Nutrient Budget in the Seasonal Wetland of the Okavango Delta, Botswana

The nutrient (P and N species) and chloride budgets were investigated in a representative floodplain in the seasonal wetlands of the Okavango Delta, Botswana. A variety of sources of nutrients in the surface water were considered, namely ion species coming with the floodwater, those generated from dry floodplain soils and those from water-soluble dust deposition (both local and long-range sources). Concentrations of total-nitrogen and chloride in surface water were below 1 mg l⁻¹. Total-phosphorus concentrations were 0.05 mg l⁻¹, reflecting the oligotrophic character of the system. Dust deposition rates were highest for chloride at 2.44 g m⁻² year⁻¹ followed by 0.79 g m⁻² year⁻¹ for total-N, 0.40 g m⁻² year⁻¹ for ammonia and only 0.02 g m⁻² year⁻¹ for total-P, respectively. Chloride was derived primarily from long-range transport, while N and P species were of more local origin. Dissolution rates for these ions combined were calculated to be 3.9 g m⁻² for the flooded area in the 1999 season and thus all dry deposits must be re-dissolved. The accumulation of dust deposits on dry surfaces and their subsequent dissolution causes 2–5 times higher concentrations of nitrogen, phosphorus and chloride with the onset of the flood, thus boosting the nutrient stock in the crucial phase of the onset of flooding. Chloride dissolved from dry soil surfaces and dust contributed approximately 40% to the overall floodplain budget. Although contributions from the soil surface and dust to the nitrogen and phosphorus pools of the floodplain are less prominent (with 10% of total), they nonetheless represent a significant source of nutrients in the entire system. Extrapolation to annually flooded swamps (10,000 km²) indicates a maximum contribution of 40% for total-nitrogen and 60% for total-phosphorus from dust deposition on wet or dry surfaces to the nutrient pool of the water body.     

Temporal variation in CO<Subscript>2</Subscript> efflux from soil and snow surfaces in a Japanese cedar (<Emphasis Type="Italic">Cryptomeria japonica</Emphasis>) plantation,central Japan

CO₂ efflux from soil and snow surfaces was measured continuously in a Japanese cedar (Cryptomeria japonica D. Don) forest in central Japan using an open dynamic chamber system. The chamber opens and closes automatically and records measurements based on an open-flow dynamic method. Between May and December, mean soil CO₂ efflux ranged from 1,529 mg CO₂ m⁻² h⁻¹ in September to 255 mg CO₂ m⁻² h⁻¹ in December. The seasonal change in CO₂ efflux from the soil paralleled the seasonal pattern of soil temperature. No marked diurnal trends in soil CO₂ efflux were observed on days without rainfall, whereas significant pulses in soil CO₂ efflux were observed on days with rainfall. In this plantation, soil CO₂ efflux frequently responded to rainfall. Measurements of changes from litter-covered soil to snow-covered surfaces revealed that CO₂ efflux decreased from values of ca. 250 mg CO₂ m⁻² h⁻¹ above soil to less than 33 mg CO₂ m⁻² h⁻¹ above snow. Soil temperature alone explained 66% of the overall variation in soil CO₂ efflux, but explained approximately 85% of the variation when data from two anomalous periods were excluded. Moreover, we found a significant correlation between soil CO₂ efflux and soil moisture (which explained 44% of the overall variation) using a second-order polynomial function. Our results suggest that the seasonality of CO₂ efflux is affected not only by soil temperature and moisture, but also by drying and rewetting cycles and by litterfall pulses.     

