Response of Oxidative Enzyme Activities to Nitrogen Deposition Affects Soil Concentrations of Dissolved Organic Carbon

Recent evidence suggests that atmospheric nitrate (NO ₃ ⁻ ) deposition can alter soil carbon (C) storage by directly affecting the activity of lignin-degrading soil fungi. In a laboratory experiment, we studied the direct influence of increasing soil NO ₃ ⁻ concentration on microbial C cycling in three different ecosystems: black oak–white oak (BOWO), sugar maple–red oak (SMRO), and sugar maple–basswood (SMBW). These ecosystems span a broad range of litter biochemistry and recalcitrance; the BOWO ecosystem contains the highest litter lignin content, SMRO had intermediate lignin content, and SMBW leaf litter has the lowest lignin content. We hypothesized that increasing soil solution NO ₃ ⁻ would reduce lignolytic activity in the BOWO ecosystem, due to a high abundance of white-rot fungi and lignin-rich leaf litter. Due to the low lignin content of litter in the SMBW, we further reasoned that the NO ₃ ⁻ repression of lignolytic activity would be less dramatic due to a lower relative abundance of white-rot basidiomycetes; the response in the SMRO ecosystem should be intermediate. We increased soil solution NO ₃ ⁻ concentrations in a 73-day laboratory incubation and measured microbial respiration and soil solution dissolved organic carbon (DOC) and phenolics concentrations. At the end of the incubation, we measured the activity of β-glucosidase, N-acetyl-glucosaminidase, phenol oxidase, and peroxidase, which are extracellular enzymes involved with cellulose and lignin degradation. We quantified the fungal biomass, and we also used fungal ribosomal intergenic spacer analysis (RISA) to gain insight into fungal community composition. In the BOWO ecosystem, increasing NO ₃ ⁻ significantly decreased oxidative enzyme activities (−30% to −54%) and increased DOC (+32% upper limit) and phenolic (+77% upper limit) concentrations. In the SMRO ecosystem, we observed a significant decrease in phenol oxidase activity (−73% lower limit) and an increase in soluble phenolic concentrations (+57% upper limit) in response to increasing NO ₃ ⁻ in soil solution, but there was no significant change in DOC concentration. In contrast to these patterns, increasing soil solution NO ₃ ⁻ in the SMBW soil resulted in significantly greater phenol oxidase activity (+700% upper limit) and a trend toward lower DOC production (−52% lower limit). Nitrate concentration had no effect on microbial respiration or β-glucosidase or N-acetyl-glucosaminidase activities. Fungal abundance and basidiomycete diversity tended to be highest in the BOWO soil and lowest in the SMBW, but neither displayed a consistent response to NO ₃ ⁻ additions. Taken together, our results demonstrate that oxidative enzyme production by microbial communities responds directly to NO ₃ ⁻ deposition, controlling extracellular enzyme activity and DOC flux. The regulation of oxidative enzymes by different microbial communities in response to NO ₃ ⁻ deposition highlights the fact that the composition and function of soil microbial communities directly control ecosystem-level responses to environmental change.     

高寒草甸植物碳氮组成及其稳定同位素特征

采用稳定同位素质谱仪Isoprime100,对采自黄河源区典型高寒草甸和人工改良草地的主要植物进行了碳、氮组成及其稳定同位素丰富度测定,判断植物光合类型,探讨稳定碳氮同位素丰富度对草地植被演替的响应。结果表明:(1)研究区58种主要植物碳元素含量在28.64%~51.55%之间,氮元素含量介于0.89%~4.04%,δ13 C值变化范围介于-29.50‰~-24.69‰,δ15 N值介于-4.57‰~8.32‰。(2)不同样地植物碳含量的大小顺序为人工草地(45.54%)未退化草甸(43.18%)轻度退化草甸(42.18%)严重退化草甸(39.68%),氮元素含量顺序为未退化草甸(2.30%)人工草地(2.28%)轻度退化草甸(2.13%)严重退化草甸(2.10%),表明草甸退化会引起植物碳氮含量的降低。(3)未退化草甸、人工草地、轻度退化草甸和严重退化草甸的δ13 C值依次为-25.63‰、-26.57‰、-26.76‰和-27.91‰,δ15 N值依次为-0.63‰、0.32‰、2.76‰和0.26‰。研究认为,黄河源区高寒草甸和人工改良草地的58种主要植物均属C3植物,没有发现C4和景天酸代谢(CAM)植物,低的年均气温可能是制约该区C4植物分布的主要因素;植物δ13 C值随草地退化程度加剧而逐渐降低,但δ15 N值的变化无规律性趋势。     

