Anthropogenic nitrogen deposition enhances carbon sequestration in boreal soils

It is proposed that carbon (C) sequestration in response to reactive nitrogen (N_r) deposition in boreal forests accounts for a large portion of the terrestrial sink for anthropogenic CO₂ emissions. While studies have helped clarify the magnitude by which N_r deposition enhances C sequestration by forest vegetation, there remains a paucity of long‐term experimental studies evaluating how soil C pools respond. We conducted a long‐term experiment, maintained since 1996, consisting of three N addition levels (0, 12.5, and 50 kg N ha^?1 yr^?1) in the boreal zone of northern Sweden to understand how atmospheric N_r deposition affects soil C accumulation, soil microbial communities, and soil respiration. We hypothesized that soil C sequestration will increase, and soil microbial biomass and soil respiration will decrease, with disproportionately large changes expected compared to low levels of N addition. Our data showed that the low N addition treatment caused a non‐significant increase in the organic horizon C pool of ~15% and a significant increase of ~30% in response to the high N treatment relative to the control. The relationship between C sequestration and N addition in the organic horizon was linear, with a slope of 10 kg C kg^?1 N. We also found a concomitant decrease in total microbial and fungal biomasses and a ~11% reduction in soil respiration in response to the high N treatment. Our data complement previous data from the same study system describing aboveground C sequestration, indicating a total ecosystem sequestration rate of 26 kg C kg^?1 N. These estimates are far lower than suggested by some previous modeling studies, and thus will help improve and validate current modeling efforts aimed at separating the effect of multiple global change factors on the C balance of the boreal region.     

Reduction of forest floor respiration by fertilization on both carbon dioxide-enriched and reference 17-year-old loblolly pine stands

Elevated atmospheric carbon dioxide (CO₂^e) increases soil respiration rates in forest, grassland, agricultural and wetland systems as a result of increased growth, root biomass and enhanced biological activity of soil microorganisms. Less is known about how forest floor fluxes respond to the combined effects of elevated CO₂ and nutrient amendments; until now no experiments have been in place with large forest trees to allow even preliminary investigations. We investigated changes in forest floor respiration (S_ff) in a Pinus taeda L. plantation fumigated with CO₂ by using free‐air CO₂ enrichment (FACE) technology and given nutrient amendments. The prototype FACE apparatus (FACEp; 707 m²) was constructed in 1993, 10 years after planting, on a moderate fertility site in Duke Forest, North Carolina, USA, enriching the stand to 55 Pa (CO₂^e). A nearby ambient CO₂ (CO₂^a) plot (117 m²) was designated at the inception of the study as a reference (Ref). Both FACEp and Ref plot were divided in half and urea fertilizer was applied to one half at an annual rate of 11.2 g N m^?2 in the spring of 1998, 1999 and 2000. Forest floor respiration was monitored continuously for 220 days – March through November 2000 – by using two Automated Carbon Efflux Systems. Thirty locations (491 cm² each) were sampled in both FACEp and Ref, about half in each fertility treatment. Forest floor respiration was strongly correlated with soil temperature at 5 cm. Rates of S_ff were greater in CO₂^e relative to CO₂^a (an enhancement of ～178 g C m^?2) during the measurement period. Application of fertilizer resulted in a statistically significant depression of respiration rates in both the CO₂^a and CO₂^e plots (a reduction of ～186 g C m^?2). The results suggest that closed canopy forests on moderate fertility sites cycle back to the atmosphere more assimilated carbon (C) than similar forests on sites of high fertility. We recognize the limitations of this non‐replicated study, but its clear results offer strong testable hypotheses for future research in this important area.     

Long-term dynamics of aboveground fungal communities in a subalpine Norway spruce forest under elevated nitrogen input

As anthropogenic N deposition has been suspected to be the main reason for the decline of macromycetous sporocarp production in forest ecosystems, various N-fertilization experiments were started in the mid 1990s. The dynamics of ectomycorrhizal (root-inhabiting) and terricolous saprobic (litter-inhabiting) fungal communities were studied by exhaustive sporocarp inventories in a substitution Norway spruce (Picea abies) forest in two 256-m² plots sampled for periods of 1 week at 1-m² resolution between 1994 and 2007. N was added to the soil twice per year in one plot from the fourth year onwards. The effects of N input and time on aboveground fungal communities were assessed using redundancy analysis, principal response curves and non-parametric multivariate ANOVA. Results of this long-term experiment revealed that both ectomycorrhizal and saprobic fungal communities responded to an increase in soil N input. The ectomycorrhizal community reacted by a fast decrease in sporocarp production and in species richness, whereas the saprobic community was less affected. The response was highly species specific, especially for the saprobic community. The difference in species composition between control and fertilized plots was significant after 1 year of N addition for ectomycorrhizal fungi and only after 3 years for saprobic fungi. An aging effect affected sporocarp production in the whole area. For both communities, this unidirectional drift in species composition was as important as the treatment effect. This result highlights the importance of considering the respective role of treatment and year effects in long-term field experiments on fungal communities.     

