Primary production and carbon allocation in relation to nutrient supply in a tropical experimental forest

Nutrient supply commonly limits aboveground plant productivity in forests, but the effects of an altered nutrient supply on gross primary production (GPP) and patterns of carbon (C) allocation remain poorly characterized. Increased nutrient supply may lead to a higher aboveground net primary production (ANPP), but a lower total belowground carbon allocation (TBCA), with little change in either aboveground plant respiration (APR) or GPP. Alternatively, increases in nutrient supply may increase GPP, with the quantity of GPP allocated aboveground increasing more steeply than the quantity of GPP allocated belowground. To examine the effects of an elevated nutrient supply on the C allocation patterns in forests, we determined whole‐ecosystem C budgets in unfertilized plots of Eucalyptus saligna and in adjacent plots receiving regular additions of 65 kg N ha^?1, 31 kg P ha^?1, 46 kg K ha^?1, and macro‐ and micronutrients. We measured the absolute flux of C allocated to the components of GPP (ANPP, TBCA and APR), as well as the fraction of GPP allocated to these components. Fertilization dramatically increased GPP. Averaged over 3 years, GPP in the fertilized plots was 34% higher than that in the unfertilized controls (3.95 vs. 2.95 kg C m^?2 yr^?1). Fertilization‐related increases in GPP were allocated entirely aboveground – ANPP was 85% higher and APR was 57% higher in the fertilized than in the control plots, while TBCA did not differ significantly between treatments. Carbon use efficiency (NPP/GPP) was slightly higher in the fertilized (0.53) compared with the control plots (0.51). Overall, fertilization increased ANPP and APR, and these increases were related to a greater GPP and an increase in the fraction of GPP allocated aboveground.     

Contribution of aboveground litter,belowground litter,and rhizosphere respiration to total soil CO<Subscript>2</Subscript> efflux in an old growth coniferous forest

In an old growth coniferous forest located in the central Cascade Mountains, Oregon, we added or removed aboveground litter and terminated live root activity by trenching to determine sources of soil respiration. Annual soil efflux from control plots ranged from 727 g C m⁻² year⁻¹ in 2002 to 841 g C m⁻² year⁻¹ in 2003. We used aboveground litter inputs (149.6 g C m⁻² year⁻¹) and differences in soil CO₂ effluxes among treatment plots to calculate contributions to total soil efflux by roots and associated rhizosphere organisms and by heterotrophic decomposition of organic matter derived from aboveground and belowground litter. On average, root and rhizospheric respiration (R_r) contributed 23%, aboveground litter decomposition contributed 19%, and belowground litter decomposition contributed 58% to total soil CO₂ efflux, respectively. These values fall within the range of values reported elsewhere, although our estimate of belowground litter contribution is higher than many published estimates, which we argue is a reflection of the high degree of mycorrhizal association and low nutrient status of this ecosystem. Additionally, we found that measured fluxes from plots with doubled needle litter led to an additional 186 g C m⁻² year⁻¹ beyond that expected based on the amount of additional carbon added; this represents a priming effect of 187%, or a 34% increase in the total carbon flux from the plots. This finding has strong implications for soil C storage, showing that it is inaccurate to assume that increases in net primary productivity will translate simply and directly into additional belowground storage.     

Linking Belowground Carbon Allocation to Anaerobic CH4 and CO2 Production in a Forested Peatland,New York State

Heterotrophic soil microorganisms rely on carbon (C) allocated belowground in plant production, but belowground C allocation (BCA) by plants is a poorly quantified part of ecosystem C cycling, especially, in peat soil. We applied a C balance approach to quantify BCA in a mixed conifer-red maple (Acer rubrum) forest on deep peat soil. Direct measurements of CH₄ and CO₂ fluxes across the soil surface (soil respiration), production of fine and small plant roots, and aboveground litterfall were used to estimate respiration by roots, by mycorrhizae and by free-living soil microorganisms. Measurements occurred in two consecutive years. Soil respiration rates averaged 1.2 bm μmol m^{? 2} s^{? 1} for CO₂ and 0.58 nmol m^{? 2} s^{? 1} for CH₄ (371 to 403 g C m^{? 2} year^{? 1}). Carbon in aboveground litter (144 g C m^{? 2} year^{? 1}) was 84% greater than C in root production (78 g C m^{? 2} year^{? 1}). Complementary in vitro assays located high rates of anaerobic microbial activity, including methanogenesis, in a dense layer of roots overlying the peat soil and in large-sized fragments within the peat matrix. Large-sized fragments were decomposing roots and aboveground leaf and twig litter, indicating that relatively fresh plant production supported most of the anaerobic microbial activity. Respiration by free-living soil microorganisms in deep peat accounted for, at most, 29 to 38 g C m^{? 2} year^{? 1}. These data emphasize the close coupling between plant production, ecosystem-level C cycling and soil microbial ecology, which BCA can help reveal.     