Influence of Litter Diversity on Dissolved Organic Matter Release and Soil Carbon Formation in a Mixed Beech Forest

We investigated the effect of leaf litter on below ground carbon export and soil carbon formation in order to understand how litter diversity affects carbon cycling in forest ecosystems. ¹³C labeled and unlabeled leaf litter of beech (Fagus sylvatica) and ash (Fraxinus excelsior), characterized by low and high decomposability, were used in a litter exchange experiment in the Hainich National Park (Thuringia, Germany). Litter was added in pure and mixed treatments with either beech or ash labeled with ¹³C. We collected soil water in 5 cm mineral soil depth below each treatment biweekly and determined dissolved organic carbon (DOC), δ¹³C values and anion contents. In addition, we measured carbon concentrations and δ¹³C values in the organic and mineral soil (collected in 1 cm increments) up to 5 cm soil depth at the end of the experiment. Litter-derived C contributes less than 1% to dissolved organic matter (DOM) collected in 5 cm mineral soil depth. Better decomposable ash litter released significantly more (0.50±0.17%) litter carbon than beech litter (0.17±0.07%). All soil layers held in total around 30% of litter-derived carbon, indicating the large retention potential of litter-derived C in the top soil. Interestingly, in mixed (ash and beech litter) treatments we did not find a higher contribution of better decomposable ash-derived carbon in DOM, O horizon or mineral soil. This suggest that the known selective decomposition of better decomposable litter by soil fauna has no or only minor effects on the release and formation of litter-derived DOM and soil organic matter. Overall our experiment showed that 1) litter-derived carbon is of low importance for dissolved organic carbon release and 2) litter of higher decomposability is faster decomposed, but litter diversity does not influence the carbon flow.     

Transport of Carbon and Nitrogen Between Litter and Soil Organic Matter in a Northern Hardwood Forest

We used sugar maple litter double-labeled with ¹³C and ¹⁵N to quantify fluxes of carbon (C) and nitrogen (N) between litter and soil in a northern hardwood forest and the retention of litter C and N in soil. Two cohorts of litter were compared, one in which the label was preferentially incorporated into non-structural tissue and the other structural tissue. Loss of ¹³C from this litter generally followed dry mass and total C loss whereas loss of ¹⁵N (20–30% in 1 year) was accompanied by large increases of total N content of this decaying litter (26–32%). Enrichment of ¹³C and ¹⁵N was detected in soil down to 10–15 cm depth. After 6 months of decay (November–May) 36–43% of the ¹³C released from the litter was recovered in the soil, with no differences between the structural and non-structural labeled litter. By October the percentage recovery of litter ¹³C in soil was much lower (16%). The C released from litter and remaining in soil organic matter (SOM) after 1 year represented over 30 g C m⁻² y⁻¹ of SOM accumulation. Recovery of litter ¹⁵N in soil was much higher than for C (over 90%) and in May ¹⁵N was mostly in organic horizons whereas by October it was mostly in 0–10 cm mineral soil. A small proportion of this N was recovered as inorganic N (2–6%). Recovery of ¹⁵N in microbial biomass was higher in May (13–15%) than in October (about 5%). The C:N ratio of the SOM and microbial biomass derived from the labeled litter was much higher for the structural than the non-structural litter and for the forest floor than mineral SOM, illustrating the interactive role of substrates and microbial activity in regulating the C:N stoichiometry of forest SOM formation. These results for a forest ecosystem long exposed to chronically high atmospheric N deposition (ca. 10 kg N ha⁻¹ y⁻¹) suggest possible mechanisms of N retention in soil: increased organic N leaching from fresh litter and reduced fungal transport of N from soil to decaying litter may promote N stabilization in mineral SOM even at a relatively low C:N ratio.     