Differential effects of warming and nitrogen fertilization on soil respiration and microbial dynamics in switchgrass croplands

The mechanistic understanding of warming and nitrogen (N) fertilization, alone or in combination, on microbially mediated decomposition is limited. In this study, soil samples were collected from previously harvested switchgrass (Panicum virgatum L.) plots that had been treated with high N fertilizer (HN: 67 kg N ha^?1) and those that had received no N fertilizer (NN) over a 3‐year period. The samples were incubated for 180 days at 15 °C and 20 °C, during which heterotrophic respiration, δ¹³C of CO₂, microbial biomass (MB), specific soil respiration rate (R_s: respiration per unit of microbial biomass), and exoenzyme activities were quantified at 10 different collections time. Employing switchgrass tissues (referred to as litter) with naturally abundant ¹³C allowed us to partition CO₂ respiration derived from soil and amended litter. Cumulative soil respiration increased significantly by 16.4% and 4.2% under warming and N fertilization, respectively. Respiration derived from soil was elevated significantly with warming, while oxidase, the agent for recalcitrant soil substrate decomposition, was not significantly affected by warming. Warming, however, significantly enhanced MB and R_s indicating a decrease in microbial growth efficiency (MGE). On the contrary, respiration derived from amended litter was elevated with N fertilization, which was consistent with the significantly elevated hydrolase. N fertilization, however, had little effect on MB and R_s, suggesting little change in microbial physiology. Temperature and N fertilization showed minimal interactive effects likely due to little differences in soil N availability between NN and HN samples, which is partly attributable to switchgrass biomass N accumulation (equivalent to ~53% of fertilizer N). Overall, the differential individual effects of warming and N fertilization may be driven by physiological adaptation and stimulated exoenzyme kinetics, respectively. The study shed insights on distinct microbial acquisition of different substrates under global temperature increase and N enrichment.     

Estimates of soil respiration and net primary production of three forests at different succession stages in South China

Soil respiration (heterotropic and autotropic respiration, R_g) and aboveground litter fall carbon were measured at three forests at different succession (early, middle and advanced) stages in Dinghushan Biosphere Reserve, Southern China. It was found that the soil respiration increases exponentially with soil temperature at 5 cm depth (T_s) according to the relation R_g=a exp(bT_s), and the more advanced forest community during succession has a higher value of a because of higher litter carbon input than the forests at early or middle succession stages. It was also found that the monthly soil respiration is linearly correlated with the aboveground litter carbon input of the previous month. Using measurements of aboveground litter and soil respiration, the net primary productions (NPPs) of three forests were estimated using nonlinear inversion. They are 475, 678 and 1148 g C m^?2 yr^?1 for the Masson pine forest (MPF), coniferous and broad‐leaf mixed forest (MF) and subtropical monsoon evergreen broad‐leaf forest (MEBF), respectively, in year 2003/2004, of which 54%, 37% and 62% are belowground NPP for those three respective forests if no change in live plant biomass is assumed. After taking account of the decrease in live plant biomass, we estimated the NPP of the subtropical MEBF is 970 g C m^?2 yr^?1 in year 2003/2004. Total amount of carbon allocated below ground for plant roots is 388 g C m^?2 yr^?1 for the MPF, 504 g C m^?2 yr^?1 for the coniferous and broad‐leaf MF and 1254 g C m^?2 yr^?1 for the subtropical MEBF in 2003/2004. Our results support the hypothesis that the amount of carbon allocation belowground increases during forest succession.     

