Nitrogen Translocation to Fresh Litter in Northern Hardwood Forest

Nitrogen immobilization in fresh litter represents a significant N flux in forest ecosystems, and changes in this process resulting from atmospheric N deposition could have important implications for ecosystem responses. We conducted two leaf decay experiments, using ¹⁵N-labeled sugar maple leaf litter, to quantify N transport from old litter and soil to fresh litter during early stages of decomposition, and we examined the influence of litter N concentration and soil N availability on upward N transfer in a northern hardwood forest. After one year of decay, the average N transfer from soil to fresh litter (2.63 mg N g^?1 litter) was much higher than the N transfer from older litter (1- to 2-years-old) to fresh litter (0.37 mg N g^?1 litter). We calculated the ratio of annual N transfer per unit of excess ¹⁵N pool for these two N sources. The ratio was not significantly different between old litter and soil, suggesting that fungi utilize N in the old litter and mineral soil pools for transport to decaying fresh litter with similar efficiency. Initial litter N concentration had a significant effect on upward N flux into decaying leaf litter, whereas no effect of soil N fertilization was observed. Reduction in the flux from soil to fresh litter owing to anthropogenic N inputs probably contributes significantly to changing soil N dynamics. Future work is needed on fungal N acquisition and transport as well as the fungal taxa involved in this process and their responses to changing environments.     

Climate Variation and Soil Carbon and Nitrogen Cycling Processes in a Northern Hardwood Forest

We exploited the natural climate gradient in the northern hardwood forest at the Hubbard Brook Experimental Forest (HBEF) to evaluate the effects of climate variation similar to what is predicted to occur with global warming over the next 50–100 years for northeastern North America on soil carbon (C) and nitrogen (N) cycle processes. Our objectives were to (1) characterize differences in soil temperature, moisture and frost associated with elevation at the HBEF and (2) evaluate variation in total soil (TSR) and microbial respiration, N mineralization, nitrification, denitrification, nitrous oxide (N₂O) flux, and methane (CH₄) uptake along this gradient. Low elevation sites were consistently warmer (1.5–2.5°C) and drier than high elevation sites. Despite higher temperatures, low elevation plots had less snow and more soil frost than high elevation plots. Net N mineralization and nitrification were slower in warmer, low elevation plots, in both summer and winter. In summer, this pattern was driven by lower soil moisture in warmer soils and in winter the pattern was linked to less snow and more soil freezing in warmer soils. These data suggest that N cycling and supply to plants in northern hardwood ecosystems will be reduced in a warmer climate due to changes in both winter and summer conditions. TSR was consistently faster in the warmer, low elevation plots. N cycling processes appeared to be more sensitive to variation in soil moisture induced by climate variation, whereas C cycling processes appeared to be more strongly influenced by temperature.     

Extracellular Enzyme Activities and Soil Organic Matter Dynamics for Northern Hardwood Forests receiving Simulated Nitrogen Deposition

Anthropogenic nitrogen enrichment alters decomposition processes that control the flux of carbon (C) and nitrogen (N) from soil organic matter (SOM) pools. To link N-driven changes in SOM to microbial responses, we measured the potential activity of several extracellular enzymes involved in SOM degradation at nine experimental sites located in northern Michigan. Each site has three treatment plots (ambient, +30 and +80 kg N ha⁻¹ y⁻¹). Litter and soil samples were collected on five dates over the third growing season of N treatment. Phenol oxidase, peroxidase and cellobiohydrolase activities showed significant responses to N additions. In the Acer saccharum–Tilia americana ecosystem, oxidative activity was 38% higher in the litter horizon of high N treatment plots, relative to ambient plots, while oxidative activity in mineral soil showed little change. In the A. saccharum–Quercus rubra and Q. velutina–Q. alba ecosystems, oxidative activities declined in both litter (15 and 23%, respectively) and soil (29 and 38%, respectively) in response to high N treatment while cellobiohydrolase activity increased (6 and 39% for litter, 29 and 18% for soil, respectively). Over 3 years, SOM content in the high N plots has decreased in the Acer–Tilia ecosystem and increased in the two Quercus ecosystems, relative to ambient plots. For all three ecosystems, differences in SOM content in relation to N treatment were directly related (r² = 0.92) to an enzyme activity factor that included both oxidative and hydrolytic enzyme responses.     