Patterns of Carbon, Nitrogen and Phosphorus Dynamics in Decomposing Foliar Litter in Canadian Forests

We examined the patterns of nitrogen (N) and phosphorus (P) gain, retention or loss in ten foliar tissues in a litterbag experiment over 6 years at 18 upland forest sites in Canada, ranging from subarctic to cool temperate. N was usually retained in the decomposing litter until about 50% of the original C remained. The peak N content in the litter was observed at between 72 and 99% of the original C remaining, with C:N mass quotients between 37 and 71 (mean 55). The rate of N release from the litters was not related to the original N concentration, which may be associated with the generally narrow range (0.59–1.28% N) in the litters. P was immediately lost from all litters, except beech leaves, with critical litter C:P mass quotients for P release being in the range 700–900. The rate of P loss was inversely correlated with the original litter P concentration, which ranged from 0.02 to 0.13%. The soil underlying the litterbags influenced the pattern of N and P dynamics in the litters; there were weak correlations between the N and P remaining at 60% C remaining in the litters and the C:N and C:P quotients of the surface layer of the soil. There was a trend for higher N and P retention in the litter at sites with lower soil C:N and N:P quotients, respectively. Although there was a large variation in C:N, C:P and N:P quotients in the original litters (29–83, 369–2122 and 5–26, respectively), and some variation in the retention or loss of N and P in the early stages of decomposition, litters converged on C:N, C:P and N:P quotients of 30, 450 and 16, when the C remaining fell below 30%. These quotients are similar to that found in the surface organic matter of these ecosystems.     

Phosphorus Limits Tropical Rain Forest Litter Fauna

The stoichiometry of resources may explain bottom-up regulation of higher trophic levels. We tested the effects of soil and litter nutrient stoichiometry on the invertebrate litter fauna of a Costa Rican tropical rain forest. Animal densities were estimated from 15 sites across a phosphorus gradient. The density of the invertebrate litter fauna varied considerably, and was strongly tied to soil and litter phosphorus concentrations. An increase in phosphorus concentrations corresponded with an equally proportionate increase in animal densities. Natural variation in nutrient levels can thus serve as a predictor of density in a highly diverse tropical animal community.     

The Role of Dissolved Organic Carbon, Dissolved Organic Nitrogen, and Dissolved Inorganic Nitrogen in a Tropical Wet Forest Ecosystem

Although tropical wet forests play an important role in the global carbon (C) and nitrogen (N) cycles, little is known about the origin, composition, and fate of dissolved organic C (DOC) and N (DON) in these ecosystems. We quantified and characterized fluxes of DOC, DON, and dissolved inorganic N (DIN) in throughfall, litter leachate, and soil solution of an old-growth tropical wet forest to assess their contribution to C stabilization (DOC) and to N export (DON and DIN) from this ecosystem. We found that the forest canopy was a major source of DOC (232 kg C ha^–1 y^–1). Dissolved organic C fluxes decreased with soil depth from 277 kg C ha^–1 y^–1 below the litter layer to around 50 kg C kg C ha^–1 y^–1 between 0.75 and 3.5m depth. Laboratory experiments to quantify biodegradable DOC and DON and to estimate the DOC sorption capacity of the soil, combined with chemical analyses of DOC, revealed that sorption was the dominant process controlling the observed DOC profiles in the soil. This sorption of DOC by the soil matrix has probably led to large soil organic C stores, especially below the rooting zone. Dissolved N fluxes in all strata were dominated by mineral N (mainly NO₃⁻). The dominance of NO₃^– relative to the total amount nitrate of N leaching from the soil shows that NO₃^– is dominant not only in forest ecosystems receiving large anthropogenic nitrogen inputs but also in this old-growth forest ecosystem, which is not N-limited.     

