Response of Net Ecosystem Productivity of Three Boreal Forest Stands to Drought

In 2000–03, continuous eddy covariance measurements of carbon dioxide (CO₂) flux were made above mature boreal aspen, black spruce, and jack pine forests in Saskatchewan, Canada, prior to and during a 3-year drought. During the 1st drought year, ecosystem respiration (R) was reduced at the aspen site due to the drying of surface soil layers. Gross ecosystem photosynthesis (GEP) increased as a result of a warm spring and a slow decrease of deep soil moisture. These conditions resulted in the highest annual net ecosystem productivity (NEP) in the 9 years of flux measurements at this site. During 2002 and 2003, a reduction of 6% and 34% in NEP, respectively, compared to 2000 was observed as the result of reductions in both R and GEP, indicating a conservative response to the drought. Although the drought affected most of western Canada, there was considerable spatial variability in summer rainfall over the 100-km extent of the study area; summer rainfalls in 2001 and 2002 at the two conifer sites minimized the impact of the drought. In 2003, however, precipitation was similarly low at all three sites. Due to low topographic position and consequent poor drainage at the black spruce site and the coarse soil with low water-holding capacity at the jack pine site almost no reduction in R, GEP, and NEP was observed at these two sites. This study shows that the impact of drought on carbon sequestration by boreal forest ecosystems strongly depends on rainfall distribution, soil characteristics, topography, and the presence of vegetation that is well adapted to these conditions. The online version of the original article can be found under doi:     

Biometric Based Carbon Flux Measurements and Net Ecosystem Production (NEP) in a Temperate Deciduous Broad-Leaved Forest Beneath a Flux Tower

Biometric based carbon flux measurements were conducted over 5 years (1999–2003) in a temperate deciduous broad-leaved forest of the AsiaFlux network to estimate net ecosystem production (NEP). Biometric based NEP, as measured by the balance between net primary production (including NPP of canopy trees and of forest floor dwarf bamboo) and heterotrophic respiration (RH), clarified the contribution of various biological processes to the ecosystem carbon budget, and also showed where and how the forest is storing C. The mean NPP of the trees was 5.4 ± 1.07 t C ha⁻¹ y⁻¹, including biomass increment (0.3 ± 0.82 t C ha⁻¹ y⁻¹), tree mortality (1.0 ± 0.61 t C ha⁻¹ y⁻¹), aboveground detritus production (2.3 ± 0.39 t C ha⁻¹ y⁻¹) and belowground fine root production (1.8 ± 0.31 t C ha⁻¹ y⁻¹). Annual biomass increment was rather small because of high tree mortality during the 5 years. Total NPP at the site was 6.5 ± 1.07 t C ha⁻¹ y⁻¹, including the NPP of the forest floor community (1.1 ± 0.06 t C ha⁻¹ y⁻¹). The soil surface CO₂ efflux (RS) was averaged across the 5 years of record using open-flow chambers. The mean estimated annual RS amounted to 7.1 ± 0.44 t C ha⁻¹, and the decomposition of soil organic matter (SOM) was estimated at 3.9 ± 0.24 t C ha⁻¹. RH was estimated at 4.4 ± 0.32 t C ha⁻¹ y⁻¹, which included decomposition of coarse woody debris. Biometric NEP in the forest was estimated at 2.1 ± 1.15 t C ha⁻¹ y⁻¹, which agreed well with the eddy-covariance based net ecosystem exchange (NEE). The contribution of woody increment (Δbiomass + mortality) of the canopy trees to NEP was rather small, and thus the SOM pool played an important role in carbon storage in the temperate forest. These results suggested that the dense forest floor of dwarf bamboo might have a critical role in soil carbon sequestration in temperate East Asian deciduous forests.     