Seasonal changes in the contribution of root respiration to total soil respiration in a cool-temperate deciduous forest

A trenching method was used to determine the contribution of root respiration to soil respiration. Soil respiration rates in a trenched plot (R _trench) and in a control plot (R _control) were measured from May 2000 to September 2001 by using an open-flow gas exchange system with an infrared gas analyser. The decomposition rate of dead roots (R _D) was estimated by using a root-bag method to correct the soil respiration measured from the trenched plots for the additional decaying root biomass. The soil respiration rates in the control plot increased from May (240–320 mg CO₂ m^–2 h^–1) to August (840–1150 mg CO₂ m^–2 h^–1) and then decreased during autumn (200–650 mg CO₂ m^–2 h^–1). The soil respiration rates in the trenched plot showed a similar pattern of seasonal change, but the rates were lower than in the control plot except during the 2 months following the trenching. Root respiration rate (R _r) and heterotrophic respiration rate (R _h) were estimated from R _control, R _trench, and R _D. We estimated that the contribution of R _r to total soil respiration in the growing season ranged from 27 to 71%. There was a significant relationship between R _h and soil temperature, whereas R _r had no significant correlation with soil temperature. The results suggest that the factors controlling the seasonal change of respiration differ between the two components of soil respiration, R _r and R _h.     

Dry deposition of ammonia gas drives species change faster than wet deposition of ammonium ions: evidence from a long‐term field manipulation

Although the effects of atmospheric nitrogen deposition on species composition are relatively well known, the roles of the different forms of nitrogen, in particular gaseous ammonia (NH₃), have not been tested in the field. Since 2002, we have manipulated the form of N deposition to an ombrotrophic bog, Whim, on deep peat in southern Scotland, with low ambient N (wet + dry = 8 kg N ha^?1 yr^?1) and S (4 kg S ha^?1 yr^?1) deposition. A gradient of ammonia (NH₃, dry N), from 70 kg N ha^?1 yr^?1 down to background, 3–4 kg N ha^?1 yr^?1 was generated by free air release. Wet ammonium (NH₄⁺, wet N) was provided to replicate plots in a fine rainwater spray (NH₄Cl at +8, +24, +56 kg N ha^?1 yr^?1). Automated treatments are coupled to meteorological conditions, in a globally unique, field experiment. Ammonia concentrations were converted to NH₃‐N deposition (kg N ha^?1) using a site/vegetation specific parameterization. Within 3 years, exposure to relatively modest deposition of NH₃, 20–56 kg NH₃‐N ha^?1 yr^?1 led to dramatic reductions in species cover, with almost total loss of Calluna vulgaris, Sphagnum capillifolium and Cladonia portentosa. These effects appear to result from direct foliar uptake and interaction with abiotic and biotic stresses, rather than via effects on the soil. Additional wet N by contrast, significantly increased Calluna cover after 5 years at the 56 kg N dose, but reduced cover of Sphagnum and Cladonia. Cover reductions caused by wet N were significantly different from and much smaller than those caused by equivalent dry N doses. The effects of gaseous NH₃ described here, highlight the potential for ammonia to destroy acid heathland and peat bog ecosystems. Separating the effects of gaseous ammonia and wet ammonium deposition, for a peat bog, has significant implications for regulatory bodies and conservation agencies.     

High retention of 15N‐labeled nitrogen deposition in a nitrogen saturated old‐growth tropical forest

The effects of increased reactive nitrogen (N) deposition in forests depend largely on its fate in the ecosystems. However, our knowledge on the fates of deposited N in tropical forest ecosystems and its retention mechanisms is limited. Here, we report the results from the first whole ecosystem ¹⁵N labeling experiment performed in a N‐rich old‐growth tropical forest in southern China. We added ¹⁵N tracer monthly as ¹⁵NH₄¹⁵NO₃ for 1 year to control plots and to N‐fertilized plots (N‐plots, receiving additions of 50 kg N ha^?1 yr^?1 for 10 years). Tracer recoveries in major ecosystem compartments were quantified 4 months after the last addition. Tracer recoveries in soil solution were monitored monthly to quantify leaching losses. Total tracer recovery in plant and soil (N retention) in the control plots was 72% and similar to those observed in temperate forests. The retention decreased to 52% in the N‐plots. Soil was the dominant sink, retaining 37% and 28% of the labeled N input in the control and N‐plots, respectively. Leaching below 20 cm was 50 kg N ha^?1 yr^?1 in the control plots and was close to the N input (51 kg N ha^?1 yr^?1), indicating N saturation of the top soil. Nitrogen addition increased N leaching to 73 kg N ha^?1 yr^?1. However, of these only 7 and 23 kg N ha^?1 yr^?1 in the control and N‐plots, respectively, originated from the labeled N input. Our findings indicate that deposited N, like in temperate forests, is largely incorporated into plant and soil pools in the short term, although the forest is N‐saturated, but high cycling rates may later release the N for leaching and/or gaseous loss. Thus, N cycling rates rather than short‐term N retention represent the main difference between temperate forests and the studied tropical forest.     

