Influence of gap size and soil properties on microbial biomass in a subtropical humid forest of north-east India

We examined the effects of treefall gap size and soil properties on microbial biomass dynamics in an undisturbed mature-phase humid subtropical broadleaved forest in north-east India. Canopy gaps had low soil moisture and low microbial biomass suggesting that belowground dynamics accompanied changes in light resources after canopy opening. High rainfall in the region causes excessive erosion/leaching of top soil and eventually soil fertility declines in treefall gaps compared to understorey. Soil microbial population was less during periods when temperature and moisture conditions are low, while it peaked during rainy season when the litter decomposition rate is at its peak on the forest floor. Greater demand for nutrients by plants during rainy season (the peak vegetative growth period) limited the availability of nutrients to soil microbes and, therefore, low microbial C, N and P. Weak correlations were also obtained for the relationships between microbial C, N and P and soil physico–chemical properties. Gap size did influence the microbial nutrients and their contribution to soil organic carbon, total Kjeldhal nitrogen and available-P. Contribution of microbial C to soil organic carbon, microbial N to total nitrogen were similar in both treefall gaps and understorey plots, while the contribution of microbial P to soil available-P was lower in gap compared to the understorey. These results indicate that any fluctuation in microbial biomass related nutrient cycling processes in conjunction with the associated microclimate variation may affect the pattern of regeneration of tree seedlings in the gaps and hence be related with their size. This revised version was published online in June 2006 with corrections to the Cover Date.     

Response of soil microbial communities to changes in a forest ecosystem brought about by pine wilt disease

Japan has suffered a lot from forestry losses due to pine wilt disease caused by pinewood nematode infestations. Studies were conducted regarding its causative agent and the effects of natural vegetation succession after pine wilt disease, but its effects on microorganisms were not given equal attention. This study determined the effects of pine wilt disease on light conditions, soil microbial biomass, litter decomposition, microbial abundance and the physical and chemical properties of the soil. Results showed that in a forest currently affected by pine wilt disease, there was higher light penetration, greater microbial biomass carbon, and a faster rate of litter decomposition. Microbial abundance was shown to be reduced in pine wilt affected areas. There were close correlations between the biological and physicochemical properties of the soil, but the reason for the decrease in microbial abundance is not yet well understood, and thus requires further study.     

人为干扰对鼎湖山马尾松林土壤细根和有机质的影响

通过处理 (根据当地习惯收割凋落物和林下层 )和保护 (无任何人为干扰 )样地的比较试验 ,1990～ 1995年期间研究了人为干扰对鼎湖山生物圈保护区马尾松 (Pinus massoniana)林土壤细根和有机质的影响。在此 5 a的研究期间 ,由于人为干扰活动而直接从处理样地取走的林下层和凋落物总量为 2 1.7t/ hm2。在保护样地 ,林下层生物量从 2 .2 t/ hm2增加至 11.10 t/ hm2 ,地表凋落物 (包括枯死的林下层 )量则从 3.0 t/ hm2 增加至 13.3t/ hm2 。收割林下层和凋落物这种人为干扰活动对林地土壤细根生物量的影响不明显 ,但却显著降低土壤轻腐殖质 (Soil lightorganic matter)量。在细根分解过程中 ,其分解速率在处理样地(试验结束时细根残存量占起始量的 4 0 .8% )显著高于在保护样地 (试验结束时细根残存量占起始量的 4 4 .3% ) ;与 Ca、Mg和K元素不同 ,N和 P两种元素的释放速率在处理样地显著高于保护样地 ,表明这种人为干扰活动不仅直接取走所收割的林下层和凋落物中的养分 ,而且还可能增加林地有效养分的流失潜力     

Effects of mixing pine and broadleaved tree/shrub litter on decomposition and N dynamics in laboratory microcosms

