Mineralization and distribution of nutrients in plants and microbes in four arctic ecosystems: responses to warming

Mineralization and nutrient distribution in plants and microbes were studied in four arctic ecosystems at Abisko, Northern Sweden and Toolik Lake, Alaska, which have been subjected to long-term warming with plastic greenhouses. Net mineralization and microbial immobilization were studied by the buried bag method and ecosystem pool sizes of C, N and P were determined by harvest methods. The highest amounts of organic N and P were bound in the soil organic matter. Microbial N and P constituted the largest labile pools often equal to (N) or exceeding (P) the amounts stored in the vegetation. Despite large pools of N and P in the soil, net mineralization of N and P was generally low during the growing season, except in the wet sedge tundra, and in most cases lower than the plant uptake requirement. In contrast, the microorganisms immobilized high amounts of nutrients in the buried bags during incubation. The same high immobilization was not observed in the surrounding soil, where the microbial nutrient content in most cases remained constant or decreased over the growing season. This suggests that the low mineralization measured in many arctic ecosystems over the growing season is due to increased immobilization by soil microbes when competition from plant roots is prevented. Furthermore, it suggests that plants compete well with microbes for nutrients in these four ecosystems. Warming increased net mineralization in several cases, which led to increased assimilation of nutrients by plants but not by the microbes.     

Soil nitrogen cycling rates in low arctic shrub tundra are enhanced by litter feedbacks

Shrub growth has increased across the Arctic in recent decades and is strongly limited by soil nitrogen (N) availability. In order to understand the role of N in controlling shrub growth, we compared N-cycling in tall birch (Betula glandulosa) and surrounding dwarf birch hummock vegetation on similar soils in a Canadian low arctic site. Stable isotope tracer analysis revealed N pools and cycling rates were ～3 times larger and faster in the tall birch ecosystem in the late growing season, just prior to leaf senescence. Gross NH ₄ ⁺ -N production rates in these ecosystems correlated positively with larger pools and production rates of dissolved soil C and N, higher quality litter inputs and lower soil C. Analyses of the soil microbial community in both ecosystems indicated similar fungal dominance (epifluorescence microscopy) and similar compositions of the principal fungal or bacterial phylotypes (denaturing gradient gel electrophoresis). Together, these results strongly suggest that vegetation feedbacks associated with larger inputs of higher quality litter promote rapid soil N-cycling and enhanced shrub growth in tall birch tundra. We conclude that these litter-related feedbacks during summer may be as important as snow-shrub feedbacks in maintaining and promoting differences in shrub growth across the arctic landscape.     

Interactive Effects of Nitrogen and Phosphorus on Soil Microbial Communities in a Tropical Forest

Elevated nitrogen (N) deposition in humid tropical regions may exacerbate phosphorus (P) deficiency in forests on highly weathered soils. However, it is not clear how P availability affects soil microbes and soil carbon (C), or how P processes interact with N deposition in tropical forests. We examined the effects of N and P additions on soil microbes and soil C pools in a N-saturated old-growth tropical forest in southern China to test the hypotheses that (1) N and P addition will have opposing effects on soil microbial biomass and activity, (2) N and P addition will alter the composition of the microbial community, (3) the addition of N and P will have interactive effects on soil microbes and (4) addition-mediated changes in microbial communities would feed back on soil C pools. Phospholipid fatty acid (PLFA) analysis was used to quantify the soil microbial community following four treatments: Control, N addition (15 g N m⁻² yr⁻¹), P addition (15 g P m⁻² yr⁻¹), and N&P addition (15 g N m⁻² yr⁻¹ plus 15 g P m⁻² yr⁻¹). These were applied from 2007 to 2011. Whereas additions of P increased soil microbial biomass, additions of N reduced soil microbial biomass. These effects, however, were transient, disappearing over longer periods. Moreover, N additions significantly increased relative abundance of fungal PLFAs and P additions significantly increased relative abundance of arbuscular mycorrhizal (AM) fungi PLFAs. Nitrogen addition had a negative effect on light fraction C, but no effect on heavy fraction C and total soil C. In contrast, P addition significantly decreased both light fraction C and total soil C. However, there were no interactions between N addition and P addition on soil microbes. Our results suggest that these nutrients are not co-limiting, and that P rather than N is limiting in this tropical forest.     