Rapid carbon turnover beneath shrub and tree vegetation is associated with low soil carbon stocks at a subarctic treeline

Climate warming at high northern latitudes has caused substantial increases in plant productivity of tundra vegetation and an expansion of the range of deciduous shrub species. However significant the increase in carbon (C) contained within above‐ground shrub biomass, it is modest in comparison with the amount of C stored in the soil in tundra ecosystems. Here, we use a ‘space‐for‐time’ approach to test the hypothesis that a shift from lower‐productivity tundra heath to higher‐productivity deciduous shrub vegetation in the sub‐Arctic may lead to a loss of soil C that out‐weighs the increase in above‐ground shrub biomass. We further hypothesize that a shift from ericoid to ectomycorrhizal systems coincident with this vegetation change provides a mechanism for the loss of soil C. We sampled soil C stocks, soil surface CO₂ flux rates and fungal growth rates along replicated natural transitions from birch forest (Betula pubescens), through deciduous shrub tundra (Betula nana) to tundra heaths (Empetrum nigrum) near Abisko, Swedish Lapland. We demonstrate that organic horizon soil organic C (SOC_org) is significantly lower at shrub (2.98 ± 0.48 kg m^?2) and forest (2.04 ± 0.25 kg m^?2) plots than at heath plots (7.03 ± 0.79 kg m^?2). Shrub vegetation had the highest respiration rates, suggesting that despite higher rates of C assimilation, C turnover was also very high and less C is sequestered in the ecosystem. Growth rates of fungal hyphae increased across the transition from heath to shrub, suggesting that the action of ectomycorrhizal symbionts in the scavenging of organically bound nutrients is an important pathway by which soil C is made available to microbial degradation. The expansion of deciduous shrubs onto potentially vulnerable arctic soils with large stores of C could therefore represent a significant positive feedback to the climate system.     

Rhizospheric and heterotrophic components of soil respiration in six Chinese temperate forests

Partitioning soil respiration (R_S) into heterotrophic (R_H) and rhizospheric (R_R) components is an important step for understanding and modeling carbon cycling in forest ecosystems, but few studies on R_R and R_H exist in Chinese temperate forests. In this study, we used a trenching plot approach to partition R_S in six temperate forests in northeastern China. Our specific objectives were to (1) examine seasonal patterns of soil surface CO₂ fluxes from trenched (R_T) and untrenched plots (R_UT) of these forests; (2) quantify annual fluxes of R_S components and their relative contributions in the forest ecosystems; and (3) examine effects of plot trenching on measurements of R_S and related environmental factors. The R_T maximized in early growing season, but the difference between R_UT and R_T peaked in later summer. The annual fluxes of R_H and R_R varied with forest types. The estimated values of R_H for the Korean pine (Pinus koraiensis Sieb. et Zucc.), Dahurian larch (Larix gmelinii Rupr.), aspen‐birch (Populous davidiana Dode and Betula platyphylla Suk.), hardwood (Fraxinus mandshurica Rupr., Juglans mandshurica Maxim. and Phellodendron amurense Rupr.), Mongolian oak (Quercus mongolica Fisch.) and mixed deciduous (no dominant tree species) forests averaged 89, 196, 187, 245, 261 and 301 g C m⁻² yr⁻¹, respectively; those of R_R averaged 424, 209, 628, 538, 524 and 483 g C m⁻² yr⁻¹, correspondingly; calculated contribution of R_R to R_S (RC) varied from 52% in the larch forest to 83% in the pine forest. The annual flux of R_R was strongly correlated to biomass of roots <0.5 cm in diameter, while that of R_H was weakly correlated to soil organic carbon concentration at A horizon. We concluded that vegetation type and associated carbon metabolisms of temperate forests should be considered in assessing and modeling R_S components. The significant impacts of changed soil physical environments and substrate availability by plot trenching should be appropriately tackled in analyzing and interpreting measurements of R_S components.     