Soil Respiration and Belowground Carbon Allocation in Mangrove Forests

Mangrove forests cover large areas of tropical and subtropical coastlines. They provide a wide range of ecosystem services that includes carbon storage in above- and below ground biomass and in soils. Carbon dioxide (CO₂) emissions from soil, or soil respiration is important in the global carbon budget and is sensitive to increasing global temperature. To understand the magnitude of mangrove soil respiration and the influence of forest structure and temperature on the variation in mangrove soil respiration I assessed soil respiration at eleven mangrove sites, ranging from latitude 27°N to 37°S. Mangrove soil respiration was similar to those observed for terrestrial forest soils. Soil respiration was correlated with leaf area index (LAI) and aboveground net primary production (litterfall), which should aid scaling up to regional and global estimates of soil respiration. Using a carbon balance model, total belowground carbon allocation (TBCA) per unit litterfall was similar in tall mangrove forests as observed in terrestrial forests, but in scrub mangrove forests TBCA per unit litter fall was greater than in terrestrial forests, suggesting mangroves allocate a large proportion of their fixed carbon below ground under unfavorable environmental conditions. The response of soil respiration to soil temperature was not a linear function of temperature. At temperatures below 26°C Q10 of mangrove soil respiration was 2.6, similar to that reported for terrestrial forest soils. However in scrub forests soil respiration declined with increasing soil temperature, largely because of reduced canopy cover and enhanced activity of photosynthetic benthic microbial communities.     

Net Primary Production and Canopy Nitrogen in a Temperate Forest Landscape: An Analysis Using Imaging Spectroscopy, Modeling and Field Data

Understanding spatial patterns of net primary production (NPP) is central to the study of terrestrial ecosystems, but efforts are frequently hampered by a lack of spatial information regarding factors such as nitrogen availability and site history. Here, we examined the degree to which canopy nitrogen can serve as an indicator of patterns of NPP at the Bartlett Experimental Forest in New Hampshire by linking canopy nitrogen estimates from two high spectral resolution remote sensing instruments with field measurements and an ecosystem model. Predicted NPP across the study area ranged from less than 700 g m⁻² year⁻¹ to greater than 1300 g m⁻² year⁻¹ with a mean of 951 g m⁻² year⁻¹. Spatial patterns corresponded with elevation, species composition and historical forest management, all of which were reflected in patterns of canopy nitrogen. The relationship between production and elevation was nonlinear, with an increase from low- to mid-elevation deciduous stands, followed by a decline in upper-elevation areas dominated by evergreens. This pattern was also evident in field measurements and mirrored an elevational trend in foliar N concentrations. The increase in production from low-to mid-elevation deciduous stands runs counter to the generally accepted pattern for the northeastern U.S. region, and suggests an importance of moisture limitations in lower-elevation forests. Field measurements of foliar N, wood production and leaf litterfall were also used to evaluate sources of error in model estimates and to determine how predictions are affected by different methods of acquiring foliar N input data. The accuracy of predictions generated from remotely sensed foliar N approached that of predictions driven by field-measured foliar N. Predictions based on the more common approach of using aggregated foliar N for individual cover types showed reasonable agreement in terms of the overall mean, but were in poor agreement on a plot-by-plot basis. Collectively, these results suggest that variation in foliar N exerts an important control on landscape-level spatial patterns and can serve as an integrator of other underlying factors that influence forest growth rates.     

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。     

Ectomycorrhizal Colonization, Biomass, and Production in a Regenerating Scrub Oak Forest in Response to Elevated CO2

The effects of CO₂ elevation on the dynamics of fine root (FR) mass and ectomycorrhizal (EM) mass and colonization were studied in situ in a Florida scrub oak system over four years of postfire regeneration. Soil cores were taken at five dates and sorted to assess the standing crop of ectomycorrhizal and fine roots. We used ingrowth bags to estimate the effects of elevated CO₂ on production of EM roots and fine roots. Elevated CO₂ tended to increase EM colonization frequency but did not affect EM mass nor FR mass in soil cores (standing mass). However, elevated CO₂ strongly increased EM mass and FR mass in ingrowth bags (production), but it did not affect the EM colonization frequency therein. An increase in belowground production with unchanged biomass indicates that elevated CO₂ may stimulate root turnover. The CO₂-stimulated increase of belowground production was initially larger than that of aboveground production. The oaks may allocate a larger portion of resources to root/mycorrhizal production in this system in elevated rather than ambient CO₂.     