天坑森林植物叶-凋落物-土壤C、N、P生态化学计量特征

为探究广西乐业大石围天坑森林群落的C、N、P养分循环特征,比较了天坑内外森林群落的植物叶片-凋落物-土壤C、N、P含量及其化学计量比,采用相关性分析和冗余分析等统计方法研究其内在联系和相互影响。结果表明,与天坑外部森林相比,天坑内部森林植物叶片和凋落物呈现出C低N、P高,土壤为C、N低P高的格局。植物叶片C:N、C:P与凋落物C、N:P显著正相关,植物叶片C与土壤P显著负相关;天坑外部森林的植物叶片N、N:P与土壤N:P显著负相关,植物叶片C:N与土壤C、C:N显著正相关,说明天坑森林内部凋落物的C、P养分可能主要来源于植物叶片,而天坑外部森林的植物叶片C、N主要来自土壤。土壤C:N:P对植物叶、凋落物的C:N:P变化的解释率分别为90.7%和50.6%,其中土壤P对植物叶和凋落物的C:N:P计量特征变化的解释度最高,坑内生境植物对P含量变化更为敏感、坑外植物对于N含量变化更为敏感,表明天坑内部森林可能是P素受限位点、天坑外部森林是N素受限位点。喀斯特天坑内部森林和外部森林植物叶-凋落物-土壤的C:N:P的差异和联系,体现了天坑内外森林群落的养分循环特征和植物群落的适应性。     

Forest Gaps Inhibit Foliar Litter Pb and Cd Release in Winter and Inhibit Pb and Cd Accumulation in Growing Season in an Alpine Forest

Aims

The release of heavy metals (such as Pb and Cd) from foliar litter play an important role in element cycling in alpine forest ecosystems. Although natural forest gaps could play important roles in the release of heavy metals from foliar litter by affecting the snow cover during the winter and solar irradiation during the growing season, few studies have examined these potential roles. The objectives of this study were to document changes in Pb and Cd dynamics during litter decomposition in the center of gaps and under closed canopies and to investigate the factors that controlled these changes during the winter and growing seasons.

Methods

Senesced foliar litter from six dominant species, including Kangding willow (Salix paraplesia), Masters larch (Larix mastersiana), Mingjiang fir (Abies faxoniana), Alpine azalea (Rhododendron lapponicum), Red birch (Betula albosinensis) and Mourning cypress (Sabina saltuaria), was placed in litterbags and incubated between the gap center and closed canopy conditions in an alpine forest in the eastern region of the Tibetan Plateau. The litterbags were sampled at the snow formation stage, snow coverage stage, snow melt stage and during the growing season. The Pb and Cd concentrations in the sampled foliar litter were determined by acid digestion (HNO₃/HClO₄).

Important findings

Over one year of decomposition, Pb accumulation and Cd release from the foliar litter occurred, regardless of the foliar litter species. However, Pb and Cd were both released from the foliar litter during the winter and accumulated during the growing season. Compared with the gap center and the canopy gap edge, the extended gap edge and the closed canopy showed higher Pb and Cd release rates in winter and higher Pb and Cd accumulation rates during the growing season, respectively. Statistical analyses indicate that the dynamics of Pb were significantly influenced by frequent freeze–thaw cycles in winter and appropriate hydrothermal conditions during the growing season, the dynamics of Cd were strongly influenced by species and the presence of a forest gap at different decomposition stages. These results show that forest gaps could inhibit Pb and Cd release from foliar litter in the alpine forest of western Sichuan. In addition, a decrease in the snow depth in the winter warming scenario would promote the release of Pb during foliar litter decomposition. There exist some difference that may be influenced by litter quality, microenvironment and microtopography during litter decomposition.     

Warming and Nitrogen Addition Increase Litter Decomposition in a Temperate Meadow Ecosystem

Background

Litter decomposition greatly influences soil structure, nutrient content and carbon sequestration, but how litter decomposition is affected by climate change is still not well understood.

