Effects of climate warming on carbon fluxes in grasslands— A global meta‐analysis

Climate warming will affect terrestrial ecosystems in many ways, and warming‐induced changes in terrestrial carbon (C) cycling could accelerate or slow future warming. So far, warming experiments have shown a wide range of C flux responses, across and within biome types. However, past meta‐analyses of C flux responses have lacked sufficient sample size to discern relative responses for a given biome type. For instance grasslands contribute greatly to global terrestrial C fluxes, and to date grassland warming experiments provide the opportunity to evaluate concurrent responses of both plant and soil C fluxes. Here, we compiled data from 70 sites (in total 622 observations) to evaluate the response of C fluxes to experimental warming across three grassland types (cold, temperate, and semi‐arid), warming methods, and short (≤3 years) and longer‐term (>3 years) experiment lengths. Overall, our meta‐analysis revealed that experimental warming stimulated C fluxes in grassland ecosystems with regard to both plant production (e.g., net primary productivity (NPP) 15.4%; aboveground NPP (ANPP) by 7.6%, belowground NPP (BNPP) by 11.6%) and soil respiration (Rs) (9.5%). However, the magnitude of C flux stimulation varied significantly across cold, temperate and semi‐arid grasslands, in that responses for most C fluxes were larger in cold than temperate or semi‐arid ecosystems. In semi‐arid and temperate grasslands, ecosystem respiration (Reco) was more sensitive to warming than gross primary productivity (GPP), while the opposite was observed for cold grasslands, where warming produced a net increase in whole‐ecosystem C storage. However, the stimulatory effect of warming on ANPP and Rs observed in short‐term studies (≤3 years) in both cold and temperate grasslands disappeared in longer‐term experiments (>3 years). These results highlight the importance of conducting long‐term warming experiments, and in examining responses across a wide range of climate.     

放牧对草地生态系统影响的研究进展

全球草地占据30%左右的陆地面积, 在全球气候变化、碳氮及养分循环、保持水土、调节畜牧业生产等方面具有重要的作用。目前草地的主要利用方式之一就是放牧, 不同的牲畜种类、放牧强度、年限、历史和制度等, 会影响草地植物群落、生物多样性及土壤微生物, 进而影响草地生态系统结构、功能和过程。该文围绕放牧对草地生态系统结构、功能和过程的影响, 1)回顾了20世纪50年代到现在各个历史阶段放牧对草地生态系统影响的研究; 2)利用文献计量分析的方法, 剖析了放牧对草地影响研究的热点内容、重要区域和关键词等; 3)阐明了放牧对草地植物生长、群落特征、碳氮及养分循环、生产力及土壤质量等的各方面影响的研究进展及国内相关研究的优势及存在的主要问题和不足; 4)基于上述分析, 从草地放牧精准管理、经典假说验证、放牧和全球变化研究相结合等方面, 提出未来研究的前沿方向和优先领域。该文在系统总结放牧对草地生态系统影响的研究进展、研究优势及存在问题的基础上, 提出未来的研究应与全球变化相结合, 为我国的草地放牧生态学研究、适应性管理和可持续利用等提供科学基础。     

Research progress on the effects of grazing on grassland ecosystem

全球草地占据30%左右的陆地面积, 在全球气候变化、碳氮及养分循环、保持水土、调节畜牧业生产等方面具有重要的作用。目前草地的主要利用方式之一就是放牧, 不同的牲畜种类、放牧强度、年限、历史和制度等, 会影响草地植物群落、生物多样性及土壤微生物, 进而影响草地生态系统结构、功能和过程。该文围绕放牧对草地生态系统结构、功能和过程的影响, 1)回顾了20世纪50年代到现在各个历史阶段放牧对草地生态系统影响的研究; 2)利用文献计量分析的方法, 剖析了放牧对草地影响研究的热点内容、重要区域和关键词等; 3)阐明了放牧对草地植物生长、群落特征、碳氮及养分循环、生产力及土壤质量等的各方面影响的研究进展及国内相关研究的优势及存在的主要问题和不足; 4)基于上述分析, 从草地放牧精准管理、经典假说验证、放牧和全球变化研究相结合等方面, 提出未来研究的前沿方向和优先领域。该文在系统总结放牧对草地生态系统影响的研究进展、研究优势及存在问题的基础上, 提出未来的研究应与全球变化相结合, 为我国的草地放牧生态学研究、适应性管理和可持续利用等提供科学基础。     

