Carbon fluxes in coniferous and deciduous forest soils

Aims

Our aims were to identify responsible factors for the site-to-site variability in soil CO₂ efflux and to assess the sources of soil CO₂ of different forest types on a regional scale.

Methods

Soil CO₂ effluxes were measured over 1–4 years in four coniferous and three deciduous forests of Bavaria, Germany, and related to climate, soil properties and forest productivity. Total belowground carbon allocation (TBCA) was assessed using soil CO₂ effluxes and aboveground litterfall. Additionally, CO₂ production of organic layers was examined over 10 months under constant conditions in an incubation experiment.

Results

Annual soil CO₂ effluxes were not different among the forest sites, but predicted effluxes at a given temperature of 10°C revealed some significant differences and correlated with the phosphorous stock of the organic layers. The incubation study indicated 50% faster decomposition of organic layers from deciduous than from coniferous forests. TBCA related to soil CO₂ efflux was smaller in the deciduous than in the coniferous forests. The ratio of TBCA to soil CO₂ efflux was positively correlated with the C stock of organic layers.

Conclusions

Our results suggest that marked differences in site characteristics have little impact on soil CO₂ effluxes at the regional scale, but the contribution of soil CO₂ sources varies among the forest types.     

Gross nitrogen transformations in soils from uncut and cut boreal upland and peatland coniferous forest stands

Gross and net nitrogen (N) ammonification and nitrification were measured in soils from an uncut and recently cut upland and peatland conifer stand in northwestern Ontario, Canada. Rates of gross total inorganic N immobilization were similar to gross mineralization, resulting in low net mineralization rates in soils from all four upland and peatland conifer stands. Gross ammonification rates were variable but similar in soils from uncut and cut peatland hollows (18–19mgNkg^–1day^–1) and upland forest floor soils (14–19mgNkg^–1day^–1). Gross ammonium ( ) immobilization rates were also variable but similar to ammonification rates. Median gross nitrification rates were within 0–2mgNkg^–1day^–1 in soils from all four upland and peatland cut and uncut stands, although rates were consistently higher for the soils from the cut stands. Large variability in gross nitrification rates were observed in peatland soils, however the highest gross nitrification rates were measured in saturated peatland soils. Net rates remained low in the soils from all four stands due to high nitrate ( ) immobilization and very fast turnover (<0.2 day). Our results suggest that potential losses may be negated by high immobilization in uncut and cut boreal forest stands. This study reveals the potential for the interaction of N production and consumption processes in regulating N retention in upland and peatland conifer forests, and the resilience of the boreal forest to disturbance.     

Simulation and scaling of temporal variation in gross primary production for coniferous and deciduous temperate forests

Observations of ecosystem net carbon dioxide exchange obtained with eddy covariance techniques over a 4‐year period at spruce, beech and pine forest sites were used to derive time series data for gross primary production (GPP) and ecosystem respiration (R_eco). A detailed canopy gas exchange model (PROXEL_NEE) was inverted at half‐hour time step to estimate seasonal changes in carboxylation capacity and light utilization efficiency of the forest canopies. The parameter estimates were then used further to develop a time‐dependent modifier of physiological activity in the daily time step gas exchange model of Chen et al. (1999) , previously used for regional simulations in BOREAS. The daily model was run under a variety of assumptions and the results emphasize the need in future analyses: (1) to focus on time‐dependent seasonal changes in canopy physiology as well as in leaf area index, (2) to compare time courses of physiological change in different habitats in terms of recognizable cardinal points in the seasonal course, and (3) to develop methods for utilizing information on seasonal changes in physiology in regional and continental carbon budget simulations. The results suggest that the daily model with appropriate seasonal adjustments for physiological process regulation should be an efficient tool for use in conjunction with remote sensing for regional evaluation of global change scenarios.     