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 carbon storage, litterfall and CO2 efflux in fertilized and unfertilized larch (Larix leptolepis) plantations

This study evaluated the effects of forest fertilization on the forest carbon (C) dynamics in a 36-year-old larch (Larix leptolepis) plantation in Korea. Above- and below-ground C storage, litterfall, root decomposition and soil CO₂ efflux rates after fertilization were measured for 2 years. Fertilizers were applied to the forest floor at rates of 112 kg N ha⁻¹ year⁻¹, 75 kg P ha⁻¹ year⁻¹ and 37 kg K ha⁻¹ year⁻¹ for 2 years (May 2002, 2003). There was no significant difference in the above-ground C storage between fertilized (41.20 Mg C ha⁻¹) and unfertilized (42.25 Mg C ha⁻¹) plots, and the C increment was similar between the fertilized (1.65 Mg C ha⁻¹ year⁻¹) and unfertilized (1.52 Mg C ha⁻¹ year⁻¹) plots. There was no significant difference in the soil C storage between the fertilized and unfertilized plots at each soil depth (0–15, 15–30 and 30–50 cm). The organic C inputs due to litterfall ranged from 1.57 Mg C ha⁻¹ year⁻¹ for fertilized to 1.68 Mg C ha⁻¹ year⁻¹ for unfertilized plots. There was no significant difference in the needle litter decomposition rates between the fertilized and unfertilized plots, while the decomposition of roots with 1–2 mm diameters increased significantly with the fertilization relative to the unfertilized plots. The mean annual soil CO₂ efflux rates for the 2 years were similar between the fertilized (0.38 g CO₂ m⁻² h⁻¹) and unfertilized (0.40 g CO₂ m⁻² h⁻¹) plots, which corresponded with the similar fluctuation in the organic carbon (litterfall, needle and root decomposition) and soil environmental parameters (soil temperature and soil water content). These results indicate that little effect on the C dynamics of the larch plantation could be attributed to the 2-year short-term fertilization trials and/or the soil fertility in the mature coniferous plantation used in this study.     

Invasion of <Emphasis Type="Italic">Spartina alterniflora</Emphasis> Enhanced Ecosystem Carbon and Nitrogen Stocks in the Yangtze Estuary,China

Whether plant invasion increases ecosystem carbon (C) stocks is controversial largely due to the lack of knowledge about differences in ecophysiological properties between invasive and native species. We conducted a field experiment in which we measured ecophysiological properties to explore the response of the ecosystem C stocks to the invasion of Spartina alterniflora (Spartina) in wetlands dominated by native Scirpus mariqueter (Scirpus) and Phragmites australis (Phragmites) in the Yangtze Estuary, China. We measured growing season length, leaf area index (LAI), net photosynthetic rate (Pn), root biomass, net primary production (NPP), litter quality and litter decomposition, plant and soil C and nitrogen (N) stocks in ecosystems dominated by the three species. Our results showed that Spartina had a longer growing season, higher LAI, higher Pn, and greater root biomass than Scirpus and Phragmites. Net primary production (NPP) was 2.16 kg C m⁻² y⁻¹ in Spartina ecosystems, which was, on average, 1.44 and 0.47 kg C m⁻² y⁻¹ greater than that in Scirpus and Phragmites ecosystems, respectively. The litter decomposition rate, particularly the belowground decomposition rate, was lower for Spartina than Scirpus and Phragmites due to the lower litter quality of Spartina. The ecosystem C stock (20.94 kg m⁻²) for Spartina was greater than that for Scirpus (17.07 kg m⁻²), Phragmites (19.51 kg m⁻²) and the mudflats (15.12 kg m⁻²). Additionally, Spartina ecosystems had a significantly greater N stock (698.8 g m⁻²) than Scirpus (597.1 g m⁻²), Phragmites ecosystems (578.2 g m⁻²) and the mudflats (375.1 g m⁻²). Our results suggest that Spartina invasion altered ecophysiological processes, resulted in changes in NPP and litter decomposition, and ultimately led to enhanced ecosystem C and N stocks in the invaded ecosystems in comparison to the ecosystems with native species.     