High arctic heath soil respiration and biogeochemical dynamics during summer and autumn freeze‐in – effects of long‐term enhanced water and nutrient supply

In High Arctic NE Greenland, temperature and precipitation are predicted to increase during this century, however, relatively little information is available on the role of increased water supply on soil CO ₂ efflux in dry, high arctic ecosystems. We measured soil respiration (R_soil) in summer and autumn of 2009 in combination with microbial biomass and nutrient availability during autumn freeze‐in at a dry, open heath in Zackenberg, NE Greenland. This tundra site has been subject to fully factorial manipulation consisting of increased soil water supply for 14 years, and occasional nitrogen (N) addition in pulses. Summer watering enhanced R_soil during summer, but decreased R_soil in the following autumn. We speculate that this is due to intensified depletion of recently fixed plant carbon by soil organisms. Hence, autumn soil microbial activity seems tightly linked to growing season plant production through plant‐associated carbon pools. Nitrogen addition alone consistently increased R_soil, but when water and nitrogen were added in combination, autumn R_soil declined similarly to when water was added alone. Despite several freeze‐thaw events, the microbial biomass carbon (C) remained constant until finally being reduced by ~60% in late September. In spite of significantly reduced microbial biomass C and phosphorus (P), microbial N did not change. This suggests N released from dead microbes was quickly assimilated by surviving microbes. We observed no change in soil organic matter content after 14 years of environmental manipulations, suggesting high ecosystem resistance to environmental changes.     

Soil microbial responses to elevated phosphorus and pH in acidic temperate deciduous forests

Although northern temperate forests are generally not considered phosphorus (P) limited, ecosystem P limitation may occur on highly weathered or strongly acidic soils where bioavailable inorganic P is low. In such environments, soil organisms may compensate by increasing the utilization of organic P via the production of extracellular enzymes to prevent limitation. In this study, we experimentally increased available P and/or pH in several acidic eastern deciduous forests underlain by glaciated and unglaciated soils in eastern Ohio, USA. We hypothesized that where inorganic P is low; soil microbes are able to access organic P by increasing production of phosphatase enzymes, thereby overcoming biogeochemical P limitations. We measured surface soil for: available P pools, N mineralization and nitrification rates, total C and N, enzymes responsible for C, N, and P hydrolysis, and microbial community composition (PLFA). Increasing surface soil pH a whole unit had little effect on microbial community composition, but increased N cycling rates in unglaciated soils. Phosphorus additions suppressed phosphatase activities over 60% in the unglaciated soils but were unchanged in the glaciated soils. All treatments had minimal influence on microbial biomass, but available pools of P strongly correlated with microbial composition. Microbes may be dependent on sources of organic P in some forest ecosystems and from a microbial perspective soil pH might be less important overall than P availability. Although our sampling was conducted less than 1 year after treatment initiation, microbial community composition was strongly influenced by available P pools and these effects may be greater than short-term increases in soil pH.     

Soil biogeochemistry during the early spring in low arctic mesic tundra and the impacts of deepened snow and enhanced nitrogen availability

