Grazing intensity significantly affects belowground carbon and nitrogen cycling in grassland ecosystems: a meta‐analysis

Livestock grazing activities potentially alter ecosystem carbon (C) and nitrogen (N) cycles in grassland ecosystems. Despite the fact that numerous individual studies and a few meta‐analyses had been conducted, how grazing, especially its intensity, affects belowground C and N cycling in grasslands remains unclear. In this study, we performed a comprehensive meta‐analysis of 115 published studies to examine the responses of 19 variables associated with belowground C and N cycling to livestock grazing in global grasslands. Our results showed that, on average, grazing significantly decreased belowground C and N pools in grassland ecosystems, with the largest decreases in microbial biomass C and N (21.62% and 24.40%, respectively). In contrast, belowground fluxes, including soil respiration, soil net N mineralization and soil N nitrification increased by 4.25%, 34.67% and 25.87%, respectively, in grazed grasslands compared to ungrazed ones. More importantly, grazing intensity significantly affected the magnitude (even direction) of changes in the majority of the assessed belowground C and N pools and fluxes, and C : N ratio as well as soil moisture. Specifically,light grazing contributed to soil C and N sequestration whereas moderate and heavy grazing significantly increased C and N losses. In addition, soil depth, livestock type and climatic conditions influenced the responses of selected variables to livestock grazing to some degree. Our findings highlight the importance of the effects of grazing intensity on belowground C and N cycling, which may need to be incorporated into regional and global models for predicting effects of human disturbance on global grasslands and assessing the climate‐biosphere feedbacks.     

Grazing enhances belowground carbon allocation,microbial biomass,and soil carbon in a subtropical grassland

Despite the large contribution of rangeland and pasture to global soil organic carbon (SOC) stocks, there is considerable uncertainty about the impact of large herbivore grazing on SOC, especially for understudied subtropical grazing lands. It is well known that root system inputs are the source of most grassland SOC, but the impact of grazing on partitioning of carbon allocation to root tissue production compared to fine root exudation is unclear. Given that different forms of root C have differing implications for SOC synthesis and decomposition, this represents a significant gap in knowledge. Root exudates should contribute to SOC primarily after microbial assimilation, and thus promote microbial contributions to SOC based on stabilization of microbial necromass, whereas root litter deposition contributes directly as plant‐derived SOC following microbial decomposition. Here, we used in situ isotope pulse‐chase methodology paired with plant and soil sampling to link plant carbon allocation patterns with SOC pools in replicated long‐term grazing exclosures in subtropical pasture in Florida, USA. We quantified allocation of carbon to root tissue and measured root exudation across grazed and ungrazed plots and quantified lignin phenols to assess the relative contribution of microbial vs. plant products to total SOC. We found that grazing exclusion was associated with dramatically less overall belowground allocation, with lower root biomass, fine root exudates, and microbial biomass. Concurrently, grazed pasture contained greater total SOC, and a larger fraction of SOC that originated from plant tissue deposition, suggesting that higher root litter deposition under grazing promotes greater SOC. We conclude that grazing effects on SOC depend on root system biomass, a pattern that may generalize to other C4‐dominated grasslands, especially in the subtropics. Improved understanding of ecological factors underlying root system biomass may be the key to forecasting SOC and optimizing grazing management to enhance SOC accumulation.     

The effects of tree rhizodeposition on soil exoenzyme activity, dissolved organic carbon, and nutrient availability in a subalpine forest ecosystem

Previous studies have found that root carbon inputs to the soil can stimulate the mineralization of existing soil carbon (C) pools. It is still uncertain, however, whether this “primed” C is derived from elevated rates of soil organic matter (SOM) decomposition, greater C release from microbial pools, or both. The goal of this research was to determine how the activities of the microbial exoenzymes that control SOM decomposition are affected by root C inputs. This was done by manipulating rhizodeposition with tree girdling in a coniferous subalpine forest in the Rocky Mountains of Colorado, USA, and following changes in the activities of nine exoenzymes involved in decomposition, as well as soil dissolved organic C, dissolved organic and inorganic nitrogen (N), and microbial biomass C and N. We found that rhizodeposition is high in the spring, when the soils are still snow-covered, and that there are large ephemeral populations of microorganisms dependent upon this C. Microbial N acquisition from peptide degradation increased with increases in microbial biomass when rhizodeposition was highest. However, our data indicate that the breakdown of cellulose, lignin, chitin, and organic phosphorus are not affected by springtime increases in soil microbial biomass associated with increases in rhizodeposition. We conclude that the priming of soil C mineralization by rhizodeposition is due to growth of the microbial biomass and an increase in the breakdown of N-rich proteins, but not due to increases in the degradation of plant litter constituents such as cellulose and lignin.     