Soil respiration and carbon balance in a subtropical native forest and two managed plantations

From 1999 to 2003, a range of carbon fluxes was measured and integrated to establish a carbon balance for a natural evergreen forest of Castanopsis kawakamii (NF) and adjacent monoculture evergreen plantations of C. kawakamii (CK) and Chinese fir (Cunninghamia lanceolata, CF) in Sanming Nature Reserve, Fujian, China. Biomass carbon increment of aboveground parts and coarse roots were measured by the allometric method. Above- and belowground litter C inputs were assessed by litter traps and sequential cores, respectively. Soil respiration (SR) was determined by the alkaline absorbance method, and the contribution from roots, above- and belowground litters was separated by the DIRT plots. Annual SR averaged 13.742 t C ha⁻¹ a⁻¹ in the NF, 9.439 t C ha⁻¹ a⁻¹ in the CK, and 4.543 t C ha⁻¹ a⁻¹ in the CF. For all forests, SR generally peaked in later spring or early summer (May or June). The contribution of root respiration ranged from 47.8% in the NF to 40.3% in the CF. On average, soil heterotrophic respiration (HR) was evenly distributed between below- (47.3∼54.5%) and aboveground litter (45.5%–52.7%). Annual C inputs (t C ha⁻¹ a⁻¹) from litterfall and root turnover averaged 4.452 and 4.295, 4.548 and 2.313, and 2.220 and 1.265, respectively, in the NF, CK, and CF. As compared to HR, annual net primary production (NPP) of 11.228, 13.264, and 6.491 t C ha⁻¹ a⁻¹ in the NF, CK, and CF brought a positive net ecosystem production (NEP) of 4.144, 7.514, and 3.677 t C ha⁻¹ a⁻¹, respectively. It suggests that native forest in subtropical China currently acts as an important carbon sink just as the timber plantation does, and converting native forest to tree plantations locally during last decades might have caused a high landscape carbon loss to the atmosphere.     

Fine Root Dynamics and Forest Production Across a Calcium Gradient in Northern Hardwood and Conifer Ecosystems

Losses of soil base cations due to acid rain have been implicated in declines of red spruce and sugar maple in the northeastern USA. We studied fine root and aboveground biomass and production in five northern hardwood and three conifer stands differing in soil Ca status at Sleepers River, VT; Hubbard Brook, NH; and Cone Pond, NH. Neither aboveground biomass and production nor belowground biomass were related to soil Ca or Ca:Al ratios across this gradient. Hardwood stands had 37% higher aboveground biomass (P = 0.03) and 44% higher leaf litter production (P < 0.01) than the conifer stands, on average. Fine root biomass (<2 mm in diameter) in the upper 35 cm of the soil, including the forest floor, was very similar in hardwoods and conifers (5.92 and 5.93 Mg ha⁻¹). The turnover coefficient (TC) of fine roots smaller than 1 mm ranged from 0.62 to 1.86 y⁻¹ and increased significantly with soil exchangeable Ca (P = 0.03). As a result, calculated fine root production was clearly higher in sites with higher soil Ca (P = 0.02). Fine root production (biomass times turnover) ranged from 1.2 to 3.7 Mg ha⁻¹ y⁻¹ for hardwood stands and from 0.9 to 2.3 Mg ha⁻¹ y⁻¹ for conifer stands. The relationship we observed between soil Ca availability and root production suggests that cation depletion might lead to reduced carbon allocation to roots in these ecosystems.     