Air temperature freeze–thaw cycles often occur during the early spring period directly after snowmelt and before budbreak in low arctic tundra. This early spring period may be associated with nitrogen (N) and carbon (C) loss from soils as leachate or as trace gases, due to the detrimental impact of soil freeze–thaw cycles and a developing active layer on soil microorganisms. We measured soil and microbial pools of C and N in early spring during a period of fluctuating air temperature (ranging from ?4 to +10°C) and in midsummer, in low arctic birch hummock tundra. In addition we measured N₂O, CH₄ and CO₂ production in the early spring. All of these biogeochemical variables were also measured in long-term snowfence (deepened snow) and N-addition plots to characterize climate-change related controls on these variables. Microbial and soil solution pools of C and N, and trace gas production varied among the five early spring sample dates, but only marginally and no more than among sample dates in midsummer. N-addition greatly elevated N₂O fluxes, indicating that although denitrifiers were present their activity during early spring was strongly limited by N-availability, but otherwise trace gas production was very low in early spring. The later thaw, warmer winter and colder spring soil temperatures resulting from deepened snow did not significantly alter N pools or rates in early spring. Together, our results indicate strong stability in microbial and soil solution C and N pool sizes in the early spring period just after snowmelt when soil temperatures are close to 0°C (?1.5 to +5°C). A review of annual temperature records from this and other sites suggests that soil freeze–thaw cycles are probably infrequent in mesic tundra in early spring. We suggest that future studies concerned with temperature controls on soil and microbial biogeochemistry should focus not on soil freeze–thaw cycles per se, but on the rapid and often stepped increases in soil temperature that occur under the thawing snowpack.     

黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征的影响

研究黄土丘陵区植被与地形特征对土壤和土壤微生物生物量生态化学计量特征影响有助于深入理解黄土丘陵区不同植被带下土壤和土壤微生物相互作用及养分循环规律.选择黄土丘陵区延河流域3个植被区(森林区、森林草原区、草原区)和5种地形部位(阴/阳沟坡、阴/阳梁峁坡、峁顶)的土壤作为研究对象,利用生态化学计量学理论研究植被和地形对土壤和土壤微生物生物量生态化学计量特征的影响.结果表明: 土壤及土壤微生物生物量碳、氮、磷含量在不同地形之间的差别主要表现在沟坡位置和阴坡高于其他坡位和阳坡.植被类型的变化对两个土层(0～10、10～20 cm)土壤和土壤微生物生物量碳、氮、磷的影响均达到显著水平,坡向对表层(0～10 cm)土壤和土壤微生物生物量碳、氮、磷的影响强于坡位,而在10～20 cm土层,坡位对土壤和土壤微生物生物量碳、氮、磷影响更显著.植被类型显著影响土壤C∶N、C∶P、N∶P和土壤微生物生物量C∶N、C∶P,坡向和坡位仅影响土壤C∶P和N∶P,植被类型的变化是影响土壤C∶N的主要因素.同时,植被类型对土壤养分和微生物生物量碳、氮、磷含量及其生态化学计量特征的影响大于地形因子.标准化主轴分析结果表明,黄土丘陵区不同植被带土壤微生物具有内稳性,特别在草原带,土壤微生物生物量生态化学计量学特征具有更加严格的约束比例.在黄土丘陵区,土壤微生物生物量N∶P或许可以作为判断养分限制的另一个有力工具,若将土壤微生物生物量N∶P与植物叶片N∶P配合使用可能有助于我们更加精确地判断黄土丘陵区的土壤养分限制情况.     

喀斯特区不同退化程度植被群落植物-凋落物-土壤-微生物生态化学计量特征

为摸清喀斯特植被退化对群落各组分C、N、P生态化学计量特征及内稳态特征的影响,为喀斯特退化生态系统植被恢复与重建提供科学依据,以桂西北喀斯特地区5种退化程度植被群落为研究对象,测定了不同退化程度植被群落植物叶片、凋落物、土壤和微生物生物量的C、N、P含量,分析其化学计量比特征、相互关系及植物内稳性特征。结果表明：(1)随着退化程度加剧,叶片C、N、P含量、N∶P和凋落物N∶P、微生物量C显著下降,而叶片C∶N、C∶P则显著增加,且植物叶片N∶P<14;随退化程度加剧,凋落物N、P含量、土壤C、N、P含量、微生物量N、P呈先略有增后显著降低的趋势,且不同退化程度群落土壤N∶P和微生物量C∶N无显著差异。(2)叶片N、P含量与土壤N、P含量,叶片C∶P与土壤C∶N、C∶P、N∶P,叶片N∶P与凋落物N、N∶P,叶片C、N、P含量与微生物量C呈显著或极显著正相关关系;叶片C∶N与土壤C、N,叶片C∶P与土壤N、P,叶片N∶P与土壤P呈显著或极显著负相关关系。(3)喀斯特地区植物叶片N、P元素的内稳性指数(H)平均值分别为2.74和2.31,属于弱稳态型,叶片N∶P的H值为5.14,为稳...     