Annual variation in soil respiration and its components in a coppice oak forest in Central Italy

In order to investigate the annual variation of soil respiration and its components in relation to seasonal changes in soil temperature and soil moisture in a Mediterranean mixed oak forest ecosystem, we set up a series of experimental treatments in May 1999 where litter (no litter), roots (no roots, by trenching) or both were excluded from plots of 4 m². Subsequently, we measured soil respiration, soil temperature and soil moisture in each plot over a year after the forest was coppiced. The treatments did not significantly affect soil temperature or soil moisture measured over 0–10 cm depth. Soil respiration varied markedly during the year with high rates in spring and autumn and low rates in summer, coinciding with summer drought, and in winter, with the lowest temperatures. Very high respiration rates, however, were observed during the summer immediately after rainfall events. The mean annual rate of soil respiration was 2.9 µ mol m^?2 s^?1, ranging from 1.35 to 7.03 µmol m^?2 s^?1. Soil respiration was highly correlated with temperature during winter and during spring and autumn whenever volumetric soil water content was above 20%. Below this threshold value, there was no correlation between soil respiration and soil temperature, but soil moisture was a good predictor of soil respiration. A simple empirical model that predicted soil respiration during the year, using both soil temperature and soil moisture accounted for more than 91% of the observed annual variation in soil respiration. All the components of soil respiration followed a similar seasonal trend and were affected by summer drought. The Q₁₀ value for soil respiration was 2.32, which is in agreement with other studies in forest ecosystems. However, we found a Q₁₀ value for root respiration of 2.20, which is lower than recent values reported for forest sites. The fact that the seasonal variation in root growth with temperature in Mediterranean ecosystems differs from that in temperate regions may explain this difference. In temperate regions, increases in size of root populations during the growing season, coinciding with high temperatures, may yield higher apparent Q₁₀ values than in Mediterranean regions where root growth is suppressed by summer drought. The decomposition of organic matter and belowground litter were the major components of soil respiration, accounting for almost 55% of the total soil respiration flux. This proportion is higher than has been reported for mature boreal and temperate forest and is probably the result of a short‐term C loss following recent logging at the site. The relationship proposed for soil respiration with soil temperature and soil moisture is useful for understanding and predicting potential changes in Mediterranean forest ecosystems in response to forest management and climate change.     

重庆缙云山两种林分土壤呼吸对模拟氮沉降的季节响应差异性

氮沉降对土壤呼吸的影响仍然存在着争论,需要进一步研究。选择重庆缙云山的马尾松林和柑橘林开展了氮添加实验,分别设置3个氮添加水平(低氮T_5:20 g N m~(-2)a~(-1),中氮T_(10):40 g N m~(-2)a~(-1)和高氮T_(15):60 g N m~(-2)a~(-1))和对照(T_0:0 g N m~(-2)a~(-1))共4个水平的处理,各林分每个处理各9次重复,每个处理量分4次,在每个季度开始各施1次。采用ACE(Automated Soil CO_2 Exchange Station,UK)自动土壤呼吸监测系统测定两林分土壤表层(0—10 cm)的呼吸、温度和湿度,分别在当年的7月、9月、11月、第2年的1月、2月、3月、5月、6月各连续测定4d,每天(8:00—18:00)4次,以揭示两种林分土壤呼吸对模拟氮沉降的季节动态响应及其差异性。结果表明:(1)柑橘林与马尾松林林下土壤表层呼吸表现出一致的季节变化动态趋势:夏季春季秋季冬季,但柑橘林土壤呼吸显著高于马尾松林(P0.05)。(2)总体上氮沉降抑制了2种林分土壤表层呼吸,而N沉降量大抑制程度越高。只在冬季土壤湿度低的马尾松林下氮沉降促进了土壤呼吸。(3)土壤温度与土壤呼吸有极显著的正相关指数关系(P0.01),而土壤水分与土壤呼吸有显著的二次模型拟合关系,但均受到氮沉降量处理的影响。综合分析表明,在亚热带山区2类森林下的典型案例结果支持氮沉降抑制土壤呼吸的认识。     

Foliar maintenance respiration of subalpine and boreal trees and shrubs in relation to nitrogen content