Nitrification in coniferous forest soils

Net nitrification rates tend to be low or negligible in the forest floor of many coniferous forests of North-East Scotland. The most likely process controls are substrate availability, pH, allelopathy, water potential, nutrient status and temperature. These are discussed in relation to field and laboratory studies of net and potential rates of nitrification.Fungi make up by far the largest part of the nitrifier community in the coniferous forest floor. Very little is known about the distribution and activity of autotrophs in these systems, although it is certain that in vitro evidence suggesting autotrophs cannot nitrify at pH levels characteristic of coniferous forest soils is unrealistic.Because of the metabolic diversity of nitrifying fungi, a variety of organic and inorganic nitrification pathways may exist in coniferous forests. The possible involvement of free radicles in fungal nitrification in coniferous forest soils is also suggested.A complete understanding of nitrification in coniferous forest soils can only result from field characterisation of N flux such as through the use of ¹⁵N. This must be combined with ecophysiological characterisation of the organisms involved in order that the complexity of nitrification in coniferous forest soils can be resolved.     

Nitrogen oxide gas emissions from temperate forest soils receiving long-term nitrogen inputs

From spring 2000 through fall 2001, we measured nitric oxide (NO) and nitrous oxide (N₂O) fluxes in two temperate forest sites in Massachusetts, USA that have been treated since 1988 with different levels of nitrogen (N) to simulate elevated rates of atmospheric N deposition. Plots within a pine stand that were treated with either 50 or 150 kg N ha^?1 yr^?1 above background displayed consistently elevated NO fluxes (100–200 µg N m^?2 h^?1) compared to control plots, while only the higher N treatment plot within a mixed hardwood stand displayed similarly elevated NO fluxes. Annual NO emissions estimated from monthly sampling accounted for 3.0–3.7% of N inputs to the high‐N plots and 8.3% of inputs to the Pine low‐N plot. Nitrous oxide fluxes in the N‐treated plots were generally < 10% of NO fluxes. Net nitrification rates (NRs) and NO production rates measured in the laboratory displayed patterns that were consistent with field NO fluxes. Total N oxide gas flux was positively correlated with contemporaneous measurements of NR and concentration. Acetylene inhibited both nitrification and NO production, indicating that autotrophic nitrification was responsible for the elevated NO production. Soil pH was negatively correlated with N deposition rate. Low levels (3–11 µg N kg^?1) of nitrite () were detected in mineral soils from both sites. Kinetic models describing NO production as a function of the protonated form of (nitrous acid [HNO₂]) adequately described the mineral soil data. The results indicate that atmospheric deposition may generate losses of gaseous NO from forest soils by promoting nitrification, and that the response may vary significantly between forest types under similar climatic regimes. The lowering of pH resulting from nitrification and/or directly from deposition may also play a role by promoting reactions involving HNO₂.     

Long-term succession in a Danish temperate deciduous forest

Forest successional trajectories covering the last 2000 yr from a mixed deciduous forest in Denmark show a gradual shift in dominance from Tilia cordata to Fagus sylvatica and a recent increase in total forest basal area since direct management ceased in 1948. The successions are reconstructed by combining a fifty-year record of direct tree observations with local pollen diagrams from Draved Forest, Denmark. Five of the seven successions record a heathland phase of Viking Age dating from 830 AD. The anthropogenic influence is considerable throughout the period of study even though Draved contains some of the most pristine forest stands in Denmark. Anthropogenic influence including felling masks the underlying natural dynamics, with the least disturbed sites showing the smallest compositional change. Some effects of former management, such as loss of Tilia cordata dominance, are irreversible. Artificial disturbance, particularly drainage, has accelerated and amplified the shift towards Fagus dominance that would have occurred on a smaller scale and at a slower rate in the absence of human intervention.     

Trade-offs between seedling growth and survival in deciduous broadleaved trees in a temperate forest