Seasonal dynamics of fine root biomass, root length density, specific root length, and soil resource availability in a Larix gmelinii plantation

Fine root turnover is a major pathway for carbon and nutrient cycling in terrestrial ecosystems and is most likely sensitive to many global change factors. Despite the importance of fine root turnover in plant C allocation and nutrient cycling dynamics and the tremendous research efforts in the past, our understanding of it remains limited. This is because the dynamics processes associated with soil resources availability are still poorly understood. Soil moisture, temperature, and available nitrogen are the most important soil characteristics that impact fine root growth and mortality at both the individual root branch and at the ecosystem level. In temperate forest ecosystems, seasonal changes of soil resource availability will alter the pattern of carbon allocation to belowground. Therefore, fine root biomass, root length density (RLD) and specific root length (SRL) vary during the growing season. Studying seasonal changes of fine root biomass, RLD, and SRL associated with soil resource availability will help us understand the mechanistic controls of carbon to fine root longevity and turnover. The objective of this study was to understand whether seasonal variations of fine root biomass, RLD and SRL were associated with soil resource availability, such as moisture, temperature, and nitrogen, and to understand how these soil components impact fine root dynamics in Larix gmelinii plantation. We used a soil coring method to obtain fine root samples (⩽2 mm in diameter) every month from May to October in 2002 from a 17-year-old L. gmelinii plantation in Maoershan Experiment Station, Northeast Forestry University, China. Seventy-two soil cores (inside diameter 60 mm; depth intervals: 0–10 cm, 10–20 cm, 20–30 cm) were sampled randomly from three replicates 25 m × 30 m plots to estimate fine root biomass (live and dead), and calculate RLD and SRL. Soil moisture, temperature, and nitrogen (ammonia and nitrates) at three depth intervals were also analyzed in these plots. Results showed that the average standing fine root biomass (live and dead) was 189.1 g·m⁻²·a⁻¹, 50% (95.4 g·m⁻²·a⁻¹) in the surface soil layer (0–10 cm), 33% (61.5 g·m⁻²·a⁻¹), 17% (32.2 g·m⁻²·a⁻¹) in the middle (10–20 cm) and deep layer (20–30cm), respectively. Live and dead fine root biomass was the highest from May to July and in September, but lower in August and October. The live fine root biomass decreased and dead biomass increased during the growing season. Mean RLD (7,411.56 m·m⁻³·a⁻¹) and SRL (10.83 m·g⁻¹·a⁻¹) in the surface layer were higher than RLD (1 474.68 m·m⁻³·a⁻¹) and SRL (8.56 m·g⁻¹·a⁻¹) in the deep soil layer. RLD and SRL in May were the highest (10 621.45 m·m⁻³ and 14.83m·g⁻¹) compared with those in the other months, and RLD was the lowest in September (2 198.20 m·m⁻³) and SRL in October (3.77 m·g⁻¹). Seasonal dynamics of fine root biomass, RLD, and SRL showed a close relationship with changes in soil moisture, temperature, and nitrogen availability. To a lesser extent, the temperature could be determined by regression analysis. Fine roots in the upper soil layer have a function of absorbing moisture and nutrients, while the main function of deeper soil may be moisture uptake rather than nutrient acquisition. Therefore, carbon allocation to roots in the upper soil layer and deeper soil layer was different. Multiple regression analysis showed that variation in soil resource availability could explain 71–73% of the seasonal variation of RLD and SRL and 58% of the variation in fine root biomass. These results suggested a greater metabolic activity of fine roots living in soil with higher resource availability, which resulted in an increased allocation of carbohydrate to these roots, but a lower allocation of carbohydrate to those in soil with lower resource availability. __________ Translated from Acta Phytoecologica Sinica, 2005, 29(3): 403–410 [译自: 植物生态学报, 2005, 29(3): 403–410]     

