在1.0 mg·cm ^-2左右; 适当地增加群落中豆科植物的比例, 可能有助于提高松嫩草地产量和质量。     

Nitrogen concentration and δ15N signature of ombrotrophic Sphagnum mosses at different N deposition levels in Europe

Alteration of the global nitrogen (N) cycle because of human‐enhanced N fixation is a major concern particularly for those ecosystems that are nutrient poor by nature. Because Sphagnum‐dominated mires are exclusively fed by wet and dry atmospheric deposition, they are assumed to be very sensitive to increased atmospheric N input. We assessed the consequences of increased atmospheric N deposition on total N concentration, N retention ability, and δ¹⁵N isotopic signature of Sphagnum plants collected in 16 ombrotrophic mires across 11 European countries. The mires spanned a gradient of atmospheric N deposition from about 0.1 up to about 2 g m^?2 yr^?1. Mean N concentration in Sphagnum capitula was about 6 mg g^?1 in less polluted mires and about 13 mg g^?1 in highly N‐polluted mires. The relative difference in N concentration between capitulum and stem decreased with increasing atmospheric N deposition, suggesting a possible metabolic mechanism that reduces excessive N accumulation in the capitulum. Sphagnum plants showed lower rates of N absorption under increasing atmospheric N deposition, indicating N saturation in Sphagnum tissues. The latter probably is related to a shift from N‐limited conditions to limitation by other nutrients. The capacity of the Sphagnum layer to filter atmospheric N deposition decreased exponentially along the depositional gradient resulting in enrichment of the mire pore water with inorganic N forms (i.e., NO₃^?+NH₄⁺). Sphagnum plants had δ¹⁵N signatures ranging from about ?8‰ to about ?3‰. The isotopic signatures were rather related to the ratio of reduced to oxidized N forms in atmospheric deposition than to total amount of atmospheric N deposition, indicating that δ¹⁵N signature of Sphagnum plants can be used as an integrated measure of δ¹⁵N signature of atmospheric precipitation. Indeed, mires located in areas characterized by greater emissions of NH₃ (i.e., mainly affected by agricultural activities) had Sphagnum plants with a lower δ¹⁵N signature compared with mires located in areas dominated by NO_x emissions (i.e., mainly affected by industrial activities).     

氮沉降下西南山地针叶林根际和非根际土壤微生物养分限制特征差异

长期氮(N)沉降诱导了土壤养分失衡, 深刻影响着森林生态系统养分循环过程、生态功能及其可持续发展。前期研究发现N沉降下西南森林树木生长受到不同程度的磷(P)限制, 而土壤微生物是否表现出与植物养分限制特征协同的响应仍未明确。基于此, 该研究以西南山地典型人工针叶林——华山松(Pinus armandii)林为对象, 通过野外原位模拟N沉降实验, 测定了土壤有效养分供给、土壤微生物生物量(碳(C)、N、P)含量以及胞外酶活性, 结合生态酶化学计量的3种模型(比值模型、矢量分析模型与阈值元素比率模型)验证森林根际/非根际土壤中微生物是否受P养分限制。结果表明: (1) N添加下两个土壤位置(根际和非根际土壤)酸性磷酸酶(AP)活性分别显著升高52.5%和53.2%, 导致土壤酶活性N:P分别降低7.8%和4.8%; (2)矢量模型分析发现N添加下两个土壤位置的矢量角度均大于45°, 根际土壤和非根际土壤的矢量角度分别为52.2°和49.0°; (3) N添加下两个土壤位置C:P阈值(TER_C:P)显著降低, 导致C:P阈值与土壤有效C:P的比值(TER_C:P/Av_C:P)远小于1, 且根际土壤表现更明显。综上所述, 3个模型均表明N沉降加剧了土壤微生物代谢的P限制, 且根际土壤微生物P限制程度更强, 这与土壤和微生物养分含量及其化学计量特征密切相关。该研究结果可为全球气候变化下森林生态系统的适应性管理提供重要科学依据。     