BACKGROUND AND AIMS: In spatially heterogeneous environments, a trade-off between seedling survival and relative growth rate may promote the coexistence of plant species. In temperate forests, however, little support for this hypothesis has been found under field conditions, as compared with shade-house experiments. Performance trade-offs were examined over a large resource gradient in a temperate hardwood forest. METHODS: The relationship between seedling survival and seedling relative growth rate in mass (RGR(M)) or height (RGR(H)) was examined at three levels of canopy cover (forest understorey, FU; small gap, SG; and large gap, LG) and at two microsites within each level of canopy cover (presence or absence of leaf litter) for five deciduous broad-leaved tree species with different seed sizes. KEY RESULTS: Within each species, both RGR(M) and RGR(H) usually increased with increasing light levels (in the order FU < SG < LG), whereas little difference was observed based on the presence or absence of litter. Seedling survival in FU was negatively correlated with both RGR(M) and RGR(H) in both LG and SG. The trade-off between high-light growth and low-light survival was more evident in the relationship with LG as compared with SG. An intraspecific trade-off between survival and RGR was observed along environmental gradients in Acer mono, whereas seedlings of Betula platyphylla var. japonica survived and grew better in LG. CONCLUSIONS: The results presented here strongly support the idea of light gradient partitioning (i.e. species coexistence) in spatially heterogeneous light environments in temperate forests, and that further species diversity would be promoted by increased spatial heterogeneity. The intraspecific trade-off between survival and RGR in Acer suggests that it has broad habitat requirements, whereas Betula has narrow habitat requirements and specializes in high-light environments.     

Relationships between carbon turnover and bioavailable energy fluxes in two temperate forest soils

Recent research has shown that the broad empirical relationships used in many ecosystem models to predict carbon turnover and stabilization in soils can fail to capture differences across vegetation types or climates. Theoretically, because energy flow is fundamental to the function of decomposer organisms and ecosystems, energetics could provide complimentary fundamental constraints on soil C dynamics. Often, however, C is considered as a surrogate for energy in studies of detrital decay and C turnover in soil. Bomb calorimetry has long been used to measure stored energy in organic matter, but in detritus not all of the energy is bioavailable. Here I outline an approach to quantify the flux of bioavailable energy dissipated by resident heterotrophic communities in soil organic horizons in situ. I used the principle of energy balance together with a biogeochemical process model parameterized through calorimetric analysis of field samples. I also tested relationships between C and energy across samples of forest detritus (foliar and fine root litter, well‐decayed Oea material, and woody debris), across decay stages, and between a deciduous and coniferous forest at the Harvard Forest, MA, USA. As a first approximation, energy and C concentrations were closely related (within ca. 10%), as were ratios of heterotrophic energy dissipation to C mineralization across types of detritus (within 16%). Differences in energy content and energy : C ratios were measurable in forest detritus (particularly woody vs nonwoody), but did not vary reliably enough between forest types or through detrital stages to indicate that soil C models could be improved by including energetics. Model results indicated that there are strong similarities in energy flows and storage in the O horizons of the contrasting forest types studied at this location. Future research could focus on broader patterns across climates or biomes.     

Interactions between two co-dominant, invasive plants in the understory of a temperate deciduous forest

Negative interactions between non-indigenous and native species has been an important research topic of invasion biology. However, interactions between two or more invasive species may be as important in understanding biological invasions, but they have rarely been studied. In this paper, we describe three field experiments that investigated interactions between two non-indigenous plant species invasive in the eastern United States, Lonicera japonica (a perennial vine) and Microstegium vimineum (an annual grass). A press removal experiment conducted within a deciduous forest understory community indicated that M. vimineum was a superior competitor to L. japonica. We tested the hypothesis that the competitive success of M. vimineum was because it overgrew, and reduced light available to, L. japonica, by conducting a separate light gradient experiment within the same community. Shade cloth that simulated the M. vimineum canopy reduced the performance of L. japonica. In a third complementary experiment, we added experimental support hosts to test the hypothesis that the competitive ability of L. japonica is limited by support hosts, onto which L. japonica climbs to access light. We found that the abundance of climbing branches increased with the number of support hosts. Results of this experiment indicate that these two invasive species compete asymmetrically for resources, particularly light.     