Modelling the Long-Term Stabilization of Carbon from Maize in a Silty Soil

Soil organic carbon (SOC) models have been widely used to predict SOC change with changing environmental and management conditions, but the accuracy of the prediction is often open to question. Objectives were (i) to quantify the amounts of C derived from maize in soil particle size fractions and at various depths in a long-term field experiment using ¹³C/¹²C analysis, (ii) to model changes in the organic C, and (iii) to compare measured and modelled pools of C. Maize was cultivated for 24 years on a silty Luvisol which resulted in a stock of 1.9 kg maize-derived C m⁻² (36% of the total organic C) in the Ap horizon. The storage of maize-derived C in particle size fractions of the Ap horizon decreased in the order clay (0.65 kg C m⁻²) > fine and medium silt (0.43) > coarse silt (0.33) > fine sand (0.13) > medium sand (0.12) > coarse sand (0.06) and the turnover times of C₃-derived C ranged from 26 (fine sand) to 77 years (clay). The turnover times increased with increasing soil depth. We used the Rothamsted Carbon Model to model the C dynamics and tested two model approaches: model A did not have any adjustable parameters, but included the Falloon equation for the estimation of the amount of inert organic matter (IOM) and independent estimations of C inputs into the soil. The model predicted well the changes in C₃-derived C with time but overestimated the changes in maize-derived C 1.6-fold. In model B, the amounts of IOM and C inputs were optimized to match the measured C₃- and C₄-derived SOC stocks after 24 years of continuous maize. This model described the experimental data well, but the modelled annual maize C inputs (0.41 kg C m⁻² a⁻¹) were less than the independently estimated total input of maize litter C (0.63 kg C m⁻² a⁻¹) and even less than the annual straw C incorporated into the soil (0.46 kg C m⁻² a⁻¹). These results indicated that the prediction of the Rothamsted Carbon Model with independent parameterization served only as an approximation for this site. The total amount of organic C associated with the fraction 0–63 μm agreed well with the sum of the pools ‘microbial biomass’, ‘humified-organic matter’ and IOM of the model B. However, the amount of maize-derived C in this fraction (3.4 g kg⁻¹) agreed only satisfactorily with the sum of maize-derived C in the pools ‘microbial biomass’ and ‘humified organic matter’ (2.6 g kg⁻¹).     

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.     

Response of Litter Decomposition to Simulated N Deposition in Disturbed, Rehabilitated and Mature Forests in Subtropical China

The response of decomposition of litter for the dominant tree species in disturbed (pine), rehabilitated (pine and broadleaf mixed) and mature (monsoon evergreen broadleaf) forests in subtropical China to simulated N deposition was studied to address the following hypothesis: (1) litter decomposition is faster in mature forest (high soil N availability) than in rehabilitated/disturbed forests (low soil N availability); (2) litter decomposition is stimulated by N addition in rehabilitated and disturbed forests due to their low soil N availability; (3) N addition has little effect on litter decomposition in mature forest due to its high soil N availability. The litterbag method (a total of 2880 litterbags) and N treatments: Control-no N addition, Low-N: −5 g N m⁻² y⁻¹, Medium-N: −10 g N m⁻² y⁻¹, and High-N: −15 g N m⁻² y⁻¹, were employed to evaluate decomposition_. Results indicated that mature forest, which has likely been N saturated due to both long-term high N deposition in the region and the age of the ecosystem, had the highest litter decomposition rate, and exhibited no significant positive and even some negative response to nitrogen additions. However, both disturbed and rehabilitated forests, which are still N limited due to previous land use history, exhibited slower litter decomposition rates with significant positive effects from nitrogen additions. These results suggest that litter decomposition and its responses to N addition in subtropical forests of China vary depending on the nitrogen status of the ecosystem.     

Fine Roots vs. Needles: A Comparison of 13C and 15N Dynamics in a Ponderosa Pine Forest Soil

Plant allocation patterns may affect soil C and N storage due to differences in litter quality and the depth of plant C and N inputs into the soil. We studied the dynamics of dual-labeled (¹³C/¹⁵N) Pinus ponderosa needles and fine roots placed at two soil depths (O and A horizon) in a temperate conifer forest soil during 2 y. Input of C as fine roots resulted in much more C retained in soil (70.5 ± 2.2% of applied) compared with needle C (42.9 ± 1.3% of applied) after 1.5 y. Needles showed faster mass loss, rates of soil ¹³CO₂ efflux, and more ¹⁵N immobilized into microbial biomass than did fine roots. The larger proportion of labile C compounds initially present in needles (17% more needle C was water soluble than in fine roots) likely contributed to its shorter C residence time and greater degree of transformation in the soil. A double exponential decay function best described the rate of ¹³C loss, with a smaller initial pulse of C loss from fine roots (S₁k₁) and a slower decay rate of the recalcitrant C pool for fine roots (0.03 y⁻¹) compared with (0.19 y⁻¹) for needles. Soil ¹³C respiration, representing heterotrophic respiration of litter C, was much more seasonal from the O horizon than from the A. However, offsetting seasonal patterns in ¹³C dynamics in the O horizon resulted in no net effect of soil depth on total ¹³C retention in the soil after 1.5 y for either litter. Almost 90% of applied litter N was retained in the soil after 1.5 y, independent of litter quality or soil depth. Very small amounts of ¹³C or ¹⁵N (<3% of applied) moved to the horizon above or below the placement depth (i.e., O to A or A to O). Our results suggest that plant allocation belowground to fine roots results in more C retained and less N mineralized compared with allocation aboveground to needles, primarily due to litter quality differences.     