Influence of decomposer food web structure and nitrogen availability on plant growth

We studied the sensitivity of soil microbial communities and ecosystem processes to variation in the vertical and horizontal structure of decomposer food web under nitrogen poor and N-enriched conditions. Microcosms with humus and litter layer of boreal forest floor, birch seedlings infected with mycorrhizal fungi, and decomposer food webs with differing trophic group and species composition of soil fauna were constructed. During the second growing period for the birch, we irrigated half of the microcosms with urea solution, and the other half with de-ionised water to create two levels of N concentration in the substrate. During the experiment night time respirations of the microcosms were measured, and the water leached through the microcosms was analysed for concentration of mineral N, and nematode numbers. The microcosms were destructively sampled after 37 weeks for plant biomass and N uptake, structure of soil animal and microbial community (indicated by PLFA profiles), and physical and chemical properties of the humus and litter materials. Predatory mites and nematodes had a negative influence on the biomass of their microbivorous and microbi-detritivorous prey, and microbi-detritivores affected the biomass and community structure of microbes (indicated by PLFA-analysis). Moreover, predatory mites and nematodes increased microbial biomass and changed the microbial community structure. The decomposer food web structure affected also N uptake and growth of plants. Microbi-detritivorous fauna had a positive effect, whereas predators of microbial and detritus feeding fauna exerted a negative influence on plant N uptake and biomass production. The impact of a trophic group on the microbes and plant was also strongly dependent on species composition within the group. Nitrogen addition magnified the influence of food web structure on microbial biomass and plant N uptake. We suggest that addition of urea-N to the soil modified the animal-microbe interaction by increasing microbial growth and altering community structure of microbes. The presence of microbi-detritivores and predators reduced loss of carbon from the microcosms, and the food web structure influenced also water holding capacity of the materials. The changes in plant growth, nutrient cycling, size of N and C pools, and in the physical properties of the soil emphasize the importance and diversity of indirect consequences of decomposer food web structure. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nitrogen availability increases in a tundra ecosystem during five years of experimental permafrost thaw

Perennially frozen soil in high latitude ecosystems (permafrost) currently stores 1330–1580 Pg of carbon (C). As these ecosystems warm, the thaw and decomposition of permafrost is expected to release large amounts of C to the atmosphere. Fortunately, losses from the permafrost C pool will be partially offset by increased plant productivity. The degree to which plants are able to sequester C, however, will be determined by changing nitrogen (N) availability in these thawing soil profiles. N availability currently limits plant productivity in tundra ecosystems but plant access to N is expected improve as decomposition increases in speed and extends to deeper soil horizons. To evaluate the relationship between permafrost thaw and N availability, we monitored N cycling during 5 years of experimentally induced permafrost thaw at the Carbon in Permafrost Experimental Heating Research (CiPEHR) project. Inorganic N availability increased significantly in response to deeper thaw and greater soil moisture induced by Soil warming. This treatment also prompted a 23% increase in aboveground biomass and a 49% increase in foliar N pools. The sedge Eriophorum vaginatum responded most strongly to warming: this species explained 91% of the change in aboveground biomass during the 5 year period. Air warming had little impact when applied alone, but when applied in combination with Soil warming, growing season soil inorganic N availability was significantly reduced. These results demonstrate that there is a strong positive relationship between the depth of permafrost thaw and N availability in tundra ecosystems but that this relationship can be diminished by interactions between increased thaw, warmer air temperatures, and higher levels of soil moisture. Within 5 years of permafrost thaw, plants actively incorporate newly available N into biomass but C storage in live vascular plant biomass is unlikely to be greater than losses from deep soil C pools.     