Using Light-Use and Production Efficiency Models to Predict Photosynthesis and Net Carbon Exchange During Forest Canopy Disturbance

Vegetation growth models are used with remotely sensed and meteorological data to monitor terrestrial carbon dynamics at a range of spatial and temporal scales. Many of these models are based on a light-use efficiency equation and two-component model of whole-plant growth and maintenance respiration that have been parameterized for distinct vegetation types and biomes. This study was designed to assess the robustness of these parameters for predicting interannual plant growth and carbon exchange, and more specifically to address inconsistencies that may arise during forest disturbances and the loss of canopy foliage. A model based on the MODIS MOD17 algorithm was parameterized for a mature upland hardwood forest by inverting CO₂ flux tower observations during years when the canopy was not disturbed. This model was used to make predictions during a year when the canopy was 37% defoliated by forest tent caterpillars. Predictions improved after algorithms were modified to scale for the effects of diffuse radiation and loss of leaf area. Photosynthesis and respiration model parameters were found to be robust at daily and annual time scales regardless of canopy disturbance, and differences between modeled net ecosystem production and tower net ecosystem exchange were only approximately 2 g C m⁻² d⁻¹ and less than 23 g C m⁻² y⁻¹. Canopy disturbance events such as insect defoliations are common in temperate forests of North America, and failure to account for cyclical outbreaks of forest tent caterpillars in this stand could add an uncertainty of approximately 4–13% in long-term predictions of carbon sequestration.     

Different Population Dynamics of Microcebus murinus in Primary and Secondary Deciduous Dry Forests of Madagascar

The goal of this study was to identify causes for lower population densities of mouse lemurs (Microcebus murinus) in secondary than in primary dry deciduous forests of western Madagascar. Variations in the abundance of Microcebus murinus are linked to the capacity to enter energy-saving torpor during the dry season. Under natural conditions in primary forest, Microcebus murinus can maintain daily torpor (and possibly hibernation) as long as body temperatures remain below 28°C. Females are more likely to hibernate than males, resulting in skewed sex ratios of captured Microcebus murinus in the primary forest. In the secondary forest the sex ratio of subjects captured during the dry season is more balanced than in primary forest, indicating that fewer females go into torpor in secondary than in primary forest. Secondary forests have fewer large standing or fallen trees that might provide holes as shelter for Microcebus murinus. Ambient temperatures are higher in secondary than in primary forests and higher outside than inside tree holes. These high ambient temperatures might hinder the ability of Microcebus murinus to maintain torpor for prolonged periods in secondary forests. Mouse lemurs from secondary forest have lower body mass than in primary forest. The year-to-year recapture rate is zero in secondary forest and thus significantly lower than in primary forest. This indicates that survival rates are lower in secondary than in primary forests. Thus, secondary forests may be of limited value as buffer zones or even corridors for mouse lemurs.     

Spatial and Temporal Dynamics of Exotic Earthworm Communities Along Invasion Fronts in a Temperate Hardwood Forest in South-Central New York (USA)