Responses of terrestrial nitrogen pools and dynamics to different patterns of freeze‐thaw cycle: A meta‐analysis

Altered freeze‐thaw cycle (FTC) patterns due to global climate change may affect nitrogen (N) cycling in terrestrial ecosystems. However, the general responses of soil N pools and fluxes to different FTC patterns are still poorly understood. Here, we compiled data of 1519 observations from 63 studies and conducted a meta‐analysis of the responses of 17 variables involved in terrestrial N pools and fluxes to FTC. Results showed that under FTC treatment, soil NH₄⁺, NO₃^?, NO₃^? leaching, and N₂O emission significantly increased by 18.5%, 18.3%, 66.9%, and 144.9%, respectively; and soil total N (TN) and microbial biomass N (MBN) significantly decreased by 26.2% and 4.7%, respectively; while net N mineralization or nitrification rates did not change. Temperate and cropland ecosystems with relatively high soil nutrient contents were more responsive to FTC than alpine and arctic tundra ecosystems with rapid microbial acclimation. Therefore, altered FTC patterns (such as increased duration of FTC, temperature of freeze, amplitude of freeze, and frequency of FTC) due to global climate warming would enhance the release of inorganic N and the losses of N via leaching and N₂O emissions. Results of this meta‐analysis help better understand the responses of N cycling to FTC and the relationships between FTC patterns and N pools and N fluxes.     

放牧对呼伦贝尔草地植物和土壤生态化学计量学特征的影响

放牧通过畜体采食、践踏和排泄物归还影响草地群落组成、植物形态和土壤养分,植物通过改变养分利用策略适应环境变化。通过分析呼伦贝尔草原放牧和围封样地中的群落植物和土壤的碳氮磷养分及化学计量比,探讨放牧对生态系统化学计量学特征和养分循环速率的影响机制。结果如下:(1)群落尺度上,放牧和围封草地植物叶片C、N和P的含量没有显著差异;但是在种群尺度上,放牧草地植物叶片N含量显著高于围封草地;(2)放牧草地土壤全C、全N、有机C、速效P含量,低于围封草地,硝态N含量高于围封草地;土壤全P和铵态N指标没有显著差异;(3)放牧草地植物C∶N比显著低于围封草地,植物残体分解速率较快,提高了生态系统养分循环速率。     

短期氮素添加和模拟放牧对青藏高原高寒草甸生态系统呼吸的影响

氮沉降和放牧是影响草地碳循环过程的重要环境因子,但很少有研究探讨这些因子交互作用对生态系统呼吸的影响。在西藏高原高寒草甸地区开展了外源氮素添加与刈割模拟放牧实验,测定了其对植物生物量分配、土壤微生物碳氮和生态系统呼吸的影响。结果表明:氮素添加显著促进生态系统呼吸,而模拟放牧对其无显著影响,且降低了氮素添加的刺激作用。氮素添加通过提高微生物氮含量和土壤微生物代谢活性,促进植物地上生产,从而增加生态系统的碳排放;而模拟放牧降低了微生物碳含量,且降低了氮素添加的作用,促进根系的补偿性生长,降低了氮素添加对生态系统碳排放的刺激作用。这表明,放牧压力的存在会抑制氮沉降对高寒草甸生态系统碳排放的促进作用,同时外源氮输入也会缓解放牧压力对高寒草甸生态系统生产的负面影响。     