A nitrogen-based model of maintenance respiration (R_m) would link R_m with nitrogen-based photosynthesis models and enable simpler estimation of dark respiration flux from forest canopies. To test whether an N-based model of R_m would apply generally to foliage of boreal and subalpine woody plants, I measured R_m (CO₂ efflux at night from fully expanded foliage) for foliage of seven species of trees and shrubs in the northern boreal forest (near Thompson, Manitoba, Canada) and seven species in the subalpine montane forest (near Fraser, Colorado, USA). At 10°C, average R_m for boreal foliage ranged from 0.94 to 6.8μmol kg^?1 s^?1 (0.18–0.58 μmol m^?2 s^?1) and for subalpine foliage it ranged from 0.99 to 7.6 μmol kg^?1 s^?1 (0.28–0.64μmol m^?2 s^?1). CO₂ efflux at 10°C for the samples was only weakly correlated with sample weight (r = 0.11) and leaf area (r = 0.58). However, CO₂ efflux per unit foliage weight was highly correlated with foliage N concentration [r = 0.83, CO₂ flux at 10°C (mol kg^?1 s^?1) = 2.62 × foliage N (mol kg^?1)J, and slopes were statistically similar for the boreal and subalpine sites (P=0.28). CO₂ efflux per unit of foliar N was 1.8 times that reported for a variety of crop and wildland species growing in warmer climates.     

The contribution of bryophytes to the carbon exchange for a temperate rainforest

Bryophytes blanket the floor of temperate rainforests in New Zealand and may influence a number of important ecosystem processes, including carbon cycling. Their contribution to forest floor carbon exchange was determined in a mature, undisturbed podocarp‐broadleaved forest in New Zealand, dominated by 100–400‐year‐old rimu (Dacrydium cupressimum) trees. Eight species of mosses and 13 species of liverworts contributed to the 62% cover of the diverse forest floor community. The bryophyte community developed a relatively thin (depth <30 mm), but dense, canopy that experienced elevated CO₂ partial pressures (median 46.6 Pa immediately below the bryophyte canopy) relative to the surrounding air (median 37.6 Pa at 100 mm above the canopy). Light‐saturated rates of net CO₂ exchange from 14 microcosms collected from the forest floor were highly variable; the maximum rate of net uptake (bryophyte photosynthesis – whole‐plant respiration) per unit ground area at saturating irradiance was 1.9 μmol m^?2 s^?1 and in one microcosm, the net rate of CO₂ exchange was negative (respiration). CO₂ exchange for all microcosms was strongly dependent on water content. The average water content in the microcosms ranged from 1375% when fully saturated to 250% when air‐dried. Reduction in water content across this range resulted in an average decrease of 85% in net CO₂ uptake per unit ground area. The results from the microcosms were used in a model to estimate annual carbon exchange for the forest floor. This model incorporated hourly variability in average irradiance reaching the forest floor, water content of the bryophyte layer, and air and soil temperature. The annual net carbon uptake by forest floor bryophytes was 103 g m^?2, compared to annual carbon efflux from the forest floor (bryophyte and soil respiration) of ?1010 g m^?2. To put this in perspective of the magnitude of the components of CO₂ exchange for the forest floor, the bryophyte layer reclaimed an amount of CO₂ equivalent to only about 10% of forest floor respiration (bryophyte plus soil) or ～11% of soil respiration. The contribution of forest floor bryophytes to productivity in this temperate rainforest was much smaller than in boreal forests, possibly because of differences in species composition and environmental limitations to photosynthesis. Because of their close dependence on water table depth, the contribution of the bryophyte community to ecosystem CO₂ exchange may be highly responsive to rapid changes in climate.     

Direct and indirect effects of elevated atmospheric CO2 on net ecosystem production in a Chesapeake Bay tidal wetland