This study was carried out to compare the ecological function of exotic pine (Pinus radiata—Pr) and native pine (Pinus tabulaeformis—Pt) in terms of litter decomposition and its related N dynamics and to evaluate if the presence of broad-leaved tree species (Cercidiphyllum japonicum—Cj) or shrub species (Ostryopsis davidiana—Od) litter would promote the decomposition of pine needles and N cycling. Mass remaining, N release of the four single-species litters and mixed-species (Pt + Cj; Pr + Cj; Pt + Od; Pr + Od) litters and soil N dynamics were measured at microcosm scale during an 84-day incubation period. The Pt and Pr litter, with poorer substrate quality, indicated slower decomposition rates than did the Cj and Od litter. Due to their high C/N ratios, the N mass of Pt and Pr litter continuously increased during the early stage of decomposition, which showed that Pt and Pr litter immobilized exogenous N by microbes. No significant differences of soil inorganic, dissolved organic and microbial biomass N were found between the Pt and Pr microcosm at each sampling. The results showed that the exotic Pr performed similar ecological function to the native Pt in terms of litter decomposition and N dynamics during the early stage. The presence of Cj or Od litter increased the decomposition rates of pine needle litter and also dramatically increased soil N availability. So it is feasible for plantation managers to consider the use of Cj as an ameliorative species or to retain Od in pine plantations to promote the decomposition of pine litter and increase nutrient circulation. The results also suggested that different species litters induced different soil dissolved organic nitrogen (DON). As a major soluble N pool in soil, DON developed a different changing tendency over time compared with inorganic N, and should be included into soil N dynamic under the condition of our study.     

高山森林三种细根分解初期微生物生物量动态

为了解川西高山森林凋落物分解过程的微生物生物量特征,采用凋落物分解袋法,测定了粗枝云杉(Picea asperata)、岷江冷杉(Abies faxoniana)和红桦(Betula albosinensi)细根分解几个关键时期微生物生物量碳(MBC)、氮(MBN)和磷(MBP)的动态特征。3个树种细根分解过程中的MBC均表现为在土壤深冻期下降至全年最低点后缓慢上升,至土壤融冻中期再次下降,到生长季节增长的趋势。然而,粗枝云杉与岷江冷杉细根分解过程中的MBC最大值出现在生长季节末期,红桦细根分解过程中的MBC最大值出现在土壤冻结初期。3个树种细根分解过程中的MBN表现出相似的动态规律:土壤深冻期急剧下降至全年最低,随后在冻融季节无显著变化,生长季节明显增加,到生长季节末期达到全年最大值。另外,粗枝云杉和岷江冷杉细根分解过程中MBP均随着分解的进行呈现增加趋势,而红桦细根分解过程中的MBP在土壤融冻末期出现最大值,在生长季节中期出现另一峰值,生长季节末期明显下降。这些结果表明冬季细根分解过程中仍存在一定的土壤微生物,但受到细根质量、温度及其驱动的环境因子的深刻影响。     

Effects of nitrogen addition on litter decomposition, soil microbial biomass, and enzyme activities between leguminous and non-leguminous forests

Anthropogenic nitrogen (N) deposition is an expanding problem that affects the functioning and composition of forest ecosystems, particularly the decomposition of forest litters. Legumes play an important role in the nitrogen cycle of forest ecosystems. Two litter types were chosen from Zijin Mountain in China: Robinia pseudoacacia leaves from a leguminous forest (LF) and Liquidambar formosana leaves from a non-leguminous forest (NF). The litter samples were mixed into original forest soils and incubated in microcosms. Then, they were treated by five forms of N addition: NH₄ ⁺, NO₃ ^?, urea, glycine, and a mixture of all four. During a 6-month incubation period, litter mass losses, soil microbial biomass, soil pH, and enzyme activities were investigated. Results showed that mixed N and NO₃ ^?-N addition significantly accelerated the litter decomposition rates of LF leaves, while mixed N, glycine-N, and urea-N addition significantly accelerated the litter decomposition rates of NF leaves. Litter decomposition rates and soil enzyme activities under mixed N addition were higher than those under single form of N additions in the two forest types. Nitrogen addition had no significant effects on soil pH and soil microbial biomass. The results indicate that nitrogen addition may alter microbial allocation to extracellular enzyme production without affecting soil microbial biomass, and then affected litter decomposition process. The results further reveal that mixed N is a more important factor in controlling litter decomposition process than single form of N, and may seriously affect soil N cycle and the release of carbon stored belowground.     