氮添加对昆仑山高山草地土壤、微生物和植物生态化学计量特征的影响

全球范围内的氮沉降增加改变了生态系统氮(N)素循环过程,由此带来的生态学效应已成为当前研究的热点。以昆仑山高山草地生态系统2种优势植物黄花棘豆(Oxytropis ochrocephala)和针茅(Stipa capillata)为研究对象,开展人工氮肥添加试验,研究土壤-微生物-植物系统各组分生态化学计量特征对氮添加的响应特征。结果表明:①氮添加显著提高了土壤NH4^+-N和土壤NO3^--N含量(P<0.05),土壤全N、全磷(P)、速效P含量没有明显变化。②氮添加条件下针茅叶片N含量增加,P含量降低,而黄花棘豆N和P含量无明显变化。③土壤微生物量碳(MBC)和微生物量氮(MBN)随着施氮量的增加呈现出先增加后降低的趋势,当施氮量为6N·m^-2·y^-1时呈现出最高值。土壤NH4+-N含量与土壤微生物量N含量有显著的正相关关系(P<0.01)。综合分析表明,短期氮添加有利于土壤养分和微生物量的积累,促进植物和微生物养分吸收利用。2种优势植物的生态化学计量特征对氮沉降的响应不同,过量的氮输入将会造成植物生长受到P限制,氮沉降会改变昆仑山高山草地生态系统的生物地球化学循环过程。     

Linkages of plant stoichiometry to ecosystem production and carbon fluxes with increasing nitrogen inputs in an alpine steppe

Unprecedented levels of nitrogen (N) have entered terrestrial ecosystems over the past century, which substantially influences the carbon (C) exchange between the atmosphere and biosphere. Temperature and moisture are generally regarded as the major controllers over the N effects on ecosystem C uptake and release. N‐phosphorous (P) stoichiometry regulates the growth and metabolisms of plants and soil organisms, thereby affecting many ecosystem C processes. However, it remains unclear how the N‐induced shift in the plant N:P ratio affects ecosystem production and C fluxes and its relative importance. We conducted a field manipulative experiment with eight N addition levels in a Tibetan alpine steppe and assessed the influences of N on aboveground net primary production (ANPP), gross ecosystem productivity (GEP), ecosystem respiration (ER), and net ecosystem exchange (NEE); we used linear mixed‐effects models to further determine the relative contributions of various factors to the N‐induced changes in these parameters. Our results showed that the ANPP, GEP, ER, and NEE all exhibited nonlinear responses to increasing N additions. Further analysis demonstrated that the plant N:P ratio played a dominate role in shaping these C exchange processes. There was a positive relationship between the N‐induced changes in ANPP (ΔANPP) and the plant N:P ratio (ΔN:P), whereas the ΔGEP, ΔER, and ΔNEE exhibited quadratic correlations with the ΔN:P. In contrast, soil temperature and moisture were only secondary predictors for the changes in ecosystem production and C fluxes along the N addition gradient. These findings highlight the importance of plant N:P ratio in regulating ecosystem C exchange, which is crucial for improving our understanding of C cycles under the scenarios of global N enrichment.     

Nutrient concentration ratios and co-limitation in South African grasslands

*Assessing plant nutrient limitation is a fundamental part of understanding grassland dynamics. The ratio of concentrations of nitrogen (N) and phosphorus (P) in vegetation has been proposed as an index of the relative limitation of biomass production by N and P, but its utility has not been tested well in grasslands. *At five sites in Kruger National Park, South Africa, across soil and precipitation contrasts, N and P were added in a factorial design to grass-dominated plots. *Although the N:P ratio of unfertilized vegetation across all sites (5.8) would have indicated that production was N-limited, aboveground production was consistently co-limited by N and P. Aboveground production was still greater in plots fertilized with N and P than in those fertilized with just N, but the N:P ratio did not exceed standard thresholds for P limitation in N-fertilized vegetation. Comparisons among sites showed little pattern between site N:P ratio and relative responses to N and P. *When combined with results from other grassland fertilization studies, these data suggest that the N:P ratio of grasses has little ability to predict limitation in upland grasslands. Co-limitation between N and P appears to be much more widespread than would be predicted from simple assumptions of vegetative N:P ratios.     