C:N:P stoichiometry in soil: is there a “Redfield ratio” for the microbial biomass?

Well-constrained carbon:nitrogen:phosphorus (C:N:P) ratios in planktonic biomass, and their importance in advancing our understanding of biological processes and nutrient cycling in marine ecosystems, has motivated ecologists to search for similar patterns in terrestrial ecosystems. Recent analyses indicate the existence of “Redfield-like” ratios in plants, and such data may provide insight into the nature of nutrient limitation in terrestrial ecosystems. We searched for analogous patterns in the soil and the soil microbial biomass by conducting a review of the literature. Although soil is characterized by high biological diversity, structural complexity and spatial heterogeneity, we found remarkably consistent C:N:P ratios in both total soil pools and the soil microbial biomass. Our analysis indicates that, similar to marine phytoplankton, element concentrations of individual phylogenetic groups within the soil microbial community may vary, but on average, atomic C:N:P ratios in both the soil (186:13:1) and the soil microbial biomass (60:7:1) are well-constrained at the global scale. We did see significant variation in soil and microbial element ratios between vegetation types (i.e., forest versus grassland), but in most cases, the similarities in soil and microbial element ratios among sites and across large scales were more apparent than the differences. Consistent microbial biomass element ratios, combined with data linking specific patterns of microbial element stoichiometry with direct evidence of microbial nutrient limitation, suggest that measuring the proportions of C, N and P in the microbial biomass may represent another useful tool for assessing nutrient limitation of ecosystem processes in terrestrial ecosystems.     

Identification of General Patterns of Nutrient and Labile Carbon Control on Soil Carbon Dynamics Across a Successional Gradient

Carbon (C) inputs and nutrient availability are known to affect soil organic carbon (SOC) stocks. However, general rules regarding the operation of these factors across a range of soil nutrient availabilities and substrate qualities are unidentified. “Priming” (stimulated decomposition by labile C inputs) and ‘preferential substrate utilization’ (retarded decomposition due to shifts in community composition towards microbes that do not mineralize SOC) are two hypotheses to explain effects of labile C additions on SOC dynamics. For effects of nutrient additions (nitrogen and phosphorus) on SOC dynamics, the stoichiometric (faster decomposition of materials of low carbon-to-nutrient ratios) and ‘microbial mining’ (that is, reduced breakdown of recalcitrant C forms for nutrients under fertile conditions) hypotheses have been proposed. Using the natural gradient of soil nutrient availability and substrate quality of a chronosequence, combined with labile C and nutrient amendments, we explored the support for these contrasting hypotheses. Additions of labile C, nitrogen (N), phosphorus (P), and combinations of C and N and C and P were applied to three sites: 2-year fallow grassland, mature grassland and forest, and the effects of site and nutrient additions on litter decomposition and soil C dynamics were assessed. The response to C addition supported the preferential substrate hypothesis for easily degradable litter C and the priming hypothesis for SOC, but only in nitrogen-enriched soils of the forest site. Responses to N addition supported the microbial mining hypothesis irrespective of C substrate (litter or SOC), but only in the forest site. Further, P addition effects on SOC support the stoichiometric hypothesis; P availability appeared key to soil C release (priming) in the forest site if labile C and N is available. These results clearly link previously contrasting hypotheses of the factors controlling SOC with the natural gradient in litter quality and nutrient availability that exists in ecosystems at different successional stages. A holistic theory that incorporates this variability of responses, due to different mechanisms, depending on nutrient availability and substrate quality is essential for devising management strategies to safeguard soil C stocks.     