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.     

Skid Trail Effects on Soil Methane and Carbon Dioxide Flux in a Selection-Managed Northern Hardwood Forest

Ecosystems - Temperate forest soils are an important sink for methane (CH4); however, disturbance through forest management and the creation of skid trails may significantly decrease soil’s...     

Calcium Additions and Microbial Nitrogen Cycle Processes in a Northern Hardwood Forest

Evaluating, and possibly ameliorating, the effects of base cation depletion in forest soils caused by acid deposition is an important topic in the northeastern United States. We added 850 kg Ca ha⁻¹ as wollastonite (CaSiO₃) to an 11.8-ha watershed at the Hubbard Brook Experimental Forest (HBEF), a northern hardwood forest in New Hampshire, USA, in fall 1999 to replace calcium (Ca) leached from the ecosystem by acid deposition over the past 6 decades. Soil microbial biomass carbon (C) and nitrogen (N) concentrations, gross and potential net N mineralization and nitrification rates, soil solution and stream chemistry, soil:atmosphere trace gas (CO₂, N₂O, CH₄) fluxes, and foliar N concentrations have been monitored in the treated watershed and in reference areas at the HBEF before and since the Ca addition. We expected that rates of microbial C and N cycle processes would increase in response to the treatment. By 2000, soil pH was increased by a full unit in the Oie soil horizon, and by 2002 it was increased by nearly 0.5 units in the Oa soil horizon. However, there were declines in the N content of the microbial biomass, potential net and gross N mineralization rates, and soil inorganic N pools in the Oie horizon of the treated watershed. Stream, soil solution, and foliar concentrations of N showed no response to treatment. The lack of stimulation of N cycling by Ca addition suggests that microbes may not be stimulated by increased pH and Ca levels in the naturally acidic soils at the HBEF, or that other factors (for example, phosphorus, or Ca binding of labile organic matter) may constrain the capacity of microbes to respond to increased pH in the treated watershed. Possible fates for the approximately 10 kg N ha⁻¹ decline in microbial and soil inorganic pools include components of the plant community that we did not measure (for example, seedlings, understory shrubs), increased fluxes of N₂ and/or N storage in soil organic matter. These results raise questions about the factors regulating microbial biomass and activity in northern hardwood forests that should be considered in the context of proposals to mitigate the depletion of nutrient cations in soil.     

Effects of Calcium on the Rate and Extent of Litter Decomposition in a Northern Hardwood Forest

Cross-site syntheses of litter decomposition studies have shown that litter calcium (Ca) concentration may have a role in controlling the extent of decomposition of tree foliage. We used an ongoing watershed CaSiO₃ addition experiment at the Hubbard Brook Experimental Forest in New Hampshire, USA, to test the hypotheses that increased Ca in litter would have no effect on the initial rates of litter decay but would increase the extent or completeness (limit value) of foliar litter decomposition. We tested these hypotheses with a 6-year litter decomposition experiment using foliar litter of four tree species that are prominent at this site and in the Northern Hardwood forest type of North America: sugar maple (Acer saccharum Marsh), American beech (Fagus grandifolia Ehrh.), yellow birch (Betula alleghaniensis Britt.), and white ash (Fraxinus americana L.). The experiment used a reciprocal transplant design with the Ca-treated watershed and a control site providing two sources of litter and two placement sites. The litter from the Ca-treated site was 10–92% higher in Ca concentration, depending on species, than the litter from the control site. After about 3 years of decomposition, the Ca concentrations in the litter reflected the placement of the litter (that is, the site in which it was incubated) rather than the source of the litter. The source of the litter had no significant effect on measures of initial decomposition rate, cumulative mass loss (6 years), or limit value. However, the placement of the litter had a highly significant effect on extent of decomposition. Some litter types responded more than others; in particular, beech litter placed in the Ca-treated site had a significantly higher limit value, indicating more complete decomposition, and maple litter in the Ca-treated site had a marginally higher limit value. These results indicate that Ca may influence the extent of litter decomposition, but it is the Ca at the incubation site rather than the initial litter Ca that matters most. The results also suggest that loss of Ca from the soil due to decades of acid deposition at this site may have impeded late-stage litter decomposition, possibly leading to greater soil C storage, especially in forest stands with a substantial component of beech. Likewise, de-acidification may lead to a reduction in soil C.     