Non-parallel changes in soil microbial carbon and nitrogen dynamics due to reindeer grazing in northern boreal forests

Reindeer grazing has a considerable influence on mineralization processes in northern Fennoscandian boreal forests, but the mechanisms underlying the observed differences between grazed and ungrazed areas are not well understood. We studied the below-ground impacts of reindeer grazing by comparing the carbon and nitrogen mineralization rates inside and outside long-term fenced reindeer exclosure areas in five oligotrophic, lichen-dominated and five mesotrophic, dwarf-shrub dominated forests. The soil C mineralization rates and microbial metabolic activity (qCO₂) were significantly lower in the grazed than the ungrazed areas in both oligotrophic and mesotrophic forests. The reductions occurred irrespective of the impact on soil moisture. We conclude that reindeer grazing causes a reduction in the supply of labile C substrates to microbes, resulting in reduced organic matter decomposition rates through changes in the activity of the microbial biomass. Simultaneously, grazing had no consistent effect on the microbial N dynamics, but the impact ranged from no change to increased or decreased in N mineralization rates at the different study sites. The impact of grazing on the N mineralization potential thus seems to be site-specific and uncoupled from the impact of grazing on soil C mineralization. Reciprocal transplant incubations showed no interactions between N mineralization rates and the reindeer-mediated impact on the soil microclimate. We suggest that plant root damage due to trampling by reindeer may be an important mechanism for the deceleration of soil C cycling. In some cases, however, the impact of grazing on the soil active N pool may be strong enough to outweigh the reduction in soil organic matter decomposition, and by these means uncouple soil N dynamics from soil C quality.     

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.     

The effect of reindeer grazing on decomposition, mineralization and soil biota in a dry oligotrophic Scots pine forest

Reindeer grazing in the Fennoscandian area has a considerable influence on the ground vegetation, and this is likely in turn to have important consequences for the soil biota and decomposition processes. The effects of reindeer grazing on soil biota, decomposition and mineralization processes, and ecosystem properties in a lichen‐dominated forest in Finnish Lapland were studied inside and outside a large long term fenced reindeer exclosure area. Decomposition rates of Vaccinium myrtillus leaves in litter bags were retarded in the grazed area relative to the ungrazed area, as well as in subplots from which lichens had been artificially removed to simulate grazing. The effect of reindeer grazing on soil respiration and microbial C was positive in the lichen and litter layers of the soil profile, but retarded in the humus layer. There was no effect of grazing on gross N mineralization and microbial biomass N in the humus and upper mineral soil layer, but net N mineralization was increased by grazing. In these layers soil respiration was reduced by grazing, indicating that reindeer effects reduce the ratio of C to N mineralized by soil microorganisms. Grazing stimulated populations of all trophic groupings of nematodes in the lichen layer and microbe feeding nematodes in the litter layer, indicating that grazing by reindeer has multitrophic effects on the decomposer food‐web. Grazing decreased lichen and dwarf shrub biomasses and increased the mass of litter present in the litter layer on an areal basis, but did not significantly alter total C storage per unit area in the humus and mineral soil layers. The N concentration of lichens was increased by grazing, but the N concentrations of both living and dead Pinus sylvestris needles and Empetrum hermaphroditum leaves were not affected.
There was some evidence for each of three mechanisms which could account for the grazing effects that we observed in our study. Firstly, reindeer may have changed the composition and quality of litter input by affecting plant species composition and through addition of N from urine and faeces, resulting in a lack of available C relative to N for decomposer organisms. Secondly, the organic matter in the soil may be older in the grazed area, because of reduction of recent production of lichen litter relative to the ungrazed area. The organic matter in the grazed area may have been in a different phase of decomposition from that in the exclosure. Thirdly, the soil microclimate is likely to be affected by reindeer grazing through physical removal of lichen cover on the ground, and this can have a significant influence on soil microbial processes. This is supported by the strong observed effects of experimental removal of lichens on decomposer processes. The impact of reindeer grazing on soil processes may be a result of complex interactions between different mechanisms, and this could help to explain why the below‐ground effects of reindeer grazing have different consequences to those which have been observed in recent investigations on other grazing systems.     