The invasion of North American forests by exotic earthworms is producing profound ecosystem changes, such as alterations in soil nutrient cycling, and redistribution and loss of soil organic matter. However, the present and future extent of these invasions is difficult to evaluate without a better understanding of the factors that control the distribution and abundance of earthworms in previously non-invaded habitats. In this study, the species composition and short-term dynamics of three exotic earthworm invasion fronts were studied at a northern hardwood forest in south-central New York State (USA). Belt transects were established at each of the three locations to sample from earthworm-invaded areas through transition zones and into invasion front areas. Lumbricus rubellus, L. terrestrisandOctolasion tyrtaeum were the most common species, but their distribution was not homogeneous along the transects. Whereas, L. rubellus was the only species with relatively high adult densities at transition zones and invasion fronts, L. terrestris and O. tyrtaeum occurred mostly in the heavily earthworm-invaded areas and were rare at the invasion fronts. The density of earthworms along the transects decreased by 60–87 from June 2001 to October 2002 and then recovered in 2003 to values similar to those of 2001. This decrease was apparently caused by reduced recruitment of immature earthworms, probably related to the severe drought periods that the study area experienced in 2001 and 2002. Our data suggest that climate and topography, through their effects on soil moisture patterns, can be critical factors controlling the distribution and spread of exotic earthworms in previously non-invaded habitats.     

八大公山中亚热带山地常绿落叶阔叶混交林物种组成与结构

山地常绿落叶阔叶混交林在我国主要分布在中亚热带1000～2000 m中山地带。八大公山位于武陵山系北端,中亚热带北缘,具有保护较好的典型山地常绿落叶阔叶混交林。为了研究常绿落叶阔叶混交林动态和川东-鄂西植物多样性形成及维持机制,按照CTFS（Center for Tropical Forest Science）样地建设标准,中国科学院武汉植物园于2010-2011年在湖南八大公山（Badagongshan,BDGS）国家级自然保护区建立了一个25 hm2（500 m×500 m）中亚热带山地常绿落叶阔叶混交林动态监测样地（Forest Dynamic Plot）。本研究以样地内所有胸径（DBH）≥1 cm的木本植物（不含竹藤）为研究对象,分析该样地区系成分、群落组成、垂直结构、径级结构和空间分布。结果表明：八大公山样地属于典型的山地常绿落叶阔叶混交林,植物以泛热带分布科（24.50%）和北温带分布属（24.56%）占优势,包括9种珍稀濒危植物。样地内木本植物合计53科、114属、238种（含种下分类单位）,186556株存活个体。落叶树144种,常绿树94种。壳斗科（Fagaceae）、杜鹃花科（Ericaceae）、樟科（Lauraceae）和山茶科（Theaceae）占据优势;个体数超过1000的有38个物种,以常绿树种（63.16%）为主;稀有种（〈25株）103个（占样地物种总数43.28%）。群落垂直结构可分为乔木层（优势树种多脉青冈Cyclobalanopsis multinervis和亮叶水青冈Fagus lucida）、亚乔木层（优势种长蕊杜鹃Rhododendron stamineum和黄丹木姜子Litsea elongata）和灌木层（优势种短柱柃Eurya brevistyla和薄叶山矾Symplocos anomala）。个体数超过1000的树种和稀有种主要来自灌木层。由于生长型限制,样地内树种平均胸径较小（5.41 cm）,DBH〈5 cm的个体占优势（68.40%）。主要优势树种的径级结构呈现倒J型。     

Growth,Net Production,Litter Decomposition,and Net Nitrogen Accumulation by Epiphytic Bryophytes in a Tropical Montane Forest1

To understand the ecological roles of epiphytic bryophytes in the carbon (C) and nitrogen (N) cycles of a tropical montane forest, we used samples in enclosures to estimate rates of growth, net production, and N accumulation by shoots in the canopy, and litterbags, to estimate rates of decomposition and N dynamics of epiphytic bryophyte litter in the canopy and on the forest floor in Monteverde, Costa Rica. Growth of epiphytic bryophytes was estimated at 30.0–49.9 percent/yr, net production at 122–203 g/m²/yr, and N accumulation at 1.8–3.0 g N/m²/yr. Cumulative mass loss from litterbags after one and two years in the canopy was 17 ± 2 and 19 ± 2 percent (mean ± 1 SE) of initial sample mass, respectively, and mass loss from litter and green shoots in litterbags after one year on the forest floor was 29 ± 2 and 45 ± 3 percent, respectively. Approximately 30 percent of the initial N mass was released rapidly from litter in both locations. Nitrogen loss from green shoots on the forest floor was greater; about 47 percent of the initial N mass was lost within the first three months. There was no evidence for net N immobilization by litter or green shoots, but the remaining N in litter was apparently recalcitrant. Annual net accumulation of C and N by epiphytic bryophytes was estimated at 37–64 g C/m²/yr and 0.8–1.3 g N/m²/yr, respectively. Previous research at this site indicated that epiphytic bryophytes retain inorganic N from atmospheric deposition to the canopy. Therefore, they play a major role in transforming N from mobile to highly recalcitrant forms in this ecosystem.     