Population dynamics of coniferous and broad-leaved trees in a Japanese temperate mixed forest

The growth and survival of coniferous and broad-leaved trees were followed over a 5-yr period in a temperate old-growth mixed forest in Japan, and dynamic features of the forest were studied in relation to the life history of the dominants, the coniferous Abies homolepis and the broad-leaved Fagus crenata. During this period, the gap formation rate was 31m² ha^?1yr^?1, the mortality of trees > 2m high was 1.7%/yr, and the rate of loss in basal area 1.4%/yr. These values were much higher than the recruitment, 0.3%/yr, and the total growth of surviving and new trees, 0.6%/yr, owing to the inhibition of regeneration by understorey dwarf bamboo (Sasa borealis). A transition matrix model based on DBH size classes predicts that the basal area of the forest will decrease by 14% in 50 yr, but that the DBH distribution of trees > 10 cm diameter will change little. Equilibrium DBH distributions assuming recruitment being equal to mortality, were quite different between broad-leaved and coniferous trees, reflecting different survivorship curves of the two dominants. The composition and structure of the forest may change depending on the pattern and frequency of disturbances, or episodic events, notably the synchronous death of Sasa borealis.     

Soil seed bank composition and diversity in a managed temperate deciduous forest

Little is known about the influence of forest management on the interaction between seed bank and aboveground vegetation. We surveyed seed banks and vegetation in 10 forest stands under similar abiotic conditions but submitted either to a coppice-with-standards treatment (n=5) or to a selective-cutting system (n=5). We analyzed species composition and diversity, community ecological profile, and distribution of taxa among different life forms, strategy, morphology and functional type categories. A total of 2085 seedlings (8296 seedsm^–2) germinated-corresponding to 28 species, among which Juncus effusus was the most abundant. Fifty-seven percent of the species were also recorded in the aboveground vegetation, the dominant species being Rubus fruticosus agg., but only 28% of the aboveground species were present in the seed bank. Our results suggest that (1) vernal geophytes and shade-tolerant perennials, which group most true forest species, are not incorporated in the seed bank, (2) parent plants of most seeds were present either in the stand in an earlier dynamic stage or apart from the stand and long-distance dispersed, (3) as expected, early-successional species are well represented in the seed bank, (4) forestry vehicles seem to be a major means of dispersion for stress-tolerant species normally found in forest lanes and wheel tracks. We conclude that seed banks contain species that have a potentially negative impact on the true forest flora and, thus, forest management should minimize soil disturbance and retain remnants of old-coppice woods to conserve disturbance-sensitive true forest species.     

Soil moisture effects on gross nitrification differ between adjacent grassland and forested soils in central Alberta, Canada

Background and aims

Changes in soil moisture availability seasonally and as a result of climatic variability would influence soil nitrogen (N) cycling in different land use systems. This study aimed to understand mechanisms of soil moisture availability on gross N transformation rates.

Methods

A laboratory incubation experiment was conducted to evaluate the effects of soil moisture content (65 vs. 100% water holding capacity, WHC) on gross N transformation rates using the ¹⁵N tracing technique (calculated by the numerical model FLUAZ) in adjacent grassland and forest soils in central Alberta, Canada.

Results

Gross N mineralization and gross NH ₄ ⁺ immobilization rates were not influenced by soil moisture content for both soils. Gross nitrification rates were greater at 100 than at 65% WHC only in the forest soil. Denitrification rates during the 9 days of incubation were 2.47 and 4.91 mg N kg^-1 soil d^-1 in the grassland and forest soils, respectively, at 100% WHC, but were not different from zero at 65% WHC. In the forest soil, both the ratio of gross nitrification to gross NH ₄ ⁺ immobilization rates (N/IA) and cumulative N₂O emission were lower in the 65 than in the 100% WHC treatment, while in the grassland soil, the N/IA ratio was similar between the two soil moisture content treatments but cumulative N₂O emission was lower at 65% WHC.

Conclusions

The effect of soil moisture content on gross nitrification rates differ between forest and grassland soils and decreasing soil moisture content from 100 to 65% WHC reduced N₂O emissions in both soils.     