Effects of livestock grazing intensity on soil arbuscular mycorrhizal fungi and glomalin-related soil protein in a mountain forest steppe and a desert steppe of Mongolia

Arbuscular mycorrhizal fungi (AMF) are important components of the grassland ecosystems in terms of plant phosphorus uptake and accumulation of glomalin-related soil protein (GRSP). Though Mongolian grasslands are seriously degraded by livestock grazing, the effects of grazing on soil AMF and GRSP remain unclear. Here, we examined community composition and diversity of AMF as well as amount of GRSP at three different grazing intensities: lightly grazed (LG), moderately grazed (MG), and heavily grazed (HG) under two different types of grassland, mountain forest steppe at Hustai and desert steppe at Mandalgobi. The diversity and biomass of AMF-host and non-AMF plants strongly affected the overall AMF community composition and its diversity. When we separately analyzed the factors affecting soil AMF diversity at Hustai and Mandalgobi, decrease in the shoot biomass of Poaceae plants at Hustai and decreases in the species number and shoot biomass of AMF-host plants at Mandalgobi were significantly correlated with AMF diversity. GRSP decreased with increasing grazing intensity, which was significantly correlated with soil pH and total root biomass at Hustai. The decrease in plant biomass caused by grazing thus led to GRSP reduction. Our results showed that change in soil AMF community caused by livestock grazing were associated with change in the biomass and diversity of functional vegetation groups such as Poaeceae, AMF-host and non-AMF plants, indicating the importance to focus on such functional vegetation groups to evaluate the effect of grazing on AMF.

     

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.     

15N enrichment as an integrator of the effects of C and N on microbial metabolism and ecosystem function

Organic carbon (C) and nitrogen (N) are essential for heterotrophic soil microorganisms, and their bioavailability strongly influences ecosystem C and N cycling. We show here that the natural ¹⁵N abundance of the soil microbial biomass is affected by both the availability of C and N and ecosystem N processing. Microbial ¹⁵N enrichment correlated negatively with the C : N ratio of the soil soluble fraction and positively with net N mineralization for ecosystems spanning semiarid, temperate and tropical climates, grassland and forests, and over four million years of ecosystem development. In addition, during soil incubation, large increases in microbial ¹⁵N enrichment corresponded to high net N mineralization rates. These results support the idea that the N isotope composition of an organism is determined by the balance between N assimilation and dissimilation. Thus, ¹⁵N enrichment of the soil microbial biomass integrates the effects of C and N availability on microbial metabolism and ecosystem processes.     

半干旱草地长期封育进程中针茅植物根系格局变化特征

以云雾山不同封育年限草地针茅植物根系和土壤为研究对象,对其根系特征、土壤特性及两者关系进行研究,以探讨分析封育对针茅根系格局的影响。结果表明:(1)针茅植物根系生物量、根长密度、根表面积和根体积在封育初期轻微下降,之后缓慢上升,并在封育30 a草地得到显著增加。(2)随封育年限增加,各根系指标在3种针茅物种间的组成格局具有类似变化规律,具体表现为:长芒草在放牧草地所占比例最高,之后逐渐降低,并在封育30 a草地消失;大针茅所占比例呈先升后降变化规律,并在封育22 a草地达到最大值;甘青针茅仅出现于封育30 a草地,且占据优势地位。(3)大针茅和甘青针茅0—0.6 mm径级根系比例高于大针茅,使其根系直径显著低于大针茅,比根长和比根面积显著高于大针茅;此外,长芒草根组织密度显著高于长芒草和甘青针茅。(4)长期封育在显著提高土壤水分、养分含量和土壤氮磷比的同时显著降低土壤碳氮比,但对微生物生物量碳、氮无明显影响。(5)针茅根系特征与土壤指标的关联性分析显示针茅根系受土壤氮资源的显著影响。     