Plant phenology and life span influence soil pool dynamics: Bromus tectorum invasion of perennial C3–C4 grass communities

In water-limited ecosystems, small rainfall events can have dramatic impacts on microbial activity and soil nutrient pools. Plant community phenology and life span also affect soil resources by determining the timing and quantity of plant nutrient uptake, storage, and release. Using the replacement of C₃–C₄ perennial grasses by the invasive annual grass Bromus tectorum as a case study, we investigated the influence of phenology and life span on pulse responses and sizes of soil carbon (C) and nitrogen (N) pools. We hypothesized that available and microbial C and N would respond to small rainfall events and that B. tectorum invasion would increase soil C and N pools by reducing inter-annual plant C and N storage and alter seasonal pool dynamics by changing the timing of plant uptake and litter inputs. We tested our hypotheses by simulating small rainfall events in B. tectorum and perennial grass communities three times during the growing season. Microbial pools responded strongly to soil moisture and simulated rainfall events, but labile C and N pools were affected weakly or not at all. All pools were larger beneath B. tectorum than perennial grasses. Soil C and N pools increased after senescence in both communities. Our results suggest that transforming a perennial into a B. tectorum dominated community increases the overall size of soil C and N pools by decreasing plant C and N storage and changes seasonal pool dynamics by altering dominant plant phenology. Our results indicate strong roles for water, life span and phenology in controlling soil C and N pools and begin to elucidate the biogeochemical effects of altering plant community phenology and life span.     

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.     

The biogeochemical consequences of litter transformation by insect herbivory in the Subarctic: a microcosm simulation experiment

Warming may increase the extent and intensity of insect defoliations within Arctic ecosystems. A thorough understanding of the implications of this for litter decomposition is essential to make predictions of soil-atmosphere carbon (C) feedbacks. Soil nitrogen (N) and C cycles naturally are interlinked, but we lack a detailed understanding of how insect herbivores impact these cycles. In a laboratory microcosm study, we investigated the growth responses of heterotrophic soil fungi and bacteria as well as C and N mineralisation to simulated defoliator outbreaks (frass addition), long-term increased insect herbivory (litter addition at higher background N-level) and non-outbreak conditions (litter addition only) in soils from a Subarctic birch forest. Larger amounts of the added organic matter were mineralised in the outbreak simulations compared to a normal year; yet, the fungal and bacterial growth rates and biomass were not significantly different. In the simulation of long-term increased herbivory, less litter C was respired per unit mineralised N (C:N of mineralisation decreased to 20?±?1 from 38?±?3 for pure litter), which suggests a directed microbial mining for N-rich substrates. This was accompanied by higher fungal dominance relative to bacteria and lower total microbial biomass. In conclusion, while a higher fraction of foliar C will be respired by insects and microbes during outbreak years, predicted long-term increases in herbivory linked to climate change may facilitate soil C-accumulation, as less foliar C is respired per unit mineralised N. Further work elucidating animal-plant-soil interactions is needed to improve model predictions of C-sink capacity in high latitude forest ecosystems.     

Microbial biomass C,N and P in two arctic soils and responses to addition of NPK fertilizer and sugar: implications for plant nutrient uptake