Soil Fauna Affects Dissolved Carbon and Nitrogen in Foliar Litter in Alpine Forest and Alpine Meadow

Dissolved organic carbon (DOC) and total dissolved nitrogen (TDN) are generally considered important active biogeochemical pools of total carbon and nitrogen. Many studies have documented the contributions of soil fauna to litter decomposition, but the effects of the soil fauna on labile substances (i.e., DOC and TDN) in litter during early decomposition are not completely clear. Therefore, a field litterbag experiment was carried out from 13th November 2013 to 23rd October 2014 in an alpine forest and an alpine meadow located on the eastern Tibetan Plateau. Litterbags with different mesh sizes were used to provide access to or prohibit the access of the soil fauna, and the concentrations of DOC and TDN in the foliar litter were measured during the winter (the onset of freezing, deep freezing and thawing stage) and the growing season (early and late). After one year of field incubation, the concentration of DOC in the litter significantly decreased, whereas the TDN concentration in the litter increased. Similar dynamic patterns were detected under the effects of the soil fauna on both DOC and TDN in the litter between the alpine forest and the alpine meadow. The soil fauna showed greater positive effects on decreasing DOC concentration in the litter in the winter than in the growing season. In contrast, the dynamics of TND in the litter were related to seasonal changes in environmental factors, rather than the soil fauna. In addition, the soil fauna promoted a decrease in litter DOC/TDN ratio in both the alpine forest and the alpine meadow throughout the first year of decomposition, except for in the late growing season. These results suggest that the soil fauna can promote decreases in DOC and TDN concentrations in litter, contributing to early litter decomposition in these cold biomes.     

Initial Soil Organic Matter Content Influences the Storage and Turnover of Litter,Root and Soil Carbon in Grasslands

Grassland degradation is a worldwide problem that often leads to substantial loss of soil organic matter (SOM). To estimate the potential for carbon (C) accumulation in degraded grassland soils, we first need to understand how SOM content influences the transformation of plant C and its stabilization within the soil matrix. We conducted a greenhouse experiment using C₃ soils with six levels of SOM content; we planted the C₄ grass Cleistogenes squarrosa or added its litter to the soils to investigate how SOM content regulates the storage of new soil C derived from litter and roots, the decomposition of extant soil C, and the formation of soil aggregates. We found that with the increase in SOM content, microbial biomass carbon (MBC) and the mineralization of litter C increased. Both the litter addition and planted treatments increased the amount of new C inputs to soil. However, the mineralization of extant soil C was significantly accelerated by the presence of living roots but was not affected by litter addition. Accordingly, the soil C content was significantly higher in the litter addition treatments but was not affected by the planted treatments by the end of the experiment. The soil macroaggregate fraction increased with SOM content and was positively related to MBC. Our experiment suggests that as SOM content increases, plant growth and soil microbial activity increase, which allows microbes to process more plant-derived C and promote new soil C formation. Although long-term field experiments are needed to test the robustness of our findings, our greenhouse experiment suggests that the interactions between SOM content and plant C inputs should be considered when evaluating soil C turnover in degraded grasslands.     

Nitrogen Relations in a Forest Plantation--Soil Organic Matter Ecosystem Model

Two previously published models, after minor modification, areamalgamated to give a model that describes the major carbonand nitrogen pools and fluxes in a plantation forest soil system.The first model is a transport-resistance model of forest growthand dry-matter partitioning. The second is a soil organic mattermodel that was constructed for temperate grasslands. The combinedmodel is used to examine the relations between plantation growth,soil organic matter content, nitrogen deposition rate from theatmosphere, mineralization flux, nitrogen uptake by the plantation,dry matter partitioning between foliage and root, litter productionand the timing and quantity of fertilizer application. The highdemand for N by even-aged plantations during the period of canopybuilding is highlighted. The marked ontogenetic shifts in thegrowth pattern during plantation development is emphasized,indicating several phases of forest development. The resultsindicate that the potential growth of even-aged plantationsmay be greater than that realized in poor soils with commonlevels of atmospheric N deposition and normal fertilizer regimes.The simulations show how the concentrations of soil mineralN change during the development of a plantation, and point towardsthe importance of atmospheric N deposition. They also show thatfertilizer application must be accurately matched to growthstage if fertilizer is to be used efficiently. The nitrogencycle (N-uptake by plant     