Stimulated N2O flux from intact grassland monoliths after two growing seasons under elevated atmospheric CO2

Long-term exposure of native vegetation to elevated atmospheric CO₂ concentrations is expected to increase C inputs to the soil and, in ecosystems with seasonally dry periods, to increase soil moisture. We tested the hypothesis that these indirect effects of elevated CO₂ (600 μl l⁻¹ vs 350 μl l⁻¹) would improve conditions for microbial activity and stimulate emissions of nitrous oxide (N₂O), a very potent and long-lived greenhouse gas. After two growing seasons, the mean N₂O efflux from monoliths of calcareous grassland maintained at elevated CO₂ was twice as high as that measured from monoliths maintained at current ambient CO₂ (70 ± 9 vs 37 ± 4 μg N₂O m⁻² h⁻¹ in October, 27 ± 5 vs 13 ± 3 μg N₂O m⁻² h⁻¹ in November after aboveground harvest). The higher N₂O emission rates at elevated CO₂ were associated with increases in soil moisture, soil heterotrophic respiration, and plant biomass production, but appear to be mainly attributable to higher soil moisture. Our results suggest that rising atmospheric CO₂ may contribute more to the total greenhouse effect than is currently estimated because of its plant-mediated effects on soil processes which may ultimately lead to increased N₂O emissions from native grasslands. Received: 11 September 1997 / Accepted: 20 March 1998     

Emissions of nitrous oxide from three tropical forests in Southern China in response to simulated nitrogen deposition

Emissions of nitrous oxide (N₂O) from the soil following simulated nitrogen (N) deposition in a disturbed (pine), a rehabilitated (pine and broadleaf mixed) and a mature (monsoon evergreen broadleaf) tropical forest in southern China were studied. The following hypotheses were tested: (1) addition of N will increase soil N₂O emission in tropical forests; and (2) any observed increase will be more pronounced in the mature forest than in the disturbed or rehabilitated forest due to the relatively high initial soil N concentration in the mature forest. The experiment was designed with four N treatment levels (three replicates; 0, 50, 100, 150 kg N ha⁻¹ year⁻¹ for C (Control), LN (Low-N), MN (Medium-N), and HN (High-N) treatment, respectively) in the mature forest, but only three levels in the disturbed and rehabilitated forests (C, LN and MN). Between October 2005 to September 2006, soil N₂O flux was measured using static chamber and gas chromatography methodology. Nitrogen had been applied previously to the plots since July 2003 and continued during soil N₂O flux measurement period. The annual mean rates of soil N₂O emission in the C plots were 24.1 ± 1.5, 26.2 ± 1.4, and 29.3 ± 1.6 μg N₂O–N m⁻² h⁻¹ in the disturbed, rehabilitated and mature forest, respectively. There was a significant increase in soil N₂O emission following N additions in the mature forest (38%, 41%, and 58% when compared to the C plots for the LN, MN, and HN plots, respectively). In the disturbed forest a significant increase (35%) was observed in the MN plots, but not in the LN plots. The rehabilitated forest showed no significant response to N additions. Increases in soil N₂O emission occurred primarily in the cool-dry season (November, December and January). Our results suggest that the response of soil N₂O emission to N deposition in tropical forests in southern China may vary depending on the soil N status and land-use history of the forest.     

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.     

On-line screening of soil VOCs exchange responses to moisture, temperature and root presence

The exchanges of volatile organic compounds (VOCs) between soils and the atmosphere are poorly known. We investigated VOC exchange rates and how they were influenced by soil moisture, temperature and the presence of plant roots in a Mediterranean forest soil. We measured VOC exchange rates along a soil moisture gradient (5%–12.5%–20%–27.5% v/v) and a temperature gradient (10°C–15°C–25°C–35°C) using PTR-MS. Monoterpenes were identified with GC-MS. Soils were a sink rather than a source of VOCs in both soil moisture and temperature treatments (−2.16 ± 0.35 nmol m⁻² s⁻¹ and −4.90 ± 1.24 nmol m⁻² s⁻¹ respectively). Most compounds observed were oxygenated VOCs like alcohols, aldehydes and ketones and aromatic hydrocarbons. Other volatiles such as acetic acid and ethyl acetate were also observed. All those compounds had very low exchange rates (maximum uptake rates from −0.8 nmol m⁻² s⁻¹ to −0.6 nmol m⁻² s⁻¹ for methanol and acetic acid). Monoterpene exchange ranged only from −0.004 nmol m⁻² s⁻¹ to 0.004 nmol m⁻² s⁻¹ and limonene and α-pinene were the most abundant compounds. Increasing soil moisture resulted in higher soil sink activity possibly due to increases in microbial VOCs uptake activity. No general pattern of response was found in the temperature gradient for total VOCs. Roots decreased the emission of many compounds under increasing soil moisture and under increasing soil temperature. While our results showed that emission of some soil VOCs might be enhanced by the increases in soil temperature and that the uptake of most soil VOCs uptake might be reduced by the decreases of soil water availability, the low exchange rates measured indicated that soil-atmosphere VOC exchange in this system are unlikely to play an important role in atmospheric chemistry. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