Fine Root Production and Turnover in a Norway Spruce Stand in Northern Sweden: Effects of Nitrogen and Water Manipulation

Fine root length production, biomass production, and turnover in forest floor and mineral soil (0–30 cm) layers were studied in relation to irrigated (I) and irrigated-fertilized (IL) treatments in a Norway spruce stand in northern Sweden over a 2-year period. Fine roots (<1 mm) of both spruce and understory vegetation were studied. Minirhizotrons were used to estimate fine root length production and turnover, and soil cores were used to estimate standing biomass. Turnover was estimated as both the inverse of root longevity (RTL) and the ratio of annual root length production to observed root length (RTR). RTR values of spruce roots in the forest floor in I and IL plots were 0.6 and 0.5 y⁻¹, respectively, whereas the corresponding values for RTL were 0.8 and 0.9 y⁻¹. In mineral soil, corresponding values for I, IL, and control (C) plots were 1.2, 1.2, and 0.9 y⁻¹ (RTR) and 0.9, 1.1, and 1 y⁻¹ (RTL). RTR and RTL values of understory vegetation roots were 1 and 1.1 y⁻¹, respectively. Spruce root length production in both the forest floor and the mineral soil in I plots was higher than in IL plots. The IL-treated plots gave the highest estimates of spruce fine root biomass production in the forest floor, but, for the mineral soil, the estimates obtained for the I plots were the highest. The understory vegetation fine root production in the I and IL plots was similar for both the forest floor and the mineral soil and higher (for both layers) than in C plots. Nitrogen (N) turnover in the forest floor and mineral soil layers (summed) via spruce roots in IL, I, and C plots amounted to 2.4, 2.1, and 1.3 g N m⁻² y⁻¹, and the corresponding values for field vegetation roots were 0.6, 0.5, and 0.3 g N m⁻² y⁻¹. It was concluded that fertilization increases standing root biomass, root production, and N turnover of spruce roots in both the forest floor and mineral soil. Data on understory vegetation roots are required for estimating carbon budgets in model studies.     

Alder and lupine enhance nitrogen cycling in a degraded forest soil in Northern Sweden

Positive effects of legumes and actinorhizal plants on N-poor soils have been observed in many studies but few have been done at high latitudes, which was the location of our study. We measured N₂ fixation and several indices of soil N at a site near the Arctic Circle in northern Sweden. More than 20 years ago lupine (Lupinus nootkatensis Donn) and gray alder (Alnus incana L. Moench) were planted on this degraded forest site. We measured total soil N, net N mineralization and nitrification with a buried bag technique, and fluxes of NH⁺ ₄ and NO^– ₃ as collected on ion exchange membranes. We also estimated N₂ fixation activity of the N₂-fixing plants by the natural abundance of ¹⁵N of leaves with Betula pendula Roth. as reference species. Foliar nitrogen in the N₂-fixing plants was almost totally derived from N₂ fixation. Plots containing N₂-fixing species generally had significantly higher soil N and N availability than a control plot without N₂-fixing plants. Taken together, all measurements indicated that N₂-fixing plants can be used to effectively improve soil fertility at high latitudes in northern Sweden.     