Methodology/Principal Findings

A field experiment with increased temperature and nitrogen (N) addition was established in April 2007 to examine the effects of experimental warming, N addition and their interaction on litter decomposition in a temperate meadow steppe in northeastern China. Warming, N addition and warming plus N addition reduced the residual mass of L. chinensis litter by 3.78%, 7.51% and 4.53%, respectively, in 2008 and 2009, and by 4.73%, 24.08% and 16.1%, respectively, in 2010. Warming, N addition and warming plus N addition had no effect on the decomposition of P. communis litter in 2008 or 2009, but reduced the residual litter mass by 5.58%, 15.53% and 5.17%, respectively, in 2010. Warming and N addition reduced the cellulose percentage of L. chinensis and P. communis, specifically in 2010. The lignin percentage of L. chinensis and P. communis was reduced by warming but increased by N addition. The C, N and P contents of L. chinensis and P. communis litter increased with time. Warming and N addition reduced the C content and C:N ratios of L. chinensisand P. communis litter, but increased the N and P contents. Significant interactive effects of warming and N addition on litter decomposition were observed (P<0.01).

Conclusion/Significance

The litter decomposition rate was highly correlated with soil temperature, soil water content and litter quality. Warming and N addition significantly impacted the litter decomposition rate in the Songnen meadow ecosystem, and the effects of warming and N addition on litter decomposition were also influenced by the quality of litter. These results highlight how climate change could alter grassland ecosystem carbon, nitrogen and phosphorus contents in soil by influencing litter decomposition.     

Interactions between Litter Lignin and Nitrogenitter Lignin and Soil Nitrogen Availability during Leaf Litter Decomposition in a Hawaiian Montane Forest

Previous work in a young Hawaiian forest has shown that nitrogen (N) limits aboveground net primary production (ANPP) more strongly than it does decomposition, despite low soil N availability. In this study, I determined whether (a) poor litter C quality (that is, high litter lignin) poses an overriding constraint on decomposition, preventing decomposers from responding to added N, or (b) high N levels inhibit lignin degradation, lessening the effects of added N on decomposition overall. I obtained leaf litter from one species, Metrosideros polymorpha, which dominates a range of sites in the Hawaiian Islands and whose litter lignin concentration declines with decreasing precipitation. Litter from three dry sites had lignin concentrations of 12% or less, whereas litter from two wet sites, including the study site, had lignin concentrations of more than 18%. This litter was deployed 2.5 years in a common site in control plots (receiving no added nutrients) and in N-fertilized plots. Nitrogen fertilization stimulated decomposition of the low-lignin litter types more than that of the high-lignin litter types. However, in contrast to results from temperate forests, N did not inhibit lignin decomposition. Rather, lignin decay increased with added N, suggesting that the small effect of N on decomposition at this site results from limitation of decomposition by poor C quality rather than from N inhibition of lignin decay. Even though ANPP is limited by N, decomposers are strongly limited by C quality. My results suggest that anthropogenic N deposition may increase leaf litter decomposition more in ecosystems characterized by low-lignin litter than in those characterized by high-lignin litter. Received 26 October 1999; accepted 2 June 2000.     

Lignin Degradation in Foliar Litter of Two Shrub Species from the Gap Center to the Closed Canopy in an Alpine Fir Forest

To understand the effects of forest gaps on lignin degradation during shrub foliar litter decomposition, a field litterbag experiment was conducted in an alpine fir (Abies faxoniana) forest of the eastern Tibet Plateau. Dwarf bamboo (Fargesia nitida) and willow (Salix paraplesia) foliar litterbags were placed on the forest floor from the gap center to the closed canopy. The litterbags were sampled during snow formation, snow coverage, snow melting and the growing season from October 2010 to October 2012. The lignin concentrations and loss in the litter were measured. Over 2 years, lignin loss was lower in the bamboo litter (34.64–43.89%) than in the willow litter (38.91–55.10%). In the bamboo litter, lignin loss mainly occurred during the first decomposition year, whereas it occurred during the second decomposition year in the willow litter. Both bamboo and willow litter lignin loss decreased from the gap center to the closed canopy during the first year and over the entire 2-year decomposition period. Compared with the closed canopy, the gap center showed higher lignin loss for both bamboo and willow litter during the two winters, but lower lignin loss during the early growing period. Additionally, the dynamics of microbial biomass carbon during litter decomposition followed the same trend as litter lignin loss during the two winters and growing period. These results indicated that alpine forest gaps had significant effects on shrub litter lignin loss and that reduced snow cover during winter warming would inhibit shrub lignin degradation in this alpine forest.     