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.     

Development of ground vegetation biomass and nutrient pools in a clear-cut disc-plowed boreal forest

Nutrient leaching from forest substrate after clear-cutting and subsequent soil preparation is strongly influenced by the capacity of ground vegetation to sequester the released nutrients. We studied the rates and patterns of biomass and nutrient accumulation in ground vegetation growing on ridges, in furrows and on undisturbed surfaces for 2–5 years after disc-plowing in eastern Finland. The biomass of mosses on ridges remained significantly lower than that in furrows and on undisturbed surfaces. Field layer biomass on ridges and in furrows was significantly lower than on undisturbed surfaces throughout the study period. Field layer biomass increased more on ridges than in furrows. Root biomass on ridges and undisturbed surfaces was considerably higher than in furrows. Five years after disc-plowing, total biomass and nutrient pools for ridges (biomass 4,975 kg ha⁻¹, N 40 kg ha⁻¹, P 5 kg ha⁻¹, K 20 kg ha⁻¹ and Ca 18 kg ha⁻¹) and undisturbed surfaces (biomass 5,613 kg ha⁻¹, N 43 kg ha⁻¹, P 5 kg ha⁻¹, K 22 kg ha⁻¹ and Ca 18 kg ha⁻¹) were similar, but considerably lower for furrows (biomass 1,807 kg ha⁻¹, N 16 kg ha⁻¹, P 2 kg ha⁻¹, K 10 kg ha⁻¹ and Ca 6 kg ha⁻¹). Ridges covered 25% of the area, furrows 30 and 45% was undisturbed surfaces. Taking into account the proportion of each type of surface, values for the whole prepared clear-cut area were 4,312, 34, 4, 18 and 14 kg ha⁻¹ for biomass, N, P, K and Ca, respectively. Biomass and nutrient pools had not returned to uncut forest levels at the end of the 5-year study period. The results indicate that mosses and field layer vegetation respond differently to soil preparation, that the development of biomass on ridges, in furrows and on undisturbed surfaces proceeds at different rates, and that the biomass and nutrient uptake of ground vegetation remains below pre-site preparation levels for several years. However, ridges, which are known to be the most susceptible to leaching, revegetate rapidly. Responsible Editor: Tibor Kalapos.     

Effect of Watering and Soil Moisture on Mercury Emissions from Soils

This paper presents data from experiments that measured Mercury (Hg) flux as a function of water addition and subsequent soil drying, and maintenance of soil water content over time utilizing small dynamic gas exchange chambers and large mesocosms. When soil surfaces were dry and water was added at an amount less than that necessary to saturate the soil an immediate large (relative to dry soil flux) release of Hg occurred. Diel Hg emissions from soils, unenriched (0.02 μg g⁻¹) and enriched (3 μg g⁻¹) in Hg and wet below saturation, were significantly elevated above that occurring from dry soils (2–5 times depending on soil water content) for weeks to months. Enhancement of emissions from wet soils in direct sunlight were greater than that from soils shaded or in the dark suggesting that a synergism exists between soil moisture and light. When soils were watered to saturation Hg emissions were suppressed or remained the same depending on the degree of saturation. It is hypothesized that the addition of soil water in amounts less than that necessary to saturate the soil surface results in an immediate release of elemental Hg from soil surface as the more polar water molecule out competes Hg for binding sites. As the water moves into the soil, Hg adsorbed to soil particles is desorbed into soil gas and dissolved in the soil water. The process of evaporation facilitates movement of Hg as mass flow to the soil surface where it is made available for subsequent release. The latter is hypothesized to be an important process by which Hg is recharged at the soil–air interface.     