放牧对全球草地生态系统碳氮磷化学计量特征影响的Meta分析

为明确全球尺度下放牧管理措施对草地生态系统碳(C)、氮(N)、磷(P)化学计量特征的影响,提高草地生态系统管理水平,本研究选取国内外83篇中英文文献进行Meta分析,并通过亚组分析探讨了放牧家畜组合(羊单牧、牛单牧和牛羊混牧)和放牧强度(轻度、中度、重度)对草地生态系统叶片、凋落物、根系,以及土壤C、N、P化学计量特征的影响。结果表明: 放牧会显著降低叶片和凋落物C含量、C/N、C/P,增加N、P含量及N/P;显著降低根系和土壤C、N含量,C/P和N/P,增加P含量和C/N。叶片、凋落物化学计量特征变化对牛、羊单独放牧响应更为明显,而根系、土壤化学计量特征变化则对混牧响应更为明显,重度放牧会对草地生态系统化学计量特征产生更大的影响。放牧会降低土壤N含量,增加P含量,表明放牧对草地N、P含量的影响路径不同。进一步研究N、P含量变化对放牧活动不平衡响应机制,将放牧方式、强度的影响纳入草地生态系统预测、管理模型,能够有效提高草地生态系统管理水平。     

Experimental litterfall manipulation drives large and rapid changes in soil carbon cycling in a wet tropical forest

Global changes such as variations in plant net primary production are likely to drive shifts in leaf litterfall inputs to forest soils, but the effects of such changes on soil carbon (C) cycling and storage remain largely unknown, especially in C‐rich tropical forest ecosystems. We initiated a leaf litterfall manipulation experiment in a tropical rain forest in Costa Rica to test the sensitivity of surface soil C pools and fluxes to different litter inputs. After only 2 years of treatment, doubling litterfall inputs increased surface soil C concentrations by 31%, removing litter from the forest floor drove a 26% reduction over the same time period, and these changes in soil C concentrations were associated with variations in dissolved organic matter fluxes, fine root biomass, microbial biomass, soil moisture, and nutrient fluxes. However, the litter manipulations had only small effects on soil organic C (SOC) chemistry, suggesting that changes in C cycling, nutrient cycling, and microbial processes in response to litter manipulation reflect shifts in the quantity rather than quality of SOC. The manipulation also affected soil CO ₂ fluxes; the relative decline in CO ₂ production was greater in the litter removal plots (?22%) than the increase in the litter addition plots (+15%). Our analysis showed that variations in CO ₂ fluxes were strongly correlated with microbial biomass pools, soil C and nitrogen (N) pools, soil inorganic P fluxes, dissolved organic C fluxes, and fine root biomass. Together, our data suggest that shifts in leaf litter inputs in response to localized human disturbances and global environmental change could have rapid and important consequences for belowground C storage and fluxes in tropical rain forests, and highlight differences between tropical and temperate ecosystems, where belowground C cycling responses to changes in litterfall are generally slower and more subtle.     

Effects of livestock grazing on grassland carbon storage and release override impacts associated with global climate change

Predicting future carbon (C) dynamics in grassland ecosystems requires knowledge of how grazing and global climate change (e.g., warming, elevated CO₂, increased precipitation, drought, and N fertilization) interact to influence C storage and release. Here, we synthesized data from 223 grassland studies to quantify the individual and interactive effects of herbivores and climate change on ecosystem C pools and soil respiration (Rs). Our results showed that grazing overrode global climate change factors in regulating grassland C storage and release (i.e., Rs). Specifically, grazing significantly decreased aboveground plant C pool (APCP), belowground plant C pool (BPCP), soil C pool (SCP), and Rs by 19.1%, 6.4%, 3.1%, and 4.6%, respectively, while overall effects of all global climate change factors increased APCP, BPCP, and Rs by 6.5%, 15.3%, and 3.4% but had no significant effect on SCP. However, the combined effects of grazing with global climate change factors also significantly decreased APCP, SCP, and Rs by 4.0%, 4.7%, and 2.7%, respectively but had no effect on BPCP. Most of the interactions between grazing and global climate change factors on APCP, BPCP, SCP, and Rs were additive instead of synergistic or antagonistic. Our findings highlight the dominant effects of grazing on C storage and Rs when compared with the suite of global climate change factors. Therefore, incorporating the dominant effect of herbivore grazing into Earth System Models is necessary to accurately predict climate–grassland feedbacks in the Anthropocene.     