The soil microbial carbon (C), nitrogen (N) and phosphorus (P) pools were quantified in the organic horizon of soils from an arctic/alpine low-altitude heath and a high-altitude fellfield by the fumigation-extraction method before and after factorial addition of sugar, NPK fertilizer and benomyl, a fungicide. In unamended soil, microbial C, N and P made up 3.3–3.6%, 6.1–7.3% and 34.7% of the total soil C, N and P content, respectively. The inorganic extractable N pool was below 0.1% and the inorganic extractable P content slightly less than 1% of the total soil pool sizes. Benomyl addition in spring and summer did not affect microbial C or nutrient content analysed in the autumn. Sugar amendments increased microbial C by 15 and 37% in the two soils, respectively, but did not affect the microbial nutrient content, whereas inorganic N and P either declined significantly or tended to decline. The increased microbial C indicates that the microbial biomass also increased but without a proportional enhancement of N and P uptake. NPK addition did not affect the amount of microbial C but almost doubled the microbial N pool and more than doubled the P pool. A separate study has shown that CO₂ evolution increased by more than 50% after sugar amendment and by about 30% after NPK and NK additions to one of the soils. Hence, the microbial biomass did not increase in response to NPK addition, but the microbes immobilized large amounts of the added nutrients and, judging by the increased CO₂ evolution, their activity increased. We conclude: (1) that microbial biomass production in these soils is stimulated by labile carbon and that the microbial activity is stimulated by both labile C and by nutrients (N); (2) that the microbial biomass is a strong sink for nutrients and that the microbial community probably can withdraw substantial amounts of nutrients from the inorganic, plant-available pool, at least periodically; (3) that temporary declines in microbial populations are likely to release a flush of inorganic nutrients to the soil, particularly P of which the microbial biomass contained more than one third of the total soil pool; and (4) that the mobilization-immobilization cycles of nutrients coupled to the population dynamics of soil organisms can be a significant regulating factor for the nutrient supply to the primary producers, which are usually strongly nutrient-limited in arctic ecosystems.     

氮磷添加和栽植密度对大叶相思林土壤微生物群落功能多样性的影响

土壤微生物群落是土壤质量潜在的生态指标,在维持生态系统功能和服务中起着关键的作用。采用Biolog-ECO微平板法,探讨氮磷添加与不同栽植密度交互对大叶相思林(Acacia auriculiformis)土壤微生物功能多样性的影响,以便为建立合理的林分密度和氮磷施肥模式,提高土壤质量提供理论参考。以大叶相思林为研究对象,选择氯化铵(NH4Cl)作为氮肥模拟大气氮沉降,用二水合磷酸二氢钠(NaH2PO4·2H2O)进行磷添加。氮磷处理设置4个水平,即CK、施N、施P和施N+P。种植密度设置4个水平,即1667、2500、4444、10000株/hm²(分别以低密度、中密度、较高密度、高密度表示)。研究结果表明,施P显著提高了4个密度大叶相思林土壤微生物的AWCD值、碳源利用丰富度指数、McIntosh指数、Shannon指数和Simpson指数,而施N和N+P则相反。随着林分密度的减小,各处理的土壤微生物AWCD值趋于减少。通常低密度林分的土壤微生物的丰富度指数、Shannon-Wiener指数、McIntosh指数和Simpson指数较小。     

Annual burning of a tallgrass prairie inhibits C and N cycling in soil,increasing recalcitrant pyrogenic organic matter storage while reducing N availability

Grassland ecosystems store an estimated 30% of the world's total soil C and are frequently disturbed by wildfires or fire management. Aboveground litter decomposition is one of the main processes that form soil organic matter (SOM). However, during a fire biomass is removed or partially combusted and litter inputs to the soil are substituted with inputs of pyrogenic organic matter (py‐OM). Py‐OM accounts for a more recalcitrant plant input to SOM than fresh litter, and the historical frequency of burning may alter C and N retention of both fresh litter and py‐OM inputs to the soil. We compared the fate of these two forms of plant material by incubating ¹³C‐ and ¹⁵N‐labeled Andropogon gerardii litter and py‐OM at both an annually burned and an infrequently burned tallgrass prairie site for 11 months. We traced litter and py‐OM C and N into uncomplexed and organo‐mineral SOM fractions and CO₂ fluxes and determined how fire history affects the fate of these two forms of aboveground biomass. Evidence from CO₂ fluxes and SOM C:N ratios indicates that the litter was microbially transformed during decomposition while, besides an initial labile fraction, py‐OM added to SOM largely untransformed by soil microbes. Additionally, at the N‐limited annually burned site, litter N was tightly conserved. Together, these results demonstrate how, although py‐OM may contribute to C and N sequestration in the soil due to its resistance to microbial degradation, a long history of annual removal of fresh litter and input of py‐OM infers N limitation due to the inhibition of microbial decomposition of aboveground plant inputs to the soil. These results provide new insight into how fire may impact plant inputs to the soil, and the effects of py‐OM on SOM formation and ecosystem C and N cycling.     