Response of soil respiration to simulated N deposition in a disturbed and a rehabilitated tropical forest in southern China

Responses of soil respiration (CO₂ emission) to simulated N deposition were studied in a disturbed (reforested forest with previous understory and litter harvesting) and a rehabilitated (reforested forest with no understory and litter harvesting) tropical forest in southern China from October 2005 to September 2006. The objectives of the study were to test the following hypotheses: (1) soil respiration is higher in rehabilitated forest than in disturbed forest; (2) soil respiration in both rehabilitated and disturbed tropical forests is stimulated by N additions; and (3) soil respiration is more sensitive to N addition in disturbed forest than in rehabilitated forest due to relatively low soil nutrient status in the former, resulting from different previous human disturbance. Static chamber and gas chromatography techniques were employed to quantify the soil respiration, following different N treatments (Control, no N addition; Low-N, 5 g N m⁻² year⁻¹; Medium-N, 10 g N m⁻² year⁻¹), which had been applied continuously for 26 months before the respiration measurement. Results showed that soil respiration exhibited a strong seasonal pattern, with the highest rates observed in the hot and wet growing season (April–September) and the lowest rates in winter (December–February) in both rehabilitated and disturbed forests. Soil respiration rates exhibited significant positive exponential relationship with soil temperature and significant positive linear relationship with soil moisture. Soil respiration was also significantly higher in the rehabilitated forest than in the disturbed forest. Annual mean soil respiration rate in the rehabilitated forest was 20% lower in low-N plots (71 ± 4 mg CO₂-C m⁻² h⁻¹) and 10% lower in medium-N plots (80 ± 4 mg CO₂-C m⁻² h⁻¹) than in the control plots (89 ± 5 mg CO₂-C m⁻² h⁻¹), and the differences between the control and low-N or medium-N treatments were statistically significant. In disturbed forest, annual mean soil respiration rate was 5% lower in low-N plots (63 ± 3 mg CO₂-C m⁻² h⁻¹) and 8% lower in medium-N plots (61 ± 3 mg CO₂-C m⁻² h⁻¹) than in the control plots (66 ± 4 mg CO₂-C m⁻² h⁻¹), but the differences among treatments were not significant. The depressed effects of experimental N deposition occurred mostly in the hot and wet growing season. Our results suggest that response of soil respiration to elevated N deposition in the reforested tropical forests may vary depending on the status of human disturbance. Responsible Editor: Hans Lambers.     

Nitrogen sequestration capacity of two salt marshes from the Tagus estuary

The role of salt marshes as nitrogen sink is examined taking into consideration the seasonal variation of above and belowground biomass of Spartina martima and Halimione portulacoides in two marshes from Tagus estuary, Pancas and Corroios, and the degradation rates of belowground litter. Total nitrogen was determined in plant components, decomposing litter and sediment. Biomass was higher in Corroios, the saltier marsh, with 7190 g m⁻² y⁻¹ dw of S. maritima and 6593 g m⁻² y⁻¹ dw of H. portulacoides and the belowground component contributed to 96% and 90% of total biomass, respectively. In the other marsh, Pancas, belowground biomass contributed to 56% and 76% of total biomass for S. maritima and H. portulacoides, respectively. Litterbag experiment showed that between 25% and 50% of nitrogen is lost within the first month and remained relatively constant in the next four months. Slower decomposition is observed in sediments with higher nitrogen concentration (max. 0.7% N in the saltier marsh). Higher concentrations of N were found in the sediment upper layers. Considering the sediment-root system, most of the nitrogen is stored in the sediment compartment and only about 1–4% of the total N was found in the roots. Considering these results, Tagus salt marshes act as a sink for nitrogen.     