Interactions between Carbon and Nitrogen Mineralization and Soil Organic Matter Chemistry in Arctic Tundra Soils

We used long-term laboratory incubations and chemical fractionation to characterize the mineralization dynamics of organic soils from tussock, shrub, and wet meadow tundra communities, to determine the relationship between soil organic matter (SOM) decomposition and chemistry, and to quantify the relative proportions of carbon (C) and nitrogen (N) in tundra SOM that are biologically available for decomposition. In all soils but shrub, we found little decline in respiration rates over 1 year, although soils respired approximately a tenth to a third of total soil C. The lack of decline in respiration rates despite large C losses indicates that the quantity of organic matter available was not controlling respiration and thus suggests that something else was limiting microbial activity. To determine the nature of the respired C, we analyzed soil chemistry before and after the incubation using a peat fractionation scheme. Despite the large losses of soil C, SOM chemistry was relatively unchanged after the incubation. The decomposition dynamics we observed suggest that tundra SOM, which is largely plant detritus, fits within existing concepts of the litter decay continuum. The lack of changes in organic matter chemistry indicates that this material had already decomposed to the point where the breakdown of labile constituents was tied to lignin decomposition. N mineralization was correlated with C mineralization in our study, but shrub soil mineralized more and tussock soil less N than would have been predicted by this correlation. Our results suggest that a large proportion of tundra SOM is potentially mineralizable, despite the fact that decomposition was dependent on lignin breakdown, and that the historical accumulation of organic matter in tundra soils is the result of field conditions unfavorable to decomposition and not the result of fundamental chemical limitations to decomposition. Our study also suggests that the anticipated increases in shrub dominance may substantially alter the dynamics of SOM decomposition in the tundra. Received 31 January 2002; accepted 16 July 2002.     

Dynamics of Organic Matter Decomposition and Microflora Composition of Forest Litter in Artificial Biogeocenoses

The dynamics of litter stock, microbial biomass, and composition and structure of microbial communities, were studied in the course of soil organic matter transformation during vegetation season. The dynamics of litter stock in coniferous and deciduous forests proved to correlate with the biomass and total abundance of microorganisms, particularly, with the proportion of microfungi in the microbial community.     

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。     

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.     

Effect of Pasture Trees on Soil Nitrogen and Organic Matter: Implications for Tropical Montane Forest Restoration

In lower-montane ecosystems of Ecuador, Setaria sphacelata (foxtail grass), the predominant introduced pasture species, forms a tussock grassland that reduces soil nitrogen and resists recolonization of forest vegetation. We compared the influence of individual trees or small clusters of nitrogen-fixing ( Inga sp., Fabaceae) and non-nitrogen-fixing trees ( Psidium guajava L., guava) on the soil and abiotic conditions that affect further regeneration of forest vegetation within pastures. Pasture trees ameliorated air temperature and light intensity to levels similar to those in adjacent intact forest. Beneath Inga , soil NO₃ ⁻ -N was four times higher than in open pasture. Nitrification was five times higher under Inga canopies than in open pastures for both field and laboratory incubations. This suggests that the increased soil N transformations under Inga are derived mainly from improved soil rather than microenvironmental conditions. Psidium canopies slightly increased field nitrification but had no effect under laboratory conditions. We also compared the natural abundance ¹³C signature and the carbon and nitrogen content of subcanopy soil with adjacent open pasture soil. Inga increased the C and N content of the upper 5 cm of soil and increased by 7% the fraction of soil organic matter derived from C₃ plants. The improved soil and abiotic conditions beneath the canopies of N-fixing pasture trees favor the establishment and growth of woody montane species, suggesting that these trees could be used to accelerate forest regeneration within abandoned pastures.     