Litter and soil biodiversity jointly drive ecosystem functions

The decomposition of litter and the supply of nutrients into and from the soil are two fundamental processes through which the above- and belowground world interact. Microbial biodiversity, and especially that of decomposers, plays a key role in these processes by helping litter decomposition. Yet the relative contribution of litter diversity and soil biodiversity in supporting multiple ecosystem services remains virtually unknown. Here we conducted a mesocosm experiment where leaf litter and soil biodiversity were manipulated to investigate their influence on plant productivity, litter decomposition, soil respiration, and enzymatic activity in the littersphere. We showed that both leaf litter diversity and soil microbial diversity (richness and community composition) independently contributed to explain multiple ecosystem functions. Fungal saprobes community composition was especially important for supporting ecosystem multifunctionality (EMF), plant production, litter decomposition, and activity of soil phosphatase when compared with bacteria or other fungal functional groups and litter species richness. Moreover, leaf litter diversity and soil microbial diversity exerted previously undescribed and significantly interactive effects on EMF and multiple individual ecosystem functions, such as litter decomposition and plant production. Together, our work provides experimental evidence supporting the independent and interactive roles of litter and belowground soil biodiversity to maintain ecosystem functions and multiple services.     

小麦残茬落叶的分解与土壤因子间动态关系的研究

东北高寒地区的黑钙土土质优良肥沃 ,适合小麦、大豆和玉米等种植。近年来 ,由于人们只重视无机化肥的使用 ,忽视了地力培育 ,大量秸秆被移出田外 ,造成土壤有机质含量降低 ,土壤板结 ,使原本高产的农田逐渐变成中低产田 ,甚至有的已成为撂荒地。因此 ,研究当前农田土壤对枯枝落叶的分解现状 ,对于认识现有耕种条件下 ,农田土壤亚系统的物质转化和能量流动具有实际意义。1　研究地区和研究方法1 .1　自然概况该研究是在黑龙江省克山师专农场进行的。地理位置位于东经 1 2 5°8′～ 1 2 6°8′,北纬 47°50′～ 48°33′。年均气温 1 .3℃ ,1…     

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.     

Species-dependent responses of soil microbial properties to fresh leaf inputs in a subtropical forest soil in South China

Aims Forest disturbance from extreme weather events due to climate change could increase the contribution of fresh green leaves to the litter layer of soil and subsequently alter the composition and activity of the soil microbial properties and soil carbon cycling. The objective of this study was to compare the effect of naturally fallen litter and fresh leaves on the soil microbial community composition and their activities.Methods Fresh leaves and normal fallen litter were collected from four tree species (Pinus elliottii, Schima superba, Acacia mangium, A. auriculaeformis) in subtropical China and mixed with soil. Soil microbial community composition was determined using PLFAs, and its activity was quantified by soil respiration. During a 12-month period, the decomposition rate of litter was measured bimonthly using a litterbag method. Soil microbial samples were collected after 6 and 12 months. Soil respiration was measured monthly.Important findings We found that fresh leaves decomposed faster than their conspecific fallen litter. Although total microbial biomass and bacterial biomass were similar among treatments, soil fungal biomass was higher in fresh leaf than fallen litter treatments, resulting in greater values of the Fungal phospholipid fatty acids (PLFAs)/Bacterial PLFAs ratio. Fungal PLFA values were greater for Schima superba than the other species. The effect of litter type on soil respiration was species-dependent. Specifically, fallen litter released 35% more CO₂ than fresh leaves of the conifer P. elliottii. The opposite pattern was observed in the broadleaf species whose fresh leaf treatments emitted 17%–32% more CO₂ than fallen litter. Given future predictions that global climate change will cause more disturbances to forests, these results indicate that conifer and broadleaf forests in subtropical China may respond differently to increased fresh litter inputs, with net soil microbial respiration decreasing in conifer forests and increasing in broadleaf forests.     