Estimation of autotrophic and heterotrophic components of soil respiration by trenching is sensitive to corrections for root decomposition and changes in soil water content

This study aims to assess the effects of corrections for disturbances such as an increased amount of dead roots and an increase in volumetric soil water content on the calculation of soil CO₂ efflux partitioning. Soil CO₂ efflux, soil temperature and superficial soil water content were monitored in two young beech sites (H1 and H2) during a trenching experiment. Trenching induced a significant input of dead root mass that participated in soil CO₂ efflux and reduced the soil dissolved organic carbon content, while it increased superficial soil water content within the trenched plot. Annual soil CO₂ efflux in control plots was 528 g C m⁻² year⁻¹ at H1 and 527 g C m⁻² year⁻¹ at H2. The annual soil CO₂ efflux in trenched plots was 353 g C m⁻² year⁻¹ at H1 and 425 g C m⁻² year⁻¹ at H2. By taking into account annual CO₂ efflux from decaying trenched roots, the autotrophic contribution to total soil CO₂ efflux reached 69% at H1 and 54% at H2. The partitioning calculation was highly sensitive to the initial root mass estimated within the trenched plots. Uncertainties in the remaining root mass, the fraction of root C that is incorporated into soil organic matter during root decomposition, and the root decomposition rate constant had a limited impact on the partitioning calculation. Corrections for differences in superficial soil water content had a significant impact on annual respired CO₂ despite a limited effect on partitioning.     

Measurements and Modeling of Carbon and Nitrogen Cycling in Agroecosystems of Southern Wisconsin: Potential for SOC Sequestration during the Next 50 Years

Landmanagement practices such as no-tillage agriculture and tallgrass prairie restoration have been proposed as a possible means to sequester atmospheric carbon, helping to refurbish soil fertility and replenish organic matter lost as a result of previous agricultural management practices. However, the relationship between land-use changes and ecosystem structure and functioning is not yet understood. We studied soil and vegetation properties over a 4-year period (1995–98), and assembled measurements of microbial biomass, soil organic carbon (SOC) and nitrogen (N), N-mineralization, soil surface carbon dioxide (CO₂) flux, and leached C and N in managed (maize; Zea mays L.) and natural (prairie) ecosystems near the University of Wisconsin Agricultural Research Station at Arlington. Field data show that different management practices (tillage and fertilization) and ecosystem type (prairie vs maize) have a profound influence on biogeochemistry and water budgets between sites. These measurements were used in conjunction with a dynamic terrestrial ecosystem model, called IBIS (the Integrated Biosphere Simulator), to examine the long-term effects of land-use changes on biogeochemical cycling. Field data and modeling suggest that agricultural land management near Arlington between 1860 and 1950 caused SOC to be depleted by as much as 63% (native SOC approximately 25.1 kg C m⁻²). Reductions in N-mineralization and microbial biomass were also observed. Although IBIS simulations depict SOC recovery in no-tillage maize since the 1950s and also in the Arlington prairie since its restoration was initiated in 1976, field data suggest otherwise for the prairie. This restoration appears to have done little to increase SOC over the past 24 years. Measurements show that this prairie contained between 28% and 42% less SOC (in the top 1 m) than the no-tillage maize plots and 40%–47% less than simulated potential SOC for the site in 1999. Because IBIS simulates competition between C3 and C4 grass species, we hypothesized that current restored prairies, which include many forbs not characterized by the model, could be less capable of sequestering C than agricultural land planted entirely in monocultural grass in this region. Model output and field measurements show a potential 0.4 kg C m⁻² y⁻¹ difference in prairie net primary production (NPP). This study indicates that high-productivity C4 grasslands (NPP = 0.63 kg C m⁻² y⁻¹) and high-yield maize agroecosystems (10 Mg ha⁻¹) have the potential to sequester C at a rate of 74.5 g C m⁻² y⁻¹ and 86.3 g C m⁻² y⁻¹, respectively, during the next 50 years across southern Wisconsin. Received 28 December 1999; accepted 11 December 2000.     

Pools and fluxes of carbon in three Norway spruce ecosystems along a climatic gradient in Sweden

This paper presents an integrated analysis of organic carbon (C) pools in soils and vegetation, within-ecosystem fluxes and net ecosystem exchange (NEE) in three 40-year old Norway spruce stands along a north-south climatic gradient in Sweden, measured 2001–2004. A process-orientated ecosystem model (CoupModel), previously parameterised on a regional dataset, was used for the analysis. Pools of soil organic carbon (SOC) and tree growth rates were highest at the southernmost site (1.6 and 2.0-fold, respectively). Tree litter production (litterfall and root litter) was also highest in the south, with about half coming from fine roots (<1 mm) at all sites. However, when the litter input from the forest floor vegetation was included, the difference in total litter input rate between the sites almost disappeared (190–233 g C m⁻² year⁻¹). We propose that a higher N deposition and N availability in the south result in a slower turnover of soil organic matter than in the north. This effect seems to overshadow the effect of temperature. At the southern site, 19% of the total litter input to the O horizon was leached to the mineral soil as dissolved organic carbon, while at the two northern sites the corresponding figure was approx. 9%. The CoupModel accurately described general C cycling behaviour in these ecosystems, reproducing the differences between north and south. The simulated changes in SOC pools during the measurement period were small, ranging from −8 g C m⁻² year⁻¹ in the north to +9 g C m⁻² year⁻¹ in the south. In contrast, NEE and tree growth measurements at the northernmost site suggest that the soil lost about 90 g C m⁻² year⁻¹. An erratum to this article can be found at     