草地生态系统中土壤氮素矿化影响因素的研究进展

氮素是各种植物生长和发育所需的大量营养元素之一,也是牧草从土壤吸收最多的矿质元素．土壤中的氮大部分以有机态形式存在,而植物可以直接吸收利用的是无机态氮．这些有机态氮在土壤动物和微生物的作用下。由难以被植物直接吸收利用的有机态转化为可被植物直接吸收利用的无机态的过程就是土壤氮的矿化．氮素矿化受多种因子的影响,这些因子可以归结为生物因子和非生物因子．生物因子包括：土壤动物、土壤微生物和植物种类．土壤动物可以促进土壤有机质的矿化;土壤微生物种类、结构及功能与氮的分解、矿化有密切的关系;不同的植物种类对土壤氮素的矿化作用是不相同的,一般来说。有豆科植物生长的土壤比其它种类土氮素矿化的作用大．非生物因素一般可以分为环境因子和人类活动干扰．环境因子中土壤温度和含水量对土壤氮素矿化的影响是国内外众多科学家研究的方向．尽管如此,在此方面的研究还没有取得一致意见,仍然需要进行这方面的研究,而在其他诸如：不同的土壤质地与土壤类型方面,研究报道的结论也很不一致,草地生态系统中人类活动对土壤氮素矿化的影响主要包括,不同强度的放牧,割草以及施肥、火烧强度等．非生物因子对氮素矿化的影响非常直接和明显,尤其是人类活动．本文综述了近年来影响草地生态系统土壤氮素矿化有关因素的一些进展．     

Grazing history effects on above- and below-ground litter decomposition and nutrient cycling in two co-occurring grasses

Large herbivores may alter carbon and nutrient cycling in soil by changing above- and below-ground litter decomposition dynamics. Grazing effects may reflect changes in plant allocation patterns, and thus litter quality, or the site conditions for decomposition, but the relative roles of these broad mechanisms have rarely been tested. We examined plant and soil mediated effects of grazing history on litter mass loss and nutrient release in two grazing-tolerant grasses, Lolium multiflorum and Paspalum dilatatum, in a humid pampa grassland, Argentina. Shoot and root litters produced in a common garden by conspecific plants collected from grazed and ungrazed sites were incubated under both grazing conditions. We found that grazing history effects on litter decomposition were stronger for shoot than for root material. Root mass loss was neither affected by litter origin nor incubation site, although roots from the grazed origin immobilised more nutrients. Plants from the grazed site produced shoots with higher cell soluble contents and lower lignin:N ratios. Grazing effects mediated by shoot litter origin depended on the species, and were less apparent than incubation site effects. Lolium shoots from the grazed site decomposed and released nutrients faster, whereas Paspalum shoots from the grazed site retained more nutrient than their respective counterparts from the ungrazed site. Such divergent, species-specific dynamics did not translate into consistent differences in soil mineral N beneath decomposing litters. Indeed, shoot mass loss and nutrient release were generally faster in the grazed grassland, where soil N availability was higher. Our results show that grazing influenced nutrient cycling by modifying litter breakdown within species as well as the soil environment for decomposition. They also indicate that grazing effects on decomposition are likely to involve aerial litter pools rather than the more recalcitrant root compartment.     

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。     

Insensitivity of Soil Microbial Activity to Temporal Variation in Soil N in Subarctic Tundra: Evidence from Responses to Large Migratory Grazers

Large migratory grazers commonly influence soil processes in tundra ecosystems. However, the extent to which grazing effects are limited to intensive grazing periods associated with migration has not previously been investigated. We analyzed seasonal patterns in soil nitrogen (N), microbial respiration and extracellular enzyme activities (EEAs) in a lightly grazed tundra and a heavily grazed tundra that has been subjected to intensive grazing during reindeer (Rangifer tarandus L.) migration for the past 50 years. We hypothesized that due to the fertilizing effect of the reindeer, microbial respiration and EEAs related to microbial C acquisition should be higher in heavily grazed areas compared to lightly grazed areas and that the effects of grazing should be strongest during reindeer migration. Reindeer migration caused a dramatic peak in soil N availability, but in contrast to our predictions, the effect of grazing was more or less constant over the growing season and the seasonal patterns of microbial activities and microbial N were strikingly uniform between the lightly and heavily grazed areas. Microbial respiration and the EEAs of β-glucosidase, acid-phosphatase, and leucine-aminopeptidase were higher, whereas that of N-acetylglucosamidase was lower in the heavily grazed area. Experimental fertilization had no effect on EEAs related to C acquisition at either level of grazing intensity. Our findings suggest that soil microbial activities were independent of grazing-induced temporal variation in soil N availability. Instead, the effect of grazing on soil microbial activities appeared to be mediated by substrate availability for soil microorganisms. Following a shift in the dominant vegetation in response to grazing from dwarf shrubs to graminoids, the effect of grazing on soil processes is no longer sensitive to temporal grazing patterns; rather, grazers exert a consistent positive effect on the soil microbial potential for soil C decomposition.     

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.     