Spatial Patterns of Soil Surface C Flux in Experimental Canopy Gaps

To explore within-gap spatial patterns of soil surface CO₂ flux, we measured instantaneous soil surface CO₂ flux, soil surface temperature, and soil moisture in north–south transects across canopy gaps and in adjacent contiguous forest from April to November 2010 in a second-growth northern hardwood forest in Wisconsin, USA. Throughout the growing season, soil surface CO₂ flux was higher in the northern 1/3 and northern edge of gaps compared to the central and southern portions. These patterns were driven primarily by within-gap variation in soil temperature, which was itself driven by within-gap patterns of insolation. Most locations in the northern 1/3 and northern edge of gaps had significantly higher modeled total growing season C flux (mean 725 g C m⁻²) compared to the contiguous forest (mean 706 g C m⁻²), whereas C flux in the central and southern portions of gaps (mean 555 g C m⁻²) was significantly lower than both the contiguous forest and the northern portions of gaps.     

N2O, CH4 and CO2 emissions from seasonal tropical rainforests and a rubber plantation in Southwest China

The main focus of this study was to evaluate the effects of soil moisture and temperature on temporal variation of N₂O, CO₂ and CH₄ soil-atmosphere exchange at a primary seasonal tropical rainforest (PF) site in Southwest China and to compare these fluxes with fluxes from a secondary forest (SF) and a rubber plantation (RP) site. Agroforestry systems, such as rubber plantations, are increasingly replacing primary and secondary forest systems in tropical Southwest China and thus effect the N₂O emission in these regions on a landscape level. The mean N₂O emission at site PF was 6.0 ± 0.1 SE μg N m⁻² h⁻¹. Fluxes of N₂O increased from <5 μg N m⁻² h⁻¹ during dry season conditions to up to 24.5 μg N m⁻² h⁻¹ with re-wetting of the soil by the onset of first rainfall events. Comparable fluxes of N₂O were measured in the SF and RP sites, where mean N₂O emissions were 7.3 ± 0.7 SE μg N m⁻² h⁻¹ and 4.1 ± 0.5 SE μg N m⁻² h⁻¹, respectively. The dependency of N₂O fluxes on soil moisture levels was demonstrated in a watering experiment, however, artificial rainfall only influenced the timing of N₂O emission peaks, not the total amount of N₂O emitted. For all sites, significant positive correlations existed between N₂O emissions and both soil moisture and soil temperature. Mean CH₄ uptake rates were highest at the PF site (−29.5 ± 0.3 SE μg C m⁻² h⁻¹), slightly lower at the SF site (−25.6 ± 1.3 SE μg C m⁻² h⁻¹) and lowest for the RP site (−5.7 ± 0.5 SE μg C m⁻² h⁻¹). At all sites, CH₄ uptake rates were negatively correlated with soil moisture, which was also reflected in the lower uptake rates measured in the watering experiment. In contrast to N₂O emissions, CH₄ uptake did not significantly correlate with soil temperature at the SF and RP sites, and only weakly correlated at the PF site. Over the 2 month measurement period, CO₂ emissions at the PF site increased significantly from 50 mg C m⁻² h⁻¹ up to 100 mg C m⁻² h⁻¹ (mean value 68.8 ± 0.8 SE mg C m⁻² h⁻¹), whereas CO₂ emissions at the SF and RP site where quite stable and varied only slightly around mean values of 38.0 ± 1.8 SE mg C m⁻² h⁻¹ (SF) and 34.9 ± 1.1 SE mg C m⁻² h⁻¹ (RP). A dependency of soil CO₂ emissions on changes in soil water content could be demonstrated for all sites, thus, the watering experiment revealed significantly higher CO₂ emissions as compared to control chambers. Correlation of CO₂ emissions with soil temperature was significant at the PF site, but weak at the SF and not evident at the RP site. Even though we demonstrated that N and C trace gas fluxes significantly varied on subdaily and daily scales, weekly measurements would be sufficient if only the sink/ source strength of non-managed tropical forest sites needs to be identified.     