A meta‐analysis of soil salinization effects on nitrogen pools,cycles and fluxes in coastal ecosystems

Salinity intrusion caused by land subsidence resulting from increasing groundwater abstraction, decreasing river sediment loads and increasing sea level because of climate change has caused widespread soil salinization in coastal ecosystems. Soil salinization may greatly alter nitrogen (N) cycling in coastal ecosystems. However, a comprehensive understanding of the effects of soil salinization on ecosystem N pools, cycling processes and fluxes is not available for coastal ecosystems. Therefore, we compiled data from 551 observations from 21 peer‐reviewed papers and conducted a meta‐analysis of experimental soil salinization effects on 19 variables related to N pools, cycling processes and fluxes in coastal ecosystems. Our results showed that the effects of soil salinization varied across different ecosystem types and salinity levels. Soil salinization increased plant N content (18%), soil NH₄⁺ (12%) and soil total N (210%), although it decreased soil NO₃^? (2%) and soil microbial biomass N (74%). Increasing soil salinity stimulated soil N₂O fluxes as well as hydrological NH₄⁺ and NO₂^? fluxes more than threefold, although it decreased the hydrological dissolved organic nitrogen (DON) flux (59%). Soil salinization also increased the net N mineralization by 70%, although salinization effects were not observed on the net nitrification, denitrification and dissimilatory nitrate reduction to ammonium in this meta‐analysis. Overall, this meta‐analysis improves our understanding of the responses of ecosystem N cycling to soil salinization, identifies knowledge gaps and highlights the urgent need for studies on the effects of soil salinization on coastal agro‐ecosystem and microbial N immobilization. Additional increases in knowledge are critical for designing sustainable adaptation measures to the predicted intrusion of salinity intrusion so that the productivity of coastal agro‐ecosystems can be maintained or improved and the N losses and pollution of the natural environment can be minimized.     

Belowground carbon flux links biogeochemical cycles and resource‐use efficiency at the global scale

Nutrient limitation is pervasive in the terrestrial biosphere, although the relationship between global carbon (C) nitrogen (N) and phosphorus (P) cycles remains uncertain. Using meta‐analysis we show that gross primary production (GPP) partitioning belowground is inversely related to soil‐available N : P, increasing with latitude from tropical to boreal forests. N‐use efficiency is highest in boreal forests, and P‐use efficiency in tropical forests. High C partitioning belowground in boreal forests reflects a 13‐fold greater C cost of N acquisition compared to the tropics. By contrast, the C cost of P acquisition varies only 2‐fold among biomes. This analysis suggests a new hypothesis that the primary limitation on productivity in forested ecosystems transitions from belowground resources at high latitudes to aboveground resources at low latitudes as C‐intensive root‐ and mycorrhizal‐mediated nutrient capture is progressively replaced by rapidly cycling, enzyme‐derived nutrient fluxes when temperatures approach the thermal optimum for biogeochemical transformations.     