Microbial C, N and P in soils of Mediterranean oak forests: influence of season, canopy cover and soil depth

In Mediterranean ecosystems the effect of aboveground and belowground environmental factors on soil microbial biomass and nutrient immobilization-release cycles may be conditioned by the distinctive seasonal pattern of the Mediterranean-type climates. We studied the effects of season, canopy cover and soil depth on microbial C, N and P in soils of two Mediterranean forests using the fumigation-extraction procedure. Average microbial values recorded were 820 μg C g^?1, 115 μg N g^?1 and 19 μg P g^?1, which accounted for 2.7, 4.7 and 8.8% of the total pools in the surface soil, respectively. Microbial N and P pools were about 10 times higher than the inorganic N and P fractions available for plants. Microbial C values differed between forest sites but in each site they were similar across seasons. Both microbial and inorganic N and P showed maximum values in spring and minimum values in summer, which were positively correlated with soil moisture. Significant differences in soil microbial properties among canopy cover types were observed in the surface soil but only under favourable environmental conditions (spring) and not during summer. Soil depth affected microbial contents which decreased twofold from surface to subsurface soil. Microbial nutrient ratios (C/N, C/P and N/P) varied with seasons and soil depth. Soil moisture regime, which was intimately related to seasonality, emerged as a potential key factor for microbial biomass growth in the studied forests. Our research shows that under a Mediterranean-type climate the interaction among season, vegetation type and structure and soil properties affect microbial nutrient immobilization and thus could influence the biogeochemical cycles of C, N and P in Mediterranean forest ecosystems.     

全球森林土壤微生物生物量碳氮磷化学计量的季节动态

土壤微生物生物量在森林生态系统中充当具有生物活性的养分积累和储存库。土壤微生物转化有机质为植物提供可利用养分, 与植物的相互作用维系着陆地生态系统的生态功能。同时, 土壤微生物也与植物争夺营养元素, 在季节交替过程和植物的生长周期中呈现出复杂的互利-竞争关系。综合全球数据对温带、亚热带和热带森林土壤微生物生物量碳(C)、氮(N)、磷(P)含量及其化学计量比值的季节动态进行分析, 发现温带和亚热带森林的土壤微生物生物量C、N、P含量均呈现夏季低、冬季高的格局。热带森林四季的土壤微生物生物量C、N、P含量都低于温带和亚热带森林, 且热带森林土壤微生物生物量C含量、N含量在秋季相对最低, 土壤微生物生物量P含量四季都相对恒定。温带森林的土壤微生物生物量C:N在春季显著高于其他两个森林类型; 热带森林的土壤微生物生物量C:N在秋季显著高于其他2个森林类型。温带森林土壤微生物生物量N:P和C:P在四季都保持相对恒定, 而热带森林土壤微生物生物量N:P和C:P在夏季高于其他3个季节。阔叶树的土壤微生物生物量C含量、N含量、N:P、C:P在四季都显著高于针叶树; 而针叶树的土壤微生物生物量P含量在四季都显著高于阔叶树。在春季和冬季时, 土壤微生物生物量C:N在阔叶树和针叶树之间都没有显著差异; 但是在夏季和秋季, 针叶树的土壤微生物生物量C:N显著高于阔叶树。对于土壤微生物生物量的变化来说, 森林类型是主要的显著影响因子, 季节不是显著影响因子, 暗示土壤微生物生物量的季节波动是随着植物其内在固有的周期变化而变化。植物和土壤微生物密切作用表现出来的对养分的不同步吸收是保留养分和维持生态功能的一种权衡机制。