Production of CO2 in Soil Profiles of a California Annual Grassland

Soils play a key role in the global cycling of carbon (C), storing organic C, and releasing CO₂ to the atmosphere. Although a large number of studies have focused on the CO₂ flux at the soil–air interface, relatively few studies have examined the rates of CO₂ production in individual layers of a soil profile. Deeper soil horizons often have high concentrations of CO₂ in the soil air, but the sources of this CO₂ and the spatiotemporal dynamics of CO₂ production throughout the soil profile are poorly understood. We studied CO₂ dynamics in six soil profiles arrayed across a grassland hillslope in coastal southern California. Gas probes were installed in each profile and gas samples were collected weekly or biweekly over a three-year period. Using soil air CO₂ concentration data and a model based on Fick’s law of diffusion, we modeled the rates of CO₂ production with soil profile depth. The CO₂ diffusion constants were checked for accuracy using measured soil air ²²²Rn activities. The modeled net CO₂ production rates were compared with CO₂ fluxes measured at the soil surface. In general, the modeled and measured net CO₂ fluxes were very similar although the model consistently underestimated CO₂ production rates in the surficial soil horizons when the soils were moist. Profile CO₂ production rates were strongly affected by the inter- and intra-annual variability in rainfall; rates were generally 2–10 times higher in the wet season (December to May) than in the dry season (June to November). The El Niño event of 1997–1998, which brought above-average levels of rainfall to the study site, significantly increased CO₂ production in both the surface and subsurface soil horizons. Whole profile CO₂ production rates were approximately three times higher during the El Niño year than in the following years of near-average rainfall. During the dry season, when the net rates of CO₂ flux from the soil profiles are relatively low (4–11 mg C– CO₂ m⁻² h⁻¹), 20%–50% of the CO₂ diffusing out of the profiles appears to originate in the relatively moist soil subsurface (defined here as those horizons below 40 cm in depth). The natural abundance ¹⁴C signatures of the CO₂ and soil organic C suggest that the subsurface CO₂ is derived from the microbial mineralization of recent organic C, possibly dissolved organic C transported to the subsurface horizons during the wet season.     

Nitrogen Limits an Invasive Perennial Shrub in Forest Understory

Plant invasions can harm communities by domination of one or more vegetation layers. We studied whether Japanese barberry (Berberis thunbergii DC.) is limited by soil acidity or nitrogen availability in its domination of relatively undisturbed forest understories. In two sites, one more acid than the other, we applied lime, urea, or a sawdust–sugar mix to replicate plots in established barberry populations. We predicted that the acid site would be pH or cation limited, while the less acid site was N limited, unless N availability was inherently higher before treatment. Barberry above-ground net primary production (NPP) was estimated by a combination of harvest and allometric analysis. Foliar N increased in the urea treatment and was proportional to incubation estimates of net N mineralization and nitrification. Foliar Ca and P were unaffected by the treatments. Foliar K was proportional to foliar N. The more acid site had higher foliar Mn, but otherwise the sites differed little. Barberry NPP was proportional to pre-treatment biomass. The ratio of net production to pre-treatment woody biomass (relative production rate) increased with foliar N and soil N availability and decreased when soil N was immobilized by sawdust and sugar. There was no effect of soil pH or cation status on barberry growth, although a correlation with foliar K was reflected by the maintenance of a constant K : N ratio. Although more severely acid sites may be less invasible than those studied here, N availability is the primary limitation to invasive dominance in this landscape. This revised version was published online in July 2006 with corrections to the Cover Date.     

Natural<Superscript>15</Superscript> N Abundance of Plants and Soil N in a Temperate Coniferous Forest