硅肥对水稻-田面水-土壤氮磷含量的影响

研究硅肥对双季稻产量及土壤氮磷流失的影响,旨在为典型双季稻区施肥结构优化以及农田面源污染综合防控技术提供技术支撑.采用田间试验研究在相同氮磷肥基础上施0、750、1500、2250和3000 kg·hm^-2 (T₀、T₁、T₂、T₃、T₄)硅肥对双季稻产量和氮磷吸收、田面水氮磷含量及土壤有效硅、有机质、碱解氮和有效磷含量的影响.结果表明:与不施硅肥处理(T₀)比较,早、晚稻季水稻稻谷分别增产2.2%～30.4%和3.9%～9.2%;早、晚稻籽粒氮素积累量增加2.4%～47.3%、磷素积累量增加2.2%～41.3%,秸秆氮素积累量增加0.4%～28.3%、磷素积累量增加5.1%～31.0%;施硅肥后第1天,施硅处理田面水总氮(TN)浓度较不施硅肥处理平均降低3.4%～28.8%、铵态氮(NH₄⁺-N)降低10.4%～25.6%、总磷(TP)降低25.5%～29.2%、可溶性总磷(TDP)降低30.8%～38.0%;第45天后施硅各处理田面水总磷和可溶性总磷含量呈现上升趋势,并显著高于T₀处理.施硅肥有利于提高土壤有效硅水平以及有机质和碱解氮含量,以T₁处理效果最好,土壤速效磷含量随硅肥用量增加呈降低趋势.     

Leaf nitrogen and phosphorus stoichiometry across 753 terrestrial plant species in China

Leaf nitrogen and phosphorus stoichiometry of Chinese terrestrial plants was studied based on a national data set including 753 species across the country. Geometric means were calculated for functional groups based on life form, phylogeny and photosynthetic pathway, as well as for all 753 species. The relationships between leaf N and P stoichiometric traits and latitude (and temperature) were analysed. The geometric means of leaf N, P, and N : P ratio for the 753 species were 18.6 and 1.21 mg g(-1) and 14.4, respectively. With increasing latitude (decreasing mean annual temperature, MAT), leaf N and P increased, but the N : P ratio did not show significant changes. Although patterns of leaf N, P and N : P ratios across the functional groups were generally consistent with those reported previously, the overall N : P ratio of China's flora was considerably higher than the global averages, probably caused by a greater shortage of soil P in China than elsewhere. The relationships between leaf N, P and N : P ratio and latitude (and MAT) also suggested the existence of broad biogeographical patterns of these leaf traits in Chinese flora.     

Mycorrhizas improve nitrogen nutrition of Trifolium repens after 8 yr of selection under elevated atmospheric CO2 partial pressure

Altered environmental conditions may change populations of arbuscular mycorrhizal fungi and thereby affect mycorrhizal functioning. We investigated whether 8 yr of free-air CO2 enrichment has selected fungi that differently influence the nutrition and growth of host plants. In a controlled pot experiment, two sets of seven randomly picked single spore isolates, originating from field plots of elevated (60 Pa) or ambient CO2 partial pressure (pCO2), were inoculated on nodulated Trifolium repens (white clover) plants. Fungal isolates belonged to the Glomus claroideum or Glomus intraradices species complex, and host plants were clonal micropropagates derived from nine genets. Total nitrogen (N) concentration was increased in leaves of plants inoculated with fungal isolates from elevated-pCO2 plots. These isolates took up nearly twice as much N from the soil as isolates from ambient-pCO2 plots and showed much greater stimulation of biological N2 fixation. The morpho-species identity of isolates had a more pronounced effect on N2 fixation and on root length colonized than isolate identity. We conclude that rising atmospheric pCO2 may select for fungal strains that will help their host plants to meet increased N demands.     

Symbiotic nitrogen fixation in a tropical rainforest: 15N natural abundance measurements supported by experimental isotopic enrichment

* Leguminous trees are very common in the tropical rainforests of Guyana. Here, species-specific differences in N(2) fixation capability among nodulating legumes growing on different soils and a possible limitation of N(2) fixation by a relatively high nitrogen (N) and low phosphorus (P) availability in the forest were investigated. * Leaves of 17 nodulating species and 17 non-nodulating reference trees were sampled and their delta(15)N values measured. Estimates of N(2) fixation rates were calculated using the (15)N natural abundance method. Pot experiments were conducted on the effect of N and P availability on N(2) fixation using the (15)N-enriched isotope dilution method. * Nine species showed estimates of > 33% leaf N derived from N(2) fixation, while the others had low or undetectable N(2) fixation rates. High N and low P availability reduced N(2) fixation substantially. * The results suggest that a high N and low P availability in the forest limit N(2) fixation. At the forest ecosystem level, N(2) fixation was estimated at c. 6% of total N uptake by the tree community. We conclude that symbiotic N(2) fixation plays an important role in maintaining high amounts of soil available N in undisturbed forest.     