Soil Moisture Alters the Response of Soil Organic Carbon Mineralization to Litter Addition

Increasing rainfall and longer drought conditions lead to frequent changes in soil moisture that affect soil organic carbon (SOC) mineralization. However, how soil moisture affects response of SOC mineralization to litter addition in forest ecosystems remains unexplored. We added ¹³C-labeled litter to subtropical forest soils with three mass water contents (L, 21%; M, 33%; H, 45%). Carbon dioxide production was monitored, and the composition of soil microbial communities was determined by phospholipid fatty acid (PLFA). When no litter was added, SOC mineralization was greater in the M-treated soil. Litter addition promoted SOC mineralization, but this promotion was altered by soil moisture and litter type. Priming effects induced by P. massoniana leaf litter in the M-moistened soil were significantly (P < 0.05) higher than those in other treatments. Litter-derived C was approximately 55% incorporated into 18:1ω9c and 16:0 PLFAs, and this proportion was not significantly affected by soil moisture. Soil moisture affected the distribution of litter-¹³C in i15:0, i17:0, and cy19:0 individual PLFAs. The primed C evolution was significantly related to the ratio of Gram-positive to Gram-negative bacteria. These results suggest that changes in soil moisture could affect SOC mineralization in forest ecosystems.     

模拟氮沉降对典型阔叶红松林土壤有机碳和养分的影响简

为探讨氮沉降对典型阔叶红松（Pinus koraiensis）林的影响,从2008年6月~2010年8月进行了人工模拟氮沉降实验,实验分为对照、低N、中N、高N4个处理,每个处理3个重复。所施氮肥为CO(NH₂)₂,以溶液的形式喷施,4个处理浓度分别为0、30、60、120 kg·hm^-2·a^-1。在氮沉降进行1年后,采集各处理0～20、20～40和40～60 cm的土壤样品,测定其土壤有机C、全N、碱解N和速效P、速效K。结果表明：相同处理下,有机C和全N含量随土层的加深均逐渐减少。总体上低、中N处理显著增加了土壤有机C、碱解N和速效K含量,中、高N处理显著降低了土壤速效P含量（P<0.05）,而对全N含量影响不显著（P>0.05）。土壤有机C与土壤全N、碱解N、速效P、速效K之间存在极显著正相关关系（P<0.001）。有机C和土壤养分对氮沉降的响应说明氮沉降在短期内可能影响阔叶红松林土壤碳库积累和土壤肥力水平。     

Watershed-Level Responses to Calcium Silicate Treatment in a Northern Hardwood Forest

Watershed 1 (W1) at the Hubbard Brook Experimental Forest in New Hampshire, with chronically low pH and acid neutralizing capacity (ANC) in surface water, was experimentally treated with calcium silicate (CaSiO₃; wollastonite) in October 1999 to assess the role of calcium (Ca) supply in the structure and function of base-poor forest ecosystems. Wollastonite addition significantly increased the concentrations and fluxes of Ca, dissolved silica (Si), and ANC and decreased the concentrations and fluxes of inorganic monomeric Al (Al_i) and hydrogen ion (H⁺) in both soil solution and stream water in all sub-watersheds of W1. Mass balances indicate that 54% of the added Ca remained undissolved or was retained by vegetation during the first 6 years after treatment. Of the remaining added Ca, 44% was retained on O horizon cation exchange sites. The Ca:Si ratio in the dissolution products was greater than 2.0, more than twice the molar ratio in the applied wollastonite. This suggests that Ca was preferentially leached from the applied wollastonite and/or Si was immobilized by secondary mineral formation. Approximately 2% of the added Ca and 7% of the added Si were exported from W1 in streamwater in the first 6 years after treatment. Watershed-scale Ca amendment with wollastonite appears to be an effective approach to mitigating effects of acidic deposition. Not only does it appear to alleviate acidification stress to forest vegetation, but it also provides for the long-term supply of ANC to acid-impacted rivers and lakes downstream.