高寒草甸矮蒿草种群繁殖对策的研究

繁殖对策是指生物对环境的生殖适应趋势 ,是资源或能量向生存、生长和生殖等活动中最适分配的结果 ,在不同的环境中具有其独特的表现形式。研究植物在不同环境中的繁殖对策可以反映出植物对环境的适应能力和在该生境中的生殖潜能。国内外学者对植物繁殖对策的研究已有不少报道[2 ,4 ,5,6] 。但对高寒草甸矮嵩草 (Ｋｏｂｒｅｓｉａｈｕｍｉｌｉｓ)种群繁殖对策的研究报道甚少。矮嵩草是青藏高原矮嵩草草甸的建群种 ,它具有草质柔软、营养丰富、热值含量较高等特点 ,是青藏高原重要的可更新草地资源。本研究对矮嵩草的繁殖对策进行了较全面、…     

海北高寒草甸土壤有机碳同位素组成及C3/C4碳源的变化

通过对高寒嵩草草甸土壤剖面不同深度(0～5cm,5～15cm,15～25cm,25～35cm,35～50cm,50～65cm)有机碳稳定性碳同位素的测定发现,土壤有机碳稳定性同位素(δ^13C)随土壤深度的增加而变大。表层土壤(0～5cm,定义为现代土壤)的δ^13C值最小,基本上接近现代植被的碳同位素特征。在土层5～10cm深度以下(粗略地定为古土壤),土壤有机碳稳定性同位素骤然上升,与表层土壤的同位素特征明显不同。考虑到影响土壤碳同位素的诸多因素,通过稳定性碳同位素的质量平衡模型计算,得出初步结果：来自C4(或CAM)植物的碳源随土壤深度的增加而增大。进一步推测,该地区植被可能经历由C4植物占优势的群落向C3植物占优势的群溶演化的过程。在这个过程中,大气碳同位素的变化和土壤有机质的形成过程(有机质淋溶过程)等也会引起土壤碳同位素的升高,因此质量平衡模型可能会过多地估算C4组分,而低估C3组分。     

青海高寒草甸土壤放线菌区系研究

2001～2002年从海北高寒草甸生态系统采集土样,用不同方法从中分离放线菌300余株,根据其形态和分类特征,分别归入小单孢菌属(Micromonospora)、诺卡氏菌属(Nocardia)、糖多孢菌属(Saccharopolyspora)、原小单孢菌属(Promicromonospora)和链霉菌属(Streptomyces),并将链霉菌归入7个类群。同时对230株中温菌和110株低温菌的部分酶活性及其对真菌和细菌的拮抗性进行了测定,发现链霉菌不仅具有许多酶活性,而且对真菌和细菌有拮抗性。     

Forest Gaps Alter the Total Phenol Dynamics in Decomposing Litter in an Alpine Fir Forest

The total phenol content in decomposing litter not only acts as a crucial litter quality indicator, but is also closely related to litter humification due to its tight absorption to clay particles. However, limited attention has been focused on the total phenol dynamics in foliar litter in relation to forest gaps. Here, the foliar litter of six representative tree species was incubated on the forest floor from the gap center to the closed canopy of an alpine Minjiang fir (Abies faxoniana) forest in the upper reaches of the Yangtze River and eastern Tibetan Plateau. The dynamics of total phenol concentration in the incubated litter was measured from November 2012 to October 2014. Over two-year incubation, 78.22% to 94.06% of total phenols were lost from the foliar litter, but 52.08% to 86.41% of this occurred in the first year. Forest gaps accelerated the loss of total phenols in the foliar litter in the winter, although they inhibited the loss of total phenols during the growing season in the first year. In comparison with the effects of forest gaps, the variations of litter quality among different species were much stronger on the dynamics of total phenols in the second year. Overall, the loss of total phenols in the foliar litter was slightly higher in both the canopy gap and the expanded gap than in the gap center and under the closed canopy. The results suggest that the predicted decline in snow cover resulting from winter warming or vanishing gaps caused by forest regeneration will retard the loss of total phenol content in the foliar litter of alpine forest ecosystems, especially in the first decomposition year.     