Litterfall and fine root biomass contribution to nutrient dynamics in second- and old-growth Douglas-fir ecosystems

Litterfall and fine root production were measured for three years as part of a carbon balance study of three forest stands in the Pacific Northwest of the United States. A young second-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] stand, a second-growth Douglas-fir with red alder (Alnus rubra Bong.) stand, and an old-growth (∼550 years) Douglas-fir stand were monitored for inputs of carbon and nitrogen into the soil from litterfall and fine root production, as well as changes in soil C and N. Fine root production and soil nutrient changes were measured through the use of soil ingrowth bags containing homogenized soil from the respective stands. Litterfall biomass was greatest in the Douglas-fir-alder stand (527 g m⁻² yr⁻¹) that annually returned nearly three times the amount of N as the other stands. Mean residence time for forest floor material was also shortest at this site averaging 4.6 years and 5.5 years for C an N, respectively. Fine root production in the upper 20 cm ranged from 584 g m⁻² in the N rich Douglas-fir-alder stand to 836 g m⁻² in the old-growth stand. Fine root production (down to one meter) was always greater than litterfall with a below:above ratio ranging from 3.73 for the young Douglas-fir stand to 1.62 for the Douglas-fir-alder stand. The below:above N ratios for all three stands closely approximate those for biomass. Soil changes in both C and N differed by site, but the soil C changes in the old-growth stand mirrored those obtained in an ongoing CO₂ flux study. Results from the soil ingrowth bags strongly suggest that this method provides a simple, but sufficient device for measuring potential fine root biomass production as well as soil chemical changes.     

The effects of gap disturbance on nitrogen cycling and retention in late-successional northern hardwood–hemlock forests

Late-successional forests in the upper Great Lakes region are susceptible to nitrogen (N) saturation and subsequent nitrate (NO₃⁻) leaching loss. Endemic wind disturbances (i.e., treefall gaps) alter tree uptake and soil N dynamics; and, gaps are particular susceptible to NO₃⁻ leaching loss. Inorganic N was measured throughout two snow-free periods in throughfall, forest floor leachates, and mineral soil leachates in gaps (300–2,000 m², 6–9 years old), gap-edges, and closed forest plots in late-successional northern hardwood, hemlock, and northern hardwood–hemlock stands. Differences in forest water inorganic N among gaps, edges, and closed forest plots were consistent across these cover types: NO₃⁻ inputs in throughfall were significantly greater in undisturbed forest plots compared with gaps and edges; forest floor leachate NO₃⁻ was significantly greater in gaps compared to edges and closed forest plots; and soil leachate NO₃⁻ was significantly greater in gaps compared to the closed forest. Significant differences in forest water ammonium and pH were not detected. Compared to suspected N-saturated forests with high soil NO₃⁻ leaching, undisturbed forest plots in these late-successional forests are not losing NO₃⁻ (net annual gain of 2.8 kg ha⁻¹) and are likely not N-saturated. Net annual NO₃⁻ losses were observed in gaps (1.3 kg ha⁻¹) and gap-edges (0.2 kg ha⁻¹), but we suspect these N leaching losses are a result of decreased plant uptake and increased soil N mineralization associated with disturbance, and not N-saturation.     

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.     