杉木凋落物对土壤有机碳分解及微生物生物量碳的影响

利用¹³C稳定同位素示踪技术,研究了杉木凋落物对杉木人工林表层(0～5 cm)和深层(40～45 cm)土壤有机碳分解、微生物生物量碳和可溶性碳动态的影响.结果表明: 杉木人工林中深层土壤有机碳分解速率显著低于表层土壤,但其激发效应却显著高于表层土壤.杉木凋落物添加使土壤总微生物生物量碳和源于原有土壤的微生物生物量碳均显著增加,但对土壤可溶性碳没有显著影响.深层土壤被翻到林地表层,可能加速杉木人工林土壤中碳的损失.     

Impact of the plant community composition on labile soil organic carbon, soil microbial activity and community structure in semi-natural grassland ecosystems of different productivity

Aims

The main objective was to describe the effects of plant litter on SOC and on soil microbial activity and structure in extensively managed grasslands in Central Germany that vary in biomass production and plant community composition.

Methods

The decomposition of shoot and root litter was studied in an incubation experiment. Labile C and N were isolated by hot water extraction (C_HWE, N_HWE), while functional groups of microbes were identified by PLFA analysis and microbial activity was measured using a set of soil exo-enzymes.

Results

The plant community composition, particulary legume species affected SOC dynamics and below-ground microbial processes, especially via roots. This was reflected in about 20% lower decomposition of root litter in low productivity grassland soil. The C_HWE soil pool was found to be a key driver of the below-ground food web, controlling soil microbial processes.

Conclusions

Below-ground responses appear to be related to the presence of legume species, which affected the microbial communities, as well as the ratio between fungal and bacterial biomass and patterns of soil enzyme activity. Low productivity fungal-dominated grasslands with slow C turnover rates may play an important role in SOC accumulation. The approach used here is of particular importance, since associated biological and biochemical processes are fundamental to ecosystem functioning.     

人工油松林（Pinus tabulaeformis）恢复过程中土壤微生物生物量C、N的变化特征

采用时空替代法,选取15a（PF15）、25a（PF25）、30a（PF30）的人工油松林作为样地,并选取灌丛作为参考植被,研究了植被恢复过程中土壤微生物生物量C、N以及土壤养分的变化特征,同时探讨了它们之间的相互关系。研究结果表明随着恢复的进行,土壤质量得到了改善,主要表现为有机碳、全氮、粘粒含量、土壤含水量的上升和pH值、容重的下降。土壤微生物生物量C、N分别在155.00~885.64mg/kg和33.73~237.40mg/kg的范围内变化。土壤微生物生物量C、N在植被恢复的初期显著低于灌丛,而后随着恢复的进行逐步增长。土壤微生物生物量C、N与植被恢复时间的相关性没有达到统计学上的显著水平,但是土壤微生物生物量C与土壤有机碳、全氮、全磷呈显著正相关,这表明植被恢复过程中土壤微生物生物量与土壤养分状况关系密切,植被恢复通过改善土壤养分状况间接地影响土壤微生物生物量的变化。Cmic/TOC在1.38%~4.75%的范围内变化。Cmic/TOC随着植被恢复不断下降,Cmic/TOC与植被恢复时间和土壤有机碳呈显著负相关,这表明植被恢复过程中,惰性有机质积累导致供应土壤微生物的活性有机质减少,Cmic/TOC同时受土壤有机质的数量和质量影响。     

Drivers of spatial variability in urban plant and soil isotopic composition in the Los Angeles basin

Aims

Plant litter decomposition plays an important role in the storage of soil organic matter in terrestrial ecosystems. Conversion of native vegetation to agricultural lands and subsequent land abandonment can lead to shifts in canopy structure, and consequently influence decomposition dynamics by alterations in soil temperature and moisture conditions, solar radiation exposure, and soil erosion patterns. This study was conducted to assess which parameters were more closely related to short-term decomposition dynamics of two predominant Mediterranean leaf litter types.

Methods

Using the litterbag technique, we incubated leaf litter of Pinus halepensis and Rosmarinus officinalis in two Mediterranean land-uses with different degree of vegetation cover (open forest, abandoned agricultural field).