Changes in soil microbial biomass and community structure with addition of contrasting types of plant litter in a semiarid grassland ecosystem

Aims Elevated atmospheric CO₂ has the potential to enhance the net primary productivity of terrestrial ecosystems. However, the role of soil microorganisms on soil C cycling following this increased available C remains ambiguous. This study was conducted to determine how quality and quantity of plant litter inputs would affect soil microorganisms and consequently C turnover.Methods Soil microbial biomass and community structure, bacterial community-level physiological profile, and CO₂ emission caused by different substrate C decomposition were investigated using techniques of biological measurements, chemical and stable C isotope analysis, and BIOLOG-ECO microplates in a semiarid grassland ecosystem of northern China in 2006 and 2007 by mixing three contrasting types of plant materials, C₃ shoot litter (SC 3), C₃ root litter (RC 3), and C₄ shoot litter (SC 4), into the 10- to 20-cm soil layer at rates equivalent to 0 (C 0), 60 (C 60), 120 (C 120) and 240 g C m ?2 (C 240).Important findings Litter addition significantly enriched soil microbial biomass C and N and resulted in changes in microbial structure. Principal component analysis of microbial structure clearly differentiated among zero addition, C₃ -plant-derived litter, and C₄ -plant-derived litter and among shoot- and root-derived litter of C₃ plants; soil microorganisms mainly utilized carbohydrates without litter addition, carboxylic acids with C₃ -plant-derived litter addition and amino acids with C₄ -plant-derived litter addition. We also detected stimulated decomposition of older substrate with C₄ -plant-derived litter inputs. Our results show that both quality and quantity of belowground litter are involved in affecting soil microbial community structure in semiarid grassland ecosystem.     

Six years of in situ CO2 enrichment evoke changes in soil structure and soil biota of nutrient-poor grassland

Nutrient‐poor grassland on a silty clay loam overlying calcareous debris was exposed to elevated CO₂ for six growing seasons. The CO₂ exchange and productivity were persistently increased throughout the experiment, suggesting increases in soil C inputs. At the same time, elevated CO₂ lead to increased soil moisture due to reduced evapotransporation. Measurements related to soil microflora did not indicate increased soil C fluxes under elevated CO₂. Microbial biomass, soil basal respiration, and the metabolic quotient for CO₂ (qCO₂) were not altered significantly. PLFA analysis indicated no significant shift in the ratio of fungi to bacteria. 0.5 m KCl extractable organic C and N, indicators of changed DOC and DON concentrations, also remained unaltered. Microbial grazer populations (protozoa, bacterivorous and fungivorous nematodes, acari and collembola) and root feeding nematodes were not affected by elevated CO₂. However, total nematode numbers averaged slightly lower under elevated CO₂ (?16%, ns) and nematode mass was significantly reduced (?43%, P = 0.06). This reduction reflected a reduction in large‐diameter nematodes classified as omnivorous and predacious. Elevated CO₂ resulted in a shift towards smaller aggregate sizes at both micro‐ and macro‐aggregate scales; this was caused by higher soil moisture under elevated CO₂. Reduced aggregate sizes result in reduced pore neck diameters. Locomotion of large‐diameter nematodes depends on the presence of large enough pores; the reduction in aggregate sizes under elevated CO₂ may therefore account for the decrease in large nematodes. These animals are relatively high up the soil food web; this decline could therefore trigger top‐down effects on the soil food web. The CO₂ enrichment also affected the nitrogen cycle. The N stocks in living plants and surface litter increased at elevated CO₂, but N in soil organic matter and microbes remained unaltered. Nitrogen mineralization increased markedly, but microbial N did not differ between CO₂ treatments, indicating that net N immobilization rates were unaltered. In summary, this study did not provide evidence that soils and soil microbial communities are affected by increased soil C inputs under elevated CO₂. On the contrary, available data (¹³C tracer data, minirhizotron observations, root ingrowth cores) suggests that soil C inputs did not increase substantially. However, we provide first evidence that elevated CO₂ can reduce soil aggregation at the scale from µ m to mm scale, and that this can affect soil microfaunal populations.     