Biomass allocation and canopy development in spruce model ecosystems under elevated CO2 and increased N deposition

Ecosystem-level experiments on the effects of atmospheric CO₂ enrichment and N deposition on forest trees are urgently needed. Here we present data for nine model ecosystems of spruce (Picea abies) on natural nutrient-poor montane forest soil (0.7 m² of ground and 350 kg weight). Each system was composed of six 7-year-old (at harvest) trees each representing a different genotype, and a herbaceous understory layer (three species). The model ecosystems were exposed to three different CO₂ concentrations (280, 420, 560 μl l⁻¹) and three different rates of wet N deposition (0, 30, 90 kg ha⁻¹ year⁻¹) in a simulated annual course of Swiss montane climate for 3 years. The total ecosystem biomass was not affected by CO₂ concentration, but increased with increasing N deposition. However, biomass allocation to roots increased with increasing CO₂ leading to significantly lower leaf mass ratios (LMRs) and leaf area ratios (LARs) in trees grown at elevated CO₂. In contrast to CO₂ enrichment, N deposition increased biomass allocation to the aboveground plant parts, and thus LMR and LAR were higher with increasing N deposition. We observed no CO₂ ×  N interactions on growth, biomass production, or allocation, and there were also no genotype × treatment interactions. The final leaf area index (LAI) of the spruce canopies was 19% smaller at 420 and 27% smaller at 560 than that measured at 280 μl CO₂ l⁻¹, but was not significantly altered by increasing N deposition. Lower LAIs at elevated CO₂ largely resulted from shorter branches (less needles per individual tree) and partially from increased needle litterfall. Independently of N deposition, total aboveground N content in the spruce communities declined with increasing CO₂ (−18% at 420 and −31% at 560 compared to 280 μl CO₂ l⁻¹). N deposition had the opposite effect on total above ground N content (+18% at 30 and +52% at 90 compared to 0 kg N ha⁻¹ year⁻¹). Our results suggest that under competitive conditions on natural forest soil, atmospheric CO₂ enrichment may not lead to higher ecosystem biomass production, but N deposition is likely to do so. The reduction in LAI under elevated CO₂ suggests allometric down-regulation of photosynthetic carbon uptake at the canopy level. The strong decline in the tree nitrogen mass per unit ground area in response to elevated CO₂ may indicate CO₂-induced reductions of soil N availability. Received: 11 May 1997 / Accepted: 4 August 1997     

Decrease of phosphoribulokinase activity by antisense RNA in transgenic tobacco: definition of the light environment under which phosphoribulokinase is not in large excess

 To test the hypothesis that the contribution of phosphoribulokinase (PRK) to the control of photosynthesis changes depending on the light environment of the plant, the response of transgenic tobacco (Nicotiana tabacum L.) transformed with antisense PRK constructs to irradiance was determined. In plants grown under low irradiance (330 μmol m⁻² s⁻¹) steady-state photosynthesis was limited in plants with decreased PRK activity upon exposure to higher irradiance, with a control coefficient of PRK for CO₂ assimilation of 0.25 at and above 800 μmol m⁻² s⁻¹. The flux control coefficient of PRK for steady-state CO₂ assimilation was zero, however, at all irradiances in plant material grown at 800 μmol m⁻² s⁻¹ and in plants grown in a glasshouse during mid-summer (alternating shade and sun 300–1600 μmol m⁻² s⁻¹). To explain these differences between plants grown under low and high irradiances, Calvin cycle enzyme activities and metabolite content were determined. Activities of PRK and other non-equilibrium Calvin cycle enzymes fructose-1,6-bisphosphatase, sedoheptulose-1,7-bisphosphatase and ribulose-1,5-bisphosphate carboxylase-oxygenase were twofold higher in plants grown at 800 μmol m⁻² s⁻¹ or in the glasshouse than in plants grown at 330 μmol m⁻² s⁻¹. Activities of equilibrium enzymes transketolase, aldolase, ribulose-5-phosphate epimerase and isomerase were very similar under all growth irradiances. The flux control coefficient of 0.25 in plants grown at 330 μmol m⁻² s⁻¹ can be explained because low ribulose-5-phosphate content in combination with low PRK activity limits the synthesis of ribulose-1,5-bisphosphate. This limitation is overcome in high-light-grown plants because of the large relative increase in activities of sedoheptulose-1,7-bisphosphatase and fructose-1,6-bisphosphatase under these conditions, which facilitates the synthesis of larger amounts of ribulose-5-phosphate. This potential limitation will have maintained evolutionary selection pressure for high concentrations of PRK within the chloroplast. Received: 15 November 1999 / Accepted: 27 January 2000     