The rapid increase in atmospheric CO₂ concentrations (C_a) has resulted in extensive research efforts to understand its impact on terrestrial ecosystems, especially carbon balance. Despite these efforts, there are relatively few data comparing net ecosystem exchange of CO₂ between the atmosphere and the biosphere (NEE), under both ambient and elevated C_a. Here we report data on annual sums of CO₂ (NEE_net) for 19 years on a Chesapeake Bay tidal wetland for Scirpus olneyi (C₃ photosynthetic pathway)‐ and Spartina patens (C₄ photosynthetic pathway)‐dominated high marsh communities exposed to ambient and elevated C_a (ambient + 340 ppm). Our objectives were to (i) quantify effects of elevated C_a on seasonally integrated CO₂ assimilation (NEE_net = NEE_day + NEE_night, kg C m^?2 y^?1) for the two communities; and (ii) quantify effects of altered canopy N content on ecosystem photosynthesis and respiration. Across all years, NEE_net averaged 1.9 kg m^?2 y^?1 in ambient C_a and 2.5 kg m^?2 y^?1 in elevated C_a, for the C₃‐dominated community. Similarly, elevated C_a significantly (P < 0.01) increased carbon uptake in the C₄‐dominated community, as NEE_net averaged 1.5 kg m^?2 y^?1 in ambient C_a and 1.7 kg m^?2 y^?1 in elevated C_a. This resulted in an average CO₂ stimulation of 32% and 13% of seasonally integrated NEE_net for the C₃‐ and C₄‐dominated communities, respectively. Increased NEE_day was correlated with increased efficiencies of light and nitrogen use for net carbon assimilation under elevated C_a, while decreased NEE_night was associated with lower canopy nitrogen content. These results suggest that rising C_a may increase carbon assimilation in both C₃‐ and C₄‐dominated wetland communities. The challenge remains to identify the fate of the assimilated carbon.     

Impacts of fire on sources of soil CO2 efflux in a dry Amazon rain forest

Fire at the dry southern margin of the Amazon rainforest could have major consequences for regional soil carbon (C) storage and ecosystem carbon dioxide (CO₂) emissions, but relatively little information exists about impacts of fire on soil C cycling within this sensitive ecotone. We measured CO₂ effluxes from different soil components (ground surface litter, roots, mycorrhizae, soil organic matter) at a large‐scale burn experiment designed to simulate a severe but realistic potential future scenario for the region (Fire plot) in Mato Grosso, Brazil, over 1 year, and compared these measurements to replicated data from a nearby, unmodified Control plot. After four burns over 5 years, soil CO₂ efflux (R_s) was ~5.5 t C ha^?1 year^?1 lower on the Fire plot compared to the Control. Most of the Fire plot R_s reduction was specifically due to lower ground surface litter and root respiration. Mycorrhizal respiration on both plots was around ~20% of R_s. Soil surface temperature appeared to be more important than moisture as a driver of seasonal patterns in R_s at the site. Regular fire events decreased the seasonality of R_s at the study site, due to apparent differences in environmental sensitivities among biotic and abiotic soil components. These findings may contribute toward improved predictions of the amount and temporal pattern of C emissions across the large areas of tropical forest facing increasing fire disturbances associated with climate change and human activities.     

Regional scale gradients of climate and nitrogen deposition drive variation in ectomycorrhizal fungal communities associated with native Scots pine

Ectomycorrhizal fungi commonly associate with the roots of forest trees where they enhance nutrient and water uptake, promote seedling establishment and have an important role in forest nutrient cycling. Predicting the response of ectomycorrhizal fungi to environmental change is an important step to maintaining forest productivity in the future. These predictions are currently limited by an incomplete understanding of the relative significance of environmental drivers in determining the community composition of ectomycorrhizal (ECM) fungi at large spatial scales. To identify patterns of community composition in ECM fungi along regional scale gradients of climate and nitrogen deposition in Scotland, fungal communities were analysed from 15 seminatural Scots pine (Pinus sylvestris L.) forests. Fungal taxa were identified by sequencing of the ITS rDNA region using fungal‐specific primers. Nonmetric multidimensional scaling was used to assess the significance of 16 climatic, pollutant and edaphic variables on community composition. Vector fitting showed that there was a strong influence of rainfall and soil moisture on community composition at the species level, and a smaller impact of temperature on the abundance of ectomycorrhizal exploration types. Nitrogen deposition was also found to be important in determining community composition, but only when the forest experiencing the highest deposition (9.8 kg N ha^?1 yr^?1) was included in the analysis. This finding supports previously published critical load estimates for ectomycorrhizal fungi of 5–10 kg N ha^?1 yr^?1. This work demonstrates that both climate and nitrogen deposition can drive gradients of fungal community composition at a regional scale.     

Changes in microbial activities and biomasses over a forest floor gradient in C-to-N ratio

Background and aims

Under chronically elevated N deposition, N retention mainly occur at high soil C-to-N ratio. This may be mediated through soil microbes, such as ectomycorrhizal (EM) fungi, saprotrophic fungi and bacteria, and the aim of this study was to evaluate the relationship between soil microbes and forest floor C-to-N ratios.