Results

Fresh local litter lost between 20 and 55% of its initial mass throughout the 20-month incubation period. Rosemary litter decomposed faster than pine litter, showing net N immobilization in the early stages of decomposition, in contrast to the net N release exhibited by pine litter. Parameters related to litter quality (N content or C:N) or land-use/site conditions (ash content, an index of soil deposition on litter) were found to explain the cross-site variability in mass loss rates for rosemary and Aleppo pine litter, respectively.

Conclusions

The results from this study suggest that decomposition drivers may differ depending on litter type in this Mediterranean ecosystem. While rosemary litter was degraded mainly by microbial activity, decomposition of pine litter was likely driven primarily by abiotic processes like soil erosion.     

冬小麦生境中土壤养分对凋落物碳氮释放的影响

土壤养分影响植物生长, 进而影响凋落物质量和产量; 凋落物质量和产量影响凋落物分解过程。基于一个生长实验和一个相同环境分解实验, 研究了冬小麦(Triticum aestivum)生境中养分可利用性对凋落物碳(C)和氮(N)释放的影响。结果显示: (1)冬小麦凋落物产量、叶/根C:N比、C释放量和N释放量随土壤养分梯度呈单调变化; (2)土壤养分影响叶凋落物丢失率而不影响根凋落物丢失率; (3)初始叶/根C:N比与其C、N释放量之间存在负相关关系; (4)分解过程降低叶C:N比和根C:N比。结果表明: 生境中土壤养分的提高可加速凋落物C、N归还, 这反过来可能促进冬小麦生长, 因此这种效应是正反馈; 初始C:N比可预测凋落物C、N释放量。     

胶州湾滨海湿地枯落物分解对土壤活性有机碳含量及其三维荧光特性的影响

通过室内试验模拟胶州湾滨海湿地碱蓬、芦苇、互花米草枯落物的分解过程,测定土壤活性有机碳(可溶性有机碳、微生物生物量碳)含量变化,并利用三维荧光技术对可溶性有机碳(DOC)进行光谱分析.结果表明: 土壤活性有机碳含量呈先增加后降低最终趋于稳定的变化趋势;不同枯落物类型和添加方式对土壤活性有机碳的影响不同,表现为碱蓬＞互花米草＞芦苇,且原状混合＞表面覆盖;光谱分析表明,分解过程各三维荧光光谱的荧光峰数量、荧光中心位置和荧光强度都存在一定差异,添加枯落物后土壤未发现类酪氨酸峰;芳香类蛋白物质占比最高,腐殖质类物质次之.枯落物分解是以微生物分解作用为主导、枯落物性质为本质要素、多重因子综合作用的结果,其对提高土壤活性有机碳含量、增强土壤碳库的稳定性具有促进作用;该过程通过改变DOC的结构和化学组分,提高其在土壤中的迁移转化能力,增强生物可降解性和可利用性,促进微生物内源DOC的产生,加快湿地土壤碳循环.     

Soil organic matter,nutrient distribution,fungal and microbial biomass and enzyme activities in a forest beech stand on the Apennines of southern Italy

The aim of this study was to analyse the amount and qualitative characteristics of organic matter (OM) in the litter horizon (considering leaf litter at different decomposition stages) and underlying soil to a 30-cm depth in a beech stand on the Apennines in southern Italy. Distribution of major nutrients as well as fungal and microbial biomass were also evaluated, in addition to beech leaf nutrient content monitor from full expansion to abscission in order to estimate annual nutrient input to soil from litterfall and nutrient retranslocation before abscission. OM was significantly higher in leaf litter. C/N ratio and the Na, Mn, Fe levels also decreased along the decomposition continuum, whereas N and S contents slowly decreased with soil depth. Generally, leaf nutrient content was also significantly lower in dead leaves, indicating efficient retranslocation to persistent organs. Fungal biomass was the highest in leaf layers, with no significant changes between spring and autumn samplings. Enzyme activities did not differ significantly along the decomposition continuum but marked decreases were found in the upper soil layer; these remained relatively constant, with the exception of laccase, at deeper soil depths. No seasonal effect on enzyme activities and OM content was found.     