Measurement of nitrogen isotopic composition (¹⁵N) of plants and soil nitrogen might allow the characteristics of N transformation in an ecosystem to be detected. We tested the measurement of ¹⁵N for its ability to provide a picture of N dynamics at the ecosystem level by doing a simple comparison of ¹⁵N between soil N pools and plants, and by using an existing model. ¹⁵N of plants and soil N was measured together with foliar nitrate reductase activity (NRA) and the foliar NO₃^– pool at two sites with different nitrification rates in a temperature forest in Japan. ¹⁵N of plants was similar to that of soil NO₃^– in the high-nitrification site. Because of high foliar NRA and the large foliar NO₃^– pool at this site, we concluded that plant ¹⁵N indicated a great reliance of plants on soil NO₃^– there. However, many ¹⁵N of soil N overlapped each other at the other site, and ¹⁵N could not provide definitive evidence of the N source. The existing model was verified by measured ¹⁵N of soil inorganic N and it explained the variations of plant ¹⁵N between the two sites in the context of relative importance of nitrification, but more information about isotopic fractionations during plant N uptake is required for quantitative discussions about the plant N source. The model applied here can provide a basis to compare ¹⁵N signatures from different ecosystems and to understand N dynamics.     

Comparison of Belowground Biomass in C3- and C4-Dominated Mixed Communities in a Chesapeake Bay Brackish Marsh

Belowground biomass is a critical factor regulating ecosystem functions of coastal marshes, including soil organic matter (SOM) accumulation and the ability of these systems to keep pace with sea-level rise. Nevertheless, belowground biomass responses to environmental and vegetation changes have been given little emphasis marsh studies. Here we present a method using stable carbon isotopes and color to identify root and rhizomes of Schoenoplectus americanus (Pers.) Volk. ex Schinz and R. Keller (C₃) and Spartina patens (Ait.) Muhl. (C₄) occurring in C₃− and C₄-dominated communities in a Chesapeake Bay brackish marsh. The functional significance of the biomass classes we identified is underscored by differences in their chemistry, depth profiles, and variation in biomass and profiles relative to abiotic and biotic factors. C₃ rhizomes had the lowest concentrations of cellulose (29.19%) and lignin (14.43%) and the lowest C:N (46.97) and lignin:N (0.16) ratios. We distinguished two types of C₃ roots, and of these, the dark red C₃ roots had anomalously high C:N (195.35) and lignin:N (1.14) ratios, compared with other root and rhizome classes examined here and with previously published values. The C₄-dominated community had significantly greater belowground biomass (4119.1 g m⁻²) than the C₃-dominated community (3256.9 g m⁻²), due to greater total root biomass and a 3.6-fold higher C₃-root:rhizome ratio in the C₄-dominated community. C₃ rhizomes were distributed significantly shallower in the C₄-dominated community, while C₃ roots were significantly deeper. Variability in C₃ rhizome depth distributions was explained primarily by C₄ biomass, and C₃ roots were explained primarily by water table height. Our results suggest that belowground biomass in this system is sensitive to slight variations in water table height (across an 8 cm range), and that the reduced overlap between C₃ and C₄ root profiles in the C₄-dominated community may account for the greater total root biomass observed in that community. Given that future elevated atmospheric CO₂ and accelerated sea-level rise are likely to increase C₃ abundance in Atlantic and Gulf coast marshes, investigations that quantify how patterns of C₃ and C₄ belowground biomass respond to environmental and biological factors stand to improve our understanding of ecosystem-wide impacts of global changes on coastal wetlands.     

Subsurface CO<Subscript>2</Subscript> Dynamics in Temperate Beech and Spruce Forest Stands