青藏高原高寒草甸生态系统碳通量特征及其控制因子

青藏高原特殊的气候变化特征,在区域生态系统碳动态平衡起着重要作用。本文就青藏高原不同生态系统CO2通量的时空变化及其影响因子和控制途径进行综述,以明确青藏高原植草地生态系统的碳源汇状况及时空变化特征,为全球变化的研究提供依据。     

Original Articles: Differential Influence of Plant Species on Soil Nitrogen Transformations Within Moist Meadow Alpine Tundra

Plant species can influence nitrogen (N) cycling indirectly through the feedbacks of litter quality and quantity on soil N transformation rates. The goal of this research was to focus on small-scale (within-community) variation in soil N cycling associated with two community dominants of the moist meadow alpine tundra. Within this community, the small-scale patchiness of the two most abundant species (Acomastylis rossii and Deschampsia caespitosa) provides natural variation in species cover within a relatively similar microclimate, thus enabling estimation of the effects of plant species on soil N transformation rates. Monthly rates of soil N transformations were dependent on small-scale variation in both soil microclimate and species cover. The relative importance of species cover compared with soil microclimate increased for months 2 and 3 of the 3-month growing season. Growing-season net N mineralization rates were over ten times greater and nitrification rates were four times greater in Deschampsia patches than in Acomastylis patches. Variability in litter quality [carbon:nitrogen (C:N) and phenolic:N], litter quantity (aboveground and fine-root production), and soil quality (C:N) was associated with three principal components. Variability between the species in litter quality and fine-root production explained 31% of the variation in net N mineralization rates and 36% of net nitrification rates. Site variability across the landscape in aboveground production and soil C:N explained 33% of the variation in net N mineralization rates and 21% of net nitrification rates. Within the moist meadow community, the high spatial variability in soil N transformation rates was associated with differences in the dominant species' litter quality and fine-root production. Deschampsia-dominated patches consistently had greater soil N transformation rates than did Acomastylis-dominated patches across the landscape, despite site variability in soil moisture, soil C:N, and aboveground production. Plant species appear to be an important control of soil N transformation in the alpine tundra, and consequently may influence plant community structure and ecosystem function.     

Leaf Litter as a Source of Dissolved Organic Carbon in Streams

Dissolved organic carbon (DOC) is an abundant form of organic matter in stream ecosystems. Most research has focused on the watershed as the source of DOC in streams, but DOC also comes from leaching of organic matter stored in the stream channel. We used a whole-ecosystem experimental approach to assess the significance of leaching of organic matter in the channel as a source of DOC in a headwater stream. Inputs of leaf litter were excluded from a forested Appalachian headwater stream for 3 years. Stream-water concentration, export, and instream generation of DOC were reduced in the litter-excluded stream as compared with a nearby untreated reference stream. The proportion of high molecular weight (HMW) DOC (more than 10,000 daltons) in stream water was not altered by litter exclusion. Mean DOC concentration in stream water was directly related to benthic leaf-litter standing stock. Instream generation of DOC from leaf litter stored in the stream channel contributes approximately 30% of daily DOC exports in this forested headwater stream. This source of DOC is greatest during autumn and winter and least during spring and summer. It is higher during increasing discharge than during base flow. We conclude that elimination of litter inputs from a forested headwater stream has altered the biogeochemistry of DOC in this ecosystem. Received 2 September 1997; accepted 27 January 1998.