Characteristics of soil CO2 efflux variability in an aseasonal tropical rainforest in Borneo Island

Although soil carbon dioxide (CO₂) efflux from tropical forests may play an important role in global carbon (C) balance, our knowledge of the fluctuations and factors controlling soil CO₂ efflux in the Asian tropics is still poor. This study characterizes the temporal and spatial variability in soil CO₂ efflux in relation to temperature/moisture content and estimates annual efflux from the forest floor in an aseasonal intact tropical rainforest in Sarawak, Malaysia. Soil CO₂ efflux varied widely in space; the range of variation averaged 17.4 μmol m⁻² s⁻¹ in total. While most CO₂ flux rates were under 10 μmol m⁻² s⁻¹, exceptionally high fluxes were observed sporadically at several sampling points. Semivariogram analysis revealed little spatial dependence in soil CO₂ efflux. Temperature explained nearly half of the spatial heterogeneity, but the effect varied with time. Seasonal variation in CO₂ efflux had no fixed pattern, but was significantly correlated with soil moisture content. The correlation coefficient with soil moisture content (SMC) at 30 and 60 cm depth was higher than at 10 cm depths. The annual soil CO₂ efflux, estimated from the relationship between CO₂ efflux and SMC at 30 cm depth, was 165 mol m⁻² year⁻¹ (1,986 g C m⁻² year⁻¹). As this area is known to suffer severe drought every 4–5 years caused by the El Nino-Southern Oscillation, the results suggest that an unpredictable dry period might affect soil CO₂ efflux, leading an annual variation in soil C balance.     

Variation in leaf area index and stand leaf mass of European beech across gradients of soil acidity and precipitation

Leaf area index (LAI, the one-sided foliage area per unit ground surface area) is a key determinant of plant productivity which has a large influence on water and energy exchange between vegetation and the atmosphere. The variation in forest LAI across landscapes and environmental gradients and its causes are not sufficiently understood. We measured the LAI of European beech (Fagus sylvatica) by litter trapping in 23 closed, mature stands across gradients of rainfall and soil acidity or fertility. With a mean LAI of 7.4 m² m⁻² (minimum: 5.6, maximum: 9.5 m² m⁻²), beech stands maintained a comparably high leaf area index with relatively small variation along steep environmental gradients. Contrary to expectation, decreasing water availability (rainfall gradient from 1030 to 520 mm yr⁻¹) or increasing soil acidity (pH 3–7) had no significant effect on LAI. Stand leaf mass (M _l) increased slightly with soil fertility (C/N ratio, base saturation). We regressed parameters of site water availability (rainfall), soil fertility or acidity (pH, base saturation, C/N ratio, exchangeable Mg and Al content), and stand structure (stand age and stem density) against LAI and M _l in order to detect environmental controls of stand leaf area. Stand age was the most influential factor for both LAI and M _l (negative relationship). Stem density and the base saturation of the soil affected M _l significantly, but had a weak influence on LAI. We conclude that the leaf area index of beech is mainly under control of age-related physiological factors, whereas the influence of soil chemistry and rainfall is comparably low.     