Centelloside accumulation in leaves of Centella asiatica is determined by resource partitioning between primary and secondary metabolism while influenced by supply levels of either nitrogen,phosphorus or potassium

In the present study we aimed to investigate the relevance of either N, P or K supply for herb and leaf yield and for centelloside concentrations in Centella asiatica L. Urban leaves. In this regard, we elucidated the causal relationship between assimilation rate, leaf N, P and K concentrations, herb and leaf production, and centelloside accumulation. The experiments were conducted consecutively in a greenhouse where C. asiatica was grown in hydroponic culture and fertigated with nutrient solutions at either 0, 30, 60, 100 or 150% of the N, P or K amount in a standard Hoagland solution. In general, the increase in N, P or K supply enhanced assimilation rate and herb and leaf yield. However, exceeding specific thresholds, the high availability of one single nutrient caused lower leaf N concentrations and a decline in assimilation rate and plant growth. Irrespective of N, P and K supply, the leaf centelloside concentrations were negatively associated with herb and leaf yield, which is in accordance with the assumptions of the carbon/nutrient balance and the growth differentiation balance hypotheses. Moreover, we found strong negative correlations between saponins and leaf N concentrations, while the respective sapogenins were negatively correlated with K concentrations. Using C. asiatica as model system, our experiments reveal for the first time that the accumulation of saponins and sapogenins is affected by resource allocation between primary and secondary metabolism and that besides carbon, also nutrient availability is relevant for the regulation of the centelloside synthesis. Finally, our results highlight the huge potential of optimized and carefully controlled mineral nutrition of medicinal plants for steering the bio-production of high-quality natural products.     

14-3-3 proteins fine-tune plant nutrient metabolism

14-3-3 Proteins regulate many cellular processes by binding to phosphorylated proteins. Previous findings suggest a connection between three 14-3-3 isoforms and plant nutrient signaling. To better understand how these 14-3-3s regulate metabolism in response to changes in plant nutrient status, putative new targets involved in nitrogen (N) and sulfur (S) metabolisms have been identified. The interactions between these 14-3-3s and multiple proteins involved in N and S metabolism and altered activity of the target proteins were confirmed in planta. Using a combination of methods, this work elucidates how 14-3-3s function as modulators of plant N and S metabolic pathways.     

Carbon dioxide diffusion across stomata and mesophyll and photo-biochemical processes as affected by growth CO2 and phosphorus nutrition in cotton

Nutrients such as phosphorus may exert a major control over plant response to rising atmospheric carbon dioxide concentration (CO₂), which is projected to double by the end of the 21st century. Elevated CO₂ may overcome the diffusional limitations to photosynthesis posed by stomata and mesophyll and alter the photo-biochemical limitations resulting from phosphorus deficiency. To evaluate these ideas, cotton (Gossypium hirsutum) was grown in controlled environment growth chambers with three levels of phosphate (Pi) supply (0.2, 0.05 and 0.01 mM) and two levels of CO₂ concentration (ambient 400 and elevated 800 μmol mol⁻¹) under optimum temperature and irrigation. Phosphate deficiency drastically inhibited photosynthetic characteristics and decreased cotton growth for both CO₂ treatments. Under Pi stress, an apparent limitation to the photosynthetic potential was evident by CO₂ diffusion through stomata and mesophyll, impairment of photosystem functioning and inhibition of biochemical process including the carboxylation efficiency of ribulose-1,5-bisphosphate carboxylase/oxyganase and the rate of ribulose-1,5-bisphosphate regeneration. The diffusional limitation posed by mesophyll was up to 58% greater than the limitation due to stomatal conductance (g_s) under Pi stress. As expected, elevated CO₂ reduced these diffusional limitations to photosynthesis across Pi levels; however, it failed to reduce the photo-biochemical limitations to photosynthesis in phosphorus deficient plants. Acclimation/down regulation of photosynthetic capacity was evident under elevated CO₂ across Pi treatments. Despite a decrease in phosphorus, nitrogen and chlorophyll concentrations in leaf tissue and reduced stomatal conductance at elevated CO₂, the rate of photosynthesis per unit leaf area when measured at the growth CO₂ concentration tended to be higher for all except the lowest Pi treatment. Nevertheless, plant biomass increased at elevated CO₂ across Pi nutrition with taller plants, increased leaf number and larger leaf area.