土壤微生物群落对枯落物输入的响应

枯落物分解在陆地生态系统物质循环能量流动中起着关键性作用,明确枯落物输入对土壤微生物群落的影响有助于理解土壤微生物生物多样性和陆地生态系统功能的相互关系。本文采用整合分析方法,以中国为研究区域,以不添加枯落物为对照组,探究土壤微生物(真菌、细菌、放线菌)及微生物生物量碳、生物量氮对枯落物输入的响应。结果表明:与不添加枯落物相比,添加枯落物后土壤微生物生物量碳、生物量氮分别显著增加3.9%和4.4%;土壤真菌PLFA、细菌PLFA及总微生物PLFA分别增加4.0%、3.1%和2.4%。枯落物输入对土壤微生物的影响受到气候条件、年降水量、植被类型及土壤酸碱度等因素的显著影响;不同气候类型下,土壤微生物对枯落物输入的响应呈现出亚热带季风气候区>温带季风气候区>温带大陆气候区的趋势,以及随着年降水量的增加呈现出先升高后降低的趋势;不同植被类型下,土壤微生物对枯落物输入的响应呈现出阔叶林>草地≈混交林>针叶林的趋势。     

Influence of soil depth on the decomposition of Bouteloua gracilis roots in the shortgrass steppe

The distribution and turnover of plant litter contribute to soil structure, the availability of plant nutrients, and regional budgets of greenhouse gasses. Traditionally, studies of decomposition have focused on the upper soil profile. Other work has shown that temperature, precipitation, and soil texture are important determinates of patterns of decomposition. Since these factors all vary through a soil profile, it has been suggested that decomposition rates may vary with depth in a soil profile. In this work, we examine patterns of root decomposition through a shortgrass steppe soil profile. We buried fresh root litter from Bouteloua gracilis plants in litterbags at 10, 40, 70, and 100 cm. Litterbags were retrieved six times between July 1996 and May 1999. We found that the decomposition rate for fresh root litter was approximately 50% slower at 1 m than it was at 10 cm. After 33 months, 55% of the root mass buried at 10 cm remained, while 72% of the root mass buried at 1 m was still present. This corresponds to a 19-year residence time for roots at 10 cm and a 36-year residence time for roots at 1 m. Mass loss rates decreased linearly from 10 cm to 1 m. Patterns of total carbon and cellulose loss rates followed those of mass loss rates. Roots at 1 m tended to accumulate lignin-like compounds over the course of the experiment. Differences in the stabilization of lignin may be a consequence of differences in microbial community through a shortgrass steppe soil profile.     

Winter climate change implications for decomposition in northeastern forests: comparisons of sugar maple litter with herbivore fecal inputs

Forests in northeastern North America are influenced by varying climatic and biotic factors; however, there is concern that rapid changes in these factors may lead to important changes in ecosystem processes such as decomposition. Climate change (especially warming) is predicted to increase rates of decomposition in northern latitudes. Warming in winter may result in complex effects including decreased levels of snow cover and an increased incidence of soil freezing that will effect decomposition. Along with these changes in climate, moose densities have also been increasing in this region, likely affecting nutrient dynamics. We measured decomposition and N release from ¹⁵N‐labeled sugar maple leaf litter and moose feces over 20 months in reference and snow removal treatment (to induce soil freezing) plots in two separate experiments at the Hubbard Brook Experimental Forest in New Hampshire, USA. Snow removal/soil freezing decreased decomposition of maple litter, but stimulated N transfer to soil and microbial biomass. Feces decomposed more rapidly than maple litter, and feces N moved into the mineral soil more than N derived from litter, likely due to the lower C : N ratio of feces. Feces decomposition was not affected by the snow removal treatment. Total microbial biomass (measured as microbial N and C) was not significantly affected by the treatments in either the litter or feces plots. These results suggest that increases in soil freezing and/or large herbivore populations, increase the transfer rate of N from plant detritus or digested plants into the mineral soil. Such changes suggest that altering the spatial and temporal patterns of soil freezing and moose density have important implications for ecosystem N cycling.