Asymmetric effects of grazing intensity on macroelements and microelements in grassland soil and plants in Inner Mongolia Grazing alters nutrient dynamics of grasslands

Grazing is a traditional grassland management technique and greatly alters ecosystem nutrient cycling. The effects of grazing intensity on the nutrient dynamics of soil and plants in grassland ecosystems remain uncertain, especially among microelements. A 2‐year field grazing experiment was conducted in a typical grassland with four grazing intensities (ungrazed control, light, moderate, and heavy grazing) in Inner Mongolia, China. Nutrient concentration was assessed in soil and three dominant plant species (Stipa krylovii, Leymus chinensis, and Cleistogenes squarrosa). Assessed quantities included four macroelements (carbon (C), nitrogen (N), phosphorus (P), and magnesium (Mg)) and four microelements (copper (Cu), iron (Fe), manganese (Mn), and zinc (Zn)). Soil total C, total N, total P, available N, and available P concentrations significantly increased with grazing intensity but soil Mg, Cu, Fe, Mn, Zn concentrations had no significant response. Plant C concentration decreased but plant N, P, Mg, Cu, Fe, Mn, and Zn concentrations significantly increased with grazing intensity. In soil, macroelement dynamics (i.e., C, N, and P) exhibited higher sensitivity with grazing intensity, conversely in plants, microelements were more sensitive. This result indicates macroelements and microelements in soil and plants had asymmetric responses with grazing intensity. The slopes of nutrient linear regression in C. squarrosa were higher than that of S. krylovii and L. chinensis, indicating that C. squarrosa had higher nutrient acquisition capacity and responded more rapidly to heavy grazing. These findings indicate that short‐term heavy grazing accelerates nutrient cycling of the soil–plant system in grassland ecosystems, elucidate the multiple nutrient dynamics of soil and plants with grazing intensity, and emphasize the important function of microelements in plant adaptation in grazing management.     

放牧对草地土壤氮素循环关键过程的影响与机制研究进展

放牧是人类管理利用草地生态系统的最主要途径之一.食草动物的采食、践踏、卧息和排泄物归还等干扰不仅会改变草地地上植物群落,也会对土壤养分循环产生显著的影响.随着人类活动的加剧,放牧强度和频率也在逐渐增加,从而对草地土壤氮素循环关键过程产生重要影响.放牧主要通过改变土壤的物理性质、土壤氮库以及微生物的组成和结构,进而影响氮素在土壤中的迁移与转化.适度放牧会促进土壤氮素的矿化过程和硝化过程,加快氮素的周转,有利于植物吸收可利用氮素,而对于反硝化的影响与草地的水热条件和土壤类型等密切相关.目前,关于放牧强度对土壤氮素循环关键过程影响的研究结果不一致,其影响机制尚不明晰,尤其对于不同类型的草原仍存在较大的差异.本研究在大量查阅国内外已有研究结果的基础上,论述了放牧对土壤氮素循环关键过程的影响效应,总结了放牧对土壤氮素循环的影响机制,指出了目前研究过程中存在的不足,并对未来研究中值得重点关注和深入研究的科学问题进行了探讨与展望,为进一步理解放牧对草地土壤氮素循环的影响提供参考.     

Grazing simplifies soil micro‐food webs and decouples their relationships with ecosystem functions in grasslands

Livestock grazing often alters aboveground and belowground communities of grasslands and their mediated carbon (C) and nitrogen (N) cycling processes at the local scale. Yet, few have examined whether grazing‐induced changes in soil food webs and their ecosystem functions can be extrapolated to a regional scale. We investigated how large herbivore grazing affects soil micro‐food webs (microbes and nematodes) and ecosystem functions (soil C and N mineralization), using paired grazed and ungrazed plots at 10 locations across the Mongolian Plateau. Our results showed that grazing not only affected plant variables (e.g., biomass and C and N concentrations), but also altered soil substrates (e.g., C and N contents) and soil environment (e.g., soil pH and bulk density). Grazing had strong bottom‐up effects on soil micro‐food webs, leading to more pronounced decreases at higher trophic levels (nematodes) than at lower trophic levels (microbes). Structural equation modeling showed that changes in plant biomass and soil environment dominated grazing effects on microbes, while nematodes were mainly influenced by changes in plant biomass and soil C and N contents; the grazing effects, however, differed greatly among functional groups in the soil micro‐food webs. Grazing reduced soil C and N mineralization rates via changes in plant biomass, soil C and N contents, and soil environment across grasslands on the Mongolian Plateau. Spearman's rank correlation analysis also showed that grazing reduced the correlations between functional groups in soil micro‐food webs and then weakened the correlation between soil micro‐food webs and soil C and N mineralization. These results suggest that changes in soil micro‐food webs resulting from livestock grazing are poor predictors of soil C and N processes at regional scale, and that the relationships between soil food webs and ecosystem functions depend on spatial scales and land‐use changes.     