Pine Forest Floor Carbon Accumulation in Response to N and PK Additions: Bomb <Superscript>14</Superscript>C Modelling and Respiration Studies

The addition of nitrogen via deposition alters the carbon balance of temperate forest ecosystems by affecting both production and decomposition rates. The effects of 20 years of nitrogen (N) and phosphorus and potassium (PK) additions were studied in a 40-year-old pine stand in northern Sweden. Carbon fluxes of the forest floor were reconstructed using a combination of data on soil ¹⁴C, tree growth, and litter decomposition. N-only additions caused an increase in needle litterfall, whereas both N and PK additions reduced long-term decomposition rates. Soil respiration measurements showed a 40% reduction in soil respiration for treated compared to control plots. The average age of forest floor carbon was 17 years. Predictions of future soil carbon storage indicate an increase of around 100% in the next 100 years for the N plots and 200% for the NPK plots. As much as 70% of the increase in soil carbon was attributed to the decreased decomposition rate, whereas only 20% was attributable to increased litter production. A reduction in decomposition was observed at a rate of N addition of 30 kg C ha^–1 y^–1, which is not an uncommon rate of N deposition in central Europe. A model based on the continuous-quality decomposition theory was applied to interpret decomposer and substrate parameters. The most likely explanations for the decreased decomposition rate were a fertilizer-induced increase in decomposer efficiency (production-to-assimilation ratio), a more rapid rate of decrease in litter quality, and a decrease in decomposer basic growth rate.     

Detrital Controls on Soil Solution N and Dissolved Organic Matter in Soils: A Field Experiment

We established a long-term field study in an old growth coniferous forest at the H.J. Andrews Experimental Forest, OR, USA, to address how detrital quality and quantity control soil organic matter accumulation and stabilization. The Detritus Input and Removal Treatments (DIRT) plots consist of treatments that double leaf litter, double woody debris inputs, exclude litter inputs, or remove root inputs via trenching. We measured changes in soil solution chemistry with depth, and conducted long-term incubations of bulk soils from different treatments in order to elucidate effects of detrital inputs on the relative amounts and lability of different soil C pools. In the field, the addition of woody debris increased dissolved organic carbon (DOC) concentrations in O-horizon leachate and at 30 cm, but not at 100 cm, compared to control plots, suggesting increased rates of DOC retention with added woody debris. DOC concentrations decreased through the soil profile in all plots to a greater degree than did dissolved organic nitrogen (DON), most likely due to preferential sorption of high C:N hydrophobic dissolved organic matter (DOM) in upper horizons; percent hydrophobic DOM decreased significantly with depth, and hydrophilic DOM had a much lower and less variable C:N ratio. Although laboratory extracts of different litter types showed differences in DOM chemistry, percent hydrophobic DOM did not differ among soil solutions from different detrital treatments in the field, suggesting that microbial processing of DOM leachate in the field consumed easily degradable components, thus equalizing leachate chemistry among treatments. Total dissolved N leaching from plots with intact roots was very low (0.17 g m⁻² year⁻¹), slightly less than measured deposition to this very unpolluted forest (~s 0.2 g m⁻² year⁻¹). Total dissolved N losses showed significant increases in the two treatments without roots whereas concentrations of DOC decreased. In these plots, N losses were less than half of estimated plant uptake, suggesting that other mechanisms, such as increased microbial immobilization of N, accounted for retention of N in deep soils. In long-term laboratory incubations, soils from plots that had both above- and below-ground litter inputs excluded for 5 years showed a trend towards lower DOC loss rates, but not lower respiration rates. Soils from plots with added wood had similar respiration and DOC loss rates as control soils, suggesting that the additional DOC sorption observed in the field in these soils was stabilized in the soil and not readily lost upon incubation.