Methods

Soil samples from 33 Norway spruce (Picea abies (L.) H. Karst) forests in Denmark and southern Sweden in a forest floor C-to-N ratio gradient (ranging from 14 to 35) were analysed regarding the content of phospholipid fatty acids (PLFAs) to estimate their soil microbial community composition and the relative biomasses of different microbial groups. The relation of EM biomass to total fungal biomass was estimated as the loss of the fungal PLFA 18:2ω6,9 during incubation of soils and the production of EM mycelia was estimated using fungal in-growth mesh bags. The soil microbial variables were correlated to forest floor C-to-N ratio, NO ₃ ^- leaching, soil pH and stand age.

Results

Fungal proportions of microbial biomass, EM to total fungi and EM mycelial production were all positively related to C-to-N ratio, while NO ₃ ^- leaching was negatively related to C-to-N ratio.

Conclusions

Both EM and saprotrophic fungi change with forest floor C-to-N ratios and appear to play a central role in N retention in forest soil. A better understanding of the mechanisms behind this process may be revealed if the role of recalcitrant fungal metabolites for N retention (and soil C sequestration) can be identified. Research along this line deserves further studies.     

Soil CO2 efflux in contrasting boreal deciduous and coniferous stands and its contribution to the ecosystem carbon balance

Similar nonsteady‐state automated chamber systems were used to measure and partition soil CO₂ efflux in contrasting deciduous (trembling aspen) and coniferous (black spruce and jack pine) stands located within 100 km of each other near the southern edge of the Boreal forest in Canada. The stands were exposed to similar climate forcing in 2003, including marked seasonal variations in soil water availability, which provided a unique opportunity to investigate the influence of climate and stand characteristics on soil CO₂ efflux and to quantify its contribution to the net ecosystem CO₂ exchange (NEE) as measured with the eddy‐covariance technique. Partitioning of soil CO₂ efflux between soil respiration (including forest‐floor vegetation) and forest‐floor photosynthesis showed that short‐ and long‐term temporal variations of soil CO₂ efflux were related to the influence of (1) soil temperature and water content on soil respiration and (2) below‐canopy light availability, plant water status and forest‐floor plant species composition on forest‐floor photosynthesis. Overall, the three stands were weak to moderate sinks for CO₂ in 2003 (NEE of ?103, ?80 and ?28 g C m^?2 yr^?1 for aspen, black spruce and jack pine, respectively). Forest‐floor respiration accounted for 86%, 73% and 75% of annual ecosystem respiration, in the three respective stands, while forest‐floor photosynthesis contributed to 11% and 14% of annual gross ecosystem photosynthesis in the black spruce and jack pine stands, respectively. The results emphasize the need to perform concomitant measurements of NEE and soil CO₂ efflux at longer time scales in different ecosystems in order to better understand the impacts of future interannual climate variability and vegetation dynamics associated with climate change on each component of the carbon balance.     

Heterotrophic Soil Respiration in Relation to Environmental Factors and Microbial Biomass in Two Wet Tropical Forests

We examined the correlation between fungal and bacterial biomass, abiotic factors such as soil moisture, carbon in the light soil fraction and soil nitrogen to a depth of 0–25 cm and heterotrophic soil respiration using a trenching technique – in a secondary forest (Myrcia splendens, Miconia prasina and Casearia arborea) and a pine (Pinus caribeae) plantation in the Luquillo Experimental Forest in Puerto Rico. Soil respiration was significantly reduced where roots were excluded for 7 years in both the secondary forest and the pine plantation. Microbial biomass was also significantly reduced in the root exclusion plots. In root exclusion treatment, total fungal biomass was on average 31 and 65% lower than the control plots in the pine plantation and the secondary forest, respectively, but the total bacterial biomass was 24 and 8.3% lower than the control plots in the pine plantation and the secondary forest, respectively. Heterotrophic soil respiration was positively correlated with fungal biomass (R²=0.63, R²=0.39), bacterial biomass (R²=0.16, R²=0.45), soil moisture (R²=0.41, R²=0.56), carbon in light fraction (R²=0.45, R²=0.39) and total nitrogen (R²=0.69, R²=0.67) in the pine plantation and the secondary forest, respectively. The regression analysis suggested that fungal biomass might have a greater influence on heterotrophic soil respiration in the pine plantation, while the bacterial biomass might have a greater influence in the secondary forest. Heterotrophic soil respiration was more sensitive to total N than to carbon in the light fraction, and soil moisture was a major factor influencing heterotrophic soil respiration in these forests where temperature is high and relatively invariable.