Tree species and wood ash affect soil in Michigan’s Upper Peninsula

Tree species and wood ash application in plantations of short-rotation woody crops (SRWC) may have important effects on the soil productive capacity through their influence on soil organic matter (SOM) and exchangeable cations. An experiment was conducted to assess changes in soil C and N contents and pH within the 0–50 cm depth, and exchangeable cation (Ca²⁺, Mg²⁺, K⁺, and Na⁺) and extractable acidity concentrations within the 0–10 cm depth. The effects of different species (European larch [Larix decidua P. Mill.], aspen [Populus tremula L. × Populus tremuloides Michx.], and four poplar [Populus spp.] clones) and wood ash applications (0, 9, and 18 Mg ha⁻¹) on soil properties were evaluated, using a common garden experiment (N = 70 stands) over 7 years of management in Michigan’s Upper Peninsula. Soils were of the Onaway series (fine-loamy, mixed, active, frigid Inceptic Hapludalfs). The NM-6 poplar clone had the greatest soil C and N contents in almost all ash treatment levels. Soil C contents were 7.5, 19.4, and 10.7 Mg C ha⁻¹ greater under the NM-6 poplar than under larch in the ash-free, medium-, and high-level plots, respectively. Within the surface layer, ash application increased soil C and N contents (P < 0.05) through the addition of about 0.7 Mg C ha⁻¹ and 3 kg N ha⁻¹ with the 9 Mg ha⁻¹ ash application (twofold greater C and N amounts were added with the 18 Mg ha⁻¹ application). During a decadal time scale, tree species had no effects—except for K⁺—on the concentrations of the exchangeable cations, pH, and extractable acidity. In contrast, ash application increased soil pH and the concentration of Ca²⁺ (P < 0.05), from 5.2 ± 0.4 cmol_c kg⁻¹ (ash-free plots) to 8.6 ± 0.4 cmol_c kg⁻¹ (high-level ash plots), and tended to increase the concentration of Mg²⁺ (P < 0.1), while extractable acidity was reduced (P < 0.05) from 5.6 ± 0.2 cmol_c kg⁻¹ (ash-free plots) to 3.7 ± 0.2 cmol_c kg⁻¹ (high-level plots). Wood ash application, within certain limits, not only had a beneficial effect on soil properties important to the long-term productivity of fast-growing plantations but also enhanced long-term soil C sequestration.     

The organic carbon dynamics of a moorland catchment in N. W. England

The carbon cycle was quantified in the catchment of Doe House Gill, which drains high-relief moorland, with thin organic-rich soils (leptosols and podzols) 10–25 cm deep, in northern England. The soil C pool of 8,300 g m^-2 is due mainly to humic acid and older humin. If steady state is assumed, and a single soil C pool, the average ¹⁴C content of the whole soil (93% modern) yields a mean carbon residence time of 800 years, although this varied from 300 to 1,600 years in the four samples studied. Stream water fluxes of dissolved and particulate organic carbon (DOC, POC) were 2.5 and 0.4 g m⁻² a⁻¹ respectively in 2002–2003, lower than values for some other upland streams in the UK. The C pool, flux, and isotope data were used, with the assumption of steady state, to calibrate DyDOC, a model that simulates the soil carbon cycle, including the generation and transport of DOC. According to DyDOC, the litter pool (ca. 100 gC m⁻²) turns over quickly, and most (>90%) of the litter carbon is rapidly mineralised. The soil is calculated to gain only 16 gC m⁻² a⁻¹, and to lose the same amount, about 80% as CO₂ and 20% as DOC. From the DO¹⁴C content of 107.5% modern (due to “bomb carbon”) the model could be calibrated by assuming all DOC to come directly from litter, but DOC is more likely a mixture, derived from more than one soil C pool. The seasonal variability exhibited by stream water DOC concentration (maximum in September, minimum in January) is attributed mainly to variations in rainfall and evapotranspiration, rather than in the metabolic production rate of “potential DOC”. The model predicts that, for a Q ₁₀ of 2, the total soil organic C pool would decrease by about 5% if subjected to warming over 200 years. DyDOC predicts higher DOC fluxes in response to increased litter inputs or warming, and can simulate changes in DOC flux due to variations in sorption to soil solids, that might occur due to acidification and its reversal.