Effects of enhanced nitrogen deposition and phosphorus limitation on nitrogen budgets of semi-natural grasslands

Increased reactive atmospheric N deposition has been implicated in floristic changes in species‐rich acidic and calcareous grasslands, but the fate of this pollutant N in these ecosystems is unknown. This paper reports the first analysis of N budgets and N fluxes for two grasslands in the White Peak area of Derbyshire, one of the most heavily N‐polluted locations in the UK. N fluxes were monitored in lysimeter cores (retaining the original turfs) taken from field plots of unimproved acidic and calcareous grasslands that had received (in addition to ambient N deposition) simulated enhanced N deposition treatments of 3.5 and 14 g N m^?2 yr^?1 for 6 years. The influence of reducing phosphorus limitation was assessed by factorial additions of P. Seasonal leached losses of nitrate, ammonia and organic N were monitored in detail along with estimates of N removal through simulated grazing and gaseous losses through denitrification and volatilization. The rates of N fluxes by these pathways were used to create N budgets for the grasslands. Both grasslands were found to be accumulating much of the simulated additional N deposition: up to 89% accumulated in the calcareous grassland and up to 38% accumulated in the acidic grassland. The major fluxes of N loss from these grasslands were by simulated grazing and leaching of soluble organic N (constituting 90% of leached N under ambient conditions). Leached inorganic N (mainly nitrate) contributed significantly to the output flux of N under the highest N treatment only. Loss of N through ammonia volatilization accounted for less than 6% of the N added as simulated deposition, while denitrification contributed significantly to output fluxes only in the acidic grassland during winter. The implications of the results for ecosystem N balances and the likely consequences of N accumulation on these grasslands are discussed.     

Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands

Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change.     

Feedback of grazing on gross rates of N mineralization and inorganic N partitioning in steppe soils of Inner Mongolia

Plant-microbe interactions are crucial regulators of belowground nitrogen cycling in terrestrial ecosystems. However, such interactions have mostly been excluded from experimental setups for the investigation of gross inorganic N fluxes and N partitioning to plants and microorganisms. Ungulate grazing is likely to feed back on soil N fluxes, and hence it is of special importance to simultaneously investigate grazing effects on both plant and microbial N fluxes in intact plant-soil systems, where plant-microbe interactions persist during the experimental incubation. Based on the homogenous ¹⁵NH ₄ ⁺ labelling of intact plant-soil monoliths we investigated how various stocking rates (0, 2.35, 4.8 and 7.85 sheep ha^?1 grazing season^?1) in steppe of Inner Mongolia feedback on gross rates of N mineralization and short-term inorganic N partitioning between plant, microbial and soil N pools. Our results showed that the effect of grazing on gross N mineralization was non-uniform. At low stocking rate gross N mineralization tended to decrease but increased with higher grazing pressure. Hence, there was no significant correlation between stocking rate and gross N mineralization across the investigated grazing intensities. Grazing decreased ¹⁵N recovery both in plant and microbial N pools but strongly promoted NO ₃ ^? accumulation in the soil and thus negatively affected potential ecosystem N retention. This appeared to be closely related to the grazing-induced decline in easily degradable soil C availability at increasing stocking rate.     