Reindeer grazing and soil microbial processes in two suboceanic and two subcontinental tundra heaths

In oceanic, nutrient-rich Fennoscandian arctic-alpine tundra heaths, grazing by reindeer has been found to increase herbs and graminoids in relation to dwarf shrubs. In continental lichen heaths in the inland with nutrient-poor conditions, however, slowly decomposable dwarf shrubs are favoured by grazing. According to a hypothesis, by favouring easily decomposing plants in nutrient-rich conditions and slowly decomposing plants in nutrient-poor conditions, herbivory enhances soil nutrient cycling in nutrient-rich and retards it in nutrient-poor areas. We tested this hypothesis by comparing the impact of reindeer grazing on soil C and N mineralization between two oceanic and two continental arctic-alpine tundra heaths.
Although soil respiration and microbial metabolic activity were enhanced by grazing in the suboceanic but not in the subcontinental tundra heaths, gross N mineralization rates were higher in the grazed areas in soils from all study sites, indicating that reindeer grazing leads to increased rates of nutrient cycling in both nutrient-poor and nutrient-rich tundra heaths. Thus, in the subcontinental tundra heaths, the increase in soil N concentrations due to mammalian waste products enhances N mineralization rates, even though the organic C quality is not improved by reindeer grazing. There was some site-specific variation in the strength of the reindeer effects on various microbial processes and soil properties, which can be related to spatial variation in grazing intensity and timing, as these factors in turn affect the nutrient sink strength of the vegetation.     

Plant and soil microbial responses to defoliation in temperate semi-natural grassland

There is much interest in understanding the nature of feedback mechanisms between plants and soil organisms in grazed ecosystems. In this study, we examine the effects of different intensities of defoliation on the growth of three dominant grass species, and observe how these plant responses relate to the biomass and activity of the microbial community in the root zone. Our data show that grassland plants with varying tolerances to grazing have markedly different growth responses to defoliation, and that these responses vary with the intensity of cutting. Defoliation of grasses which are tolerant to grazing, namely Festuca rubra and Cynosurus cristatus, leads to a reduction in root mass and an increase in the allocation of resources to shoots. In contrast, defoliation of a grass with low tolerance to grazing, Anthoxanthum odoratum, had little effect on root mass, but increased the relative allocation of resources below-ground. In all plant species, defoliation led to an increase in soil microbial biomass and C use efficiency in the root zone. This response was greatest in the root zone of A. odoratum and is likely to be related to changes in root exudation pattern following defoliation. The significance of these changes in relation to soil nutrient dynamics and plant nutrient uptake during regrowth require further exploration. This revised version was published online in June 2006 with corrections to the Cover Date.     

Ungulate stimulation of nitrogen cycling and retention in Yellowstone Park grasslands

We studied how ungulates and a large variation in site conditions influenced grassland nitrogen (N) dynamics in Yellowstone National Park. In contrast to most grassland N studies that have examined one or two soil N processes, we investigated four rates, net N mineralization, nitrification, denitrification, and inorganic N leaching, at seven paired sites inside and outside long-term (33+ year) exclosures. Our focus was how N fluxes were related to one another among highly variable grasslands and how grazers influenced those relationships. In addition, we examined variation in soil δ¹⁵N among grasslands and the relationships between soil ¹⁵N abundance and N processes. Previously, ungulates were reported to facilitate net N mineralization across variable Yellowstone grasslands and denitrification at mesic sites. In this study, we found that herbivores also promoted nitrification among diverse grasslands. Furthermore, net N mineralization, nitrification, and denitrification (kg N ha^–1 year^–1, each variable) were postively and linearly related to one another among all grasslands (grazed and fenced), and grazers reduced the nitrification/net N mineralization and denitrification/net N mineralization ratios, indicating that ungulates inhibited the proportion of available NH₄ ⁺ that was nitrified and denitrified. There was no relationship between net N mineralization or nitrification with leaching (indexed by inorganic N adsorbed to resin buried at the bottom of rooting zones) and leaching was unaffected by grazers. Soil δ¹⁵N was positively and linearly related to in situ net N mineralization and nitrification in ungrazed grasslands; however, there was no relationship between isotopic composition of N and those rates among grazed grasslands. The results suggested that grazers simultaneously increased N availability (stimulated net N mineralization and nitrification per unit area) and N conservation (reduced N loss from the soil per unit net N mineralization) in Yellowstone grasslands. Grazers promoted N retention by stimulating microbial productivity, probably caused by herbivores promoting labile soil C. Process-level evidence for N retention by grazers was supported by soil δ¹⁵N data. Grazed grassland with high rates of N cycling had substantially lower soil δ¹⁵N relative to values expected for ungrazed grassland with comparable net N mineralization and nitrification rates. These soil ¹⁵N results suggest that ungulates inhibited N loss at those sites. Such documented evidence for consumer control of N availability to plants, microbial productivity, and N retention in Yellowstone Park is further testimony for the widespread regulation of grassland processes by large herbivores. Received: 5 May 1999 / Accepted: 1 November 1999