Rates of soil respiration (CO₂ effluxes), subsurface pore gas CO₂/O₂ concentrations, soil temperature and soil water content were measured for 15 months in two temperate and contrasting Danish forest ecosystems: beech (Fagus sylvatica L.) and Norway spruce (Picea abies [L.] Karst.). Soil CO₂ effluxes showed a distinct seasonal trend in the range of 0.48–3.3 μmol CO₂ m⁻² s⁻¹ for beech and 0.50–2.92 μmol CO₂ m⁻² s⁻¹ for spruce and were well-correlated with near-surface soil temperatures. The soil organic C-stock (upper 1 m including the O-horizon) was higher in the spruce stand (184±23 Mg C ha⁻¹) compared to the beech stand (93±19 Mg C ha⁻¹) and resulted in a faster turnover time as calculated by mass/flux in soil beneath the beech stand (28 years) compared to spruce stand (60 years). Observed soil CO₂ concentrations and effluxes were simulated using a Fickian diffusion-reaction model based on vertical CO₂ production rates and soil diffusivity. Temporal trends were simulated on the basis of observed trends in the distribution of soil water, temperature, and live roots as well as temperature and water content sensitivity functions. These functions were established based on controlled laboratory incubation experiments. The model was successfully validated against observed soil CO₂ effluxes and concentrations and revealed that temporal trends generally could be linked to variations in subsurface CO₂ production rates and diffusion over time and with depths. However, periods with exceptionally high CO₂ effluxes (> 20 μmol CO₂ m⁻² s⁻¹) were noted in March 2000 in relation to drying after heavy rain and after the removal of snow from collars. Both cases were considered non-steady state and could not be simulated.     

Effect of Different Carbon and Nitrogen Sources on Clostridium argentinense Toxigenicity in Coculture withPseudomonas mendocina

The effect of different carbon and nitrogen sources on the production of toxin by Clostridium argentinense was examined. The toxin production by C. argentinense in coculture with Pseudomonas mendocina increased in all the cases in relation to that produced by monocultures independent of the nature of the source. Using dextrin as carbon source C. argentinense produced the highest levels of toxin both in monocultures (300 LD₅₀/mL) and in cocultures with P. mendocina (5000 LD₅₀/mL). Experiments run in a microfermenter showed that the slow growth of cocultures associated with the assimilation of dextrin and the pH and Eh profiles favoured the production of toxin. Of the nitrogen sources assayed, corn steep liquor sustained the highest levels of toxin in both monocultures and cocultures with 3 and 2.8 fold increases with respect to that obtained using proteose peptone. The toxin production by C. argentinense cultures and C. argentinense–P. mendocina cocultures was highly dependent on the nature of the carbon and nitrogen sources used in the culture media. Growth of C. argentinense on substrates slowly assimilated stimulated the production of toxin.     

Chemistry and Dynamics of Dissolved Organic Matter in a Temperate Coniferous Forest on Andic Soils: Effects of Litter Quality

Dissolved organic matter (DOM) plays an important role in transporting carbon and nitrogen from forest floor to mineral soils in temperate forest ecosystems. Thus, the retention of DOM via sorption or microbial assimilation is one of the critical steps for soil organic matter formation in mineral soils. The chemical properties of DOM are assumed to control these processes, yet we lack fundamental information that links litter quality, DOM chemistry, and DOM retention. Here, we studied whether differences in litter quality affect solution chemistry and whether changes in litter inputs affect DOM quality and removal in the field. The effects of litter quality on solution chemistry were evaluated using chemical fractionation methods for laboratory extracts and for soil water collected from a temperate coniferous forest where litter inputs had been altered. In a laboratory extraction, litter type (needle, wood, root) and the degree of decomposition strongly influenced solution chemistry. Root litter produced more than 10 times more water-extractable dissolved organic N (DON) than any other litter type, suggesting that root litter may be most responsible for DON production in this forest ecosystem. The chemical composition of the O-horizon leachate was similar under all field treatments (doubled needle, doubled wood, and normal litter inputs). O-horizon leachate most resembled laboratory extracts of well-decomposed litter (that is, a high proportion of hydrophobic acids), in spite of the significant amount of litter C added to the forest floor and a tendency toward higher mean DOM under doubled-Litter treatments. A lag in DOM production from added litter or microbial modification might have obscured chemical differences in DOM under the different treatments. Net DOM removal in this forest soil was strong; DOM concentration in the water deep in the mineral soil was always low regardless of concentrations in water that entered the mineral soil and of litter input manipulation. High net removal of DOM from O-horizon leachate, in spite of extremely low initial hydrophilic neutral content (labile DOM), coupled with the lack of influence by season or soil depth, suggests that DOM retention in the soil was mostly by abiotic sorption.