Herbivore effects on above- and belowground plant production and soil nitrogen availability in the Trans-Himalayan shrub-steppes

Large mammalian herbivores may have positive, neutral, or negative effects on annual net aboveground plant production (NAP) in different ecosystems, depending on their indirect effects on availability of key nutrients such as soil N. In comparison, less is known about the corresponding influence of grazers, and nutrient dynamics, over annual net belowground plant production (NBP). In natural multi-species plant communities, it remains uncertain how grazing influences relative allocation in the above- and belowground compartments in relation to its effects on plant nutrients. We evaluated grazer impacts on NAP, NBP, and relative investment in the above- and belowground compartments, alongside their indirect effects on soil N availability in the multiple-use Trans-Himalayan grazing ecosystem with native grazers and livestock. Data show that a prevailing grazing intensity of 51% increases NAP (+61%), but reduces NBP (−35%). Grazing also reduced C:N ratio in shoots (−16%) and litter (−50%), but not in roots, and these changes coincided with increased plant-available inorganic soil N (+23%). Areas used by livestock and native grazers showed qualitatively similar responses since NAP was promoted, and NBP was reduced, in both cases. The preferential investment in the aboveground fraction, at the expense of the belowground fraction, was correlated positively with grazing intensity and with improvement in litter quality. These results are consistent with hypothesized herbivore-mediated positive feedbacks between soil nutrients and relative investment in above- and belowground compartments. Since potentially overlapping mechanisms, such as N mineralization rate, plant N uptake, compositional turnover, and soil microbial activity, may contribute towards these feedbacks, further studies may be able to discern their respective contributions.     

Reindeer influence on ecosystem processes in the tundra

Reindeer have been recorded to increase nutrient cycling rate and primary production in studies from fences almost 40 years old that separate areas with different grazing regimes in northern Fennoscandia. To further understand the mechanism behind the effects of herbivores on primary production, we measured the size of the major C and N pools, soil temperature, litter decomposition rate and N mineralization rate in lightly, moderately and heavily grazed areas along two of these fences.
Plant N found in new biomass, indicative of plant N assimilation, was significantly higher in moderately and heavily grazed areas than in lightly grazed areas, which corresponded to a decreased amount of N in old plant parts. The amount of N found in plant litter or organic soil layer did not differ between the grazing treatments. Together with soil N concentrations and litter decomposition rates, soil temperatures were significantly higher in moderately and heavily grazed areas.
We conclude that the changes in soil temperature are important for the litter decomposition rate and thus on the nutrient availability for plant uptake. However, the changes in plant community composition appear to be more important for the altered N pools and thus the enhanced primary production. The results provide some support for the keystone herbivore hypothesis, which states that intensive grazing can promote a transition from moss-rich tundra heath to productive grasslands. Grazing altered N fluxes and pools, but the total N pools were similar in all grazing treatments. Our study thus indicates that grazing can increase the primary production through enhancing the soil nutrient cycling rate, even in a long term perspective on an ecological timescale.     

Effects of nitrogen addition on vegetation and ecosystem carbon in a semi-arid grassland

To clarify responses of plant and soil carbon (C) and nitrogen (N) pools in grassland ecosystem to N addition, a field experiment was performed in a grassland in Keerqin Sandy Lands, Northeast China. We investigated vegetation composition and C and N pools of plant and soil (0–30 cm) after five consecutive years of N addition at a rate of 20 g N m^?2 y^?1. Vegetation composition and species diversity responded dramatically to N addition, as dominance by C₄ perennials was replaced with C₃ annuals. Carbon in aboveground pool increased significantly (over two-fold), mainly due to the increase of the C in aboveground living plants and surface litter, which increased by 98 and 134%, respectively. Although soil C did not change significantly, the root C pool decreased in response to 5 years of N addition. The total ecosystem C pool was not significantly impacted by N addition because the large soil pool did not respond to N addition, and the increase in aboveground C was offset by the decrease in root C pool. Moreover, N addition significantly increased the aboveground N pool, but had no significant effects on belowground and total ecosystem N pools. Our results suggest that in the mid-term N addition alters the C and N partitioning in above- and belowground pools, but has no significant effects on total ecosystem C and N pools in these N-limited grasslands.