Abiotic factors regulating nitrogen transformations in the organic layer of acid forest soils: Moisture and pH

The relationships between nitrogen transformations and moisture and pH in coniferous forest litter were determined using laboratory incubation experiments. A linear relation between gravimetric moisture content and nitrification was found within the whole studied range of moisture conditions (10–290% ODW). Net nitrogen mineralization increased linearly with moisture content up to 140% ODW. At higher moisture contents, net mineralization was found to be independent of moisture. Relative nitrification was found to be a linear function of moisture content. The dependence of the CO₂ production rates on moisture in the coniferous litter decreased from low to high moisture availability. Due to a nearly linear relationship between gravimetric moisture content and log-(water potential) within the investigated moisture range, the same type of relationships were found with this latter parameter as well. The relationship between nitrogen transformations and pH was studied by means of the addition of different amounts of HCl and NaOH during short incubation experiments (1 week). Nitrification was found to be a negative linear function of the H-ion concentration within the range of 0.04 (pH 4.40) and 0.36 (pH 3.45) mmol H-ion L^?1. At a higher H-ion concentration and thus at a lower pH than 3.45, no nitrate was produced any more. No relationship between net mineralization and pH was found.     

Root carbon inputs to the rhizosphere stimulate extracellular enzyme activity and increase nitrogen availability in temperate forest soils

The exudation of carbon (C) by tree roots stimulates microbial activity and the production of extracellular enzymes in the rhizosphere. Here, we investigated whether the strength of rhizosphere processes differed between temperate forest trees that vary in soil organic matter (SOM) chemistry and associate with either ectomycorrhizal (ECM) or arbuscular mycorrhizal (AM) fungi. We measured rates of root exudation, microbial and extracellular enzyme activity, and nitrogen (N) availability in samples of rhizosphere and bulk soil influenced by four temperate forest tree species (i.e., to estimate a rhizosphere effect). Although not significantly different between species, root exudation ranged from 0.36 to 1.10 g C m^?2 day^?1, representing a small but important transfer of C to rhizosphere microbes. The magnitude of the rhizosphere effects could not be easily characterized by mycorrhizal associations or SOM chemistry. Ash had the lowest rhizosphere effects and beech had the highest rhizosphere effects, representing one AM and one ECM species, respectively. Hemlock and sugar maple had equivalent rhizosphere effects on enzyme activity. However, the form of N produced in the rhizosphere varied with mycorrhizal association. Enhanced enzyme activity primarily increased amino acid availability in ECM rhizospheres and increased inorganic N availability in AM rhizospheres. These results show that the exudation of C by roots can enhance extracellular enzyme activity and soil-N cycling. This work suggests that global changes that alter belowground C allocation have the potential to impact the form and amount of N to support primary production in ECM and AM stands.     

Modelling carbon dynamics in coniferous forest soils in a temperature gradient

Climate change and changes in land use will alter the stores of carbon and turnover of soil organic matter. We have used a theory for carbon cycles in terrestrial ecosystems to analyse changes in soil organic matter turnover in coniferous forests. The central concepts of the theory are a continuously changing substrate quality, a constant decomposer efficiency and a climatically controlled decomposer growth rate. Measurements on litter production and soil carbon stores from field experiments have been used to successfully validate the model predictions. Measured litter production increased with increasing temperature but the response was not identical for forests of different vegetation types which reflect variations in productivity. The temperature response of needle-litter production and decomposition rate were strongest in the most productive forests and weakest for the low productive forests. Initial decay rates of soil C store from steady state showed the same trend in temperature response as decay of a single litter cohort did, but the absolute values are 16% of the decay rates of a single litter cohort. Predicted soil C ranged from 5 to 9 kg C m^–2. There exists a remarkable variation in forest soil C store response to temperature; the magnitude and even the sign depends on productivity as defined by vegetation type. The assumption that, in general, decomposition rates increase more than NPP with temperature, and consequently, soil C stores should decrease in response to a climate warming, seems therefore too simplistic.     

Vegetation structure and diversity of an ancient temperate deciduous forest in SW Denmark

Bolderslev Skov (113 ha) is the largest contiguous ancient forest remnant in Denmark. The forest has been preserved since 1998 as a strict non-intervention forest reserve. We studied vegetation structure, floristic gradients, and diversity of the forest in 50 plots of 100 m² placed according to a restricted random sampling design. Dominant tree species were Fagus sylvatica, Fraxinus excelsior, Tilia cordata and Quercus robur, which in most parts of the forest form a mixed canopy. Most stands appeared to be of moderate age, 55-80 years old, and large old trees were rare. pH in association with light and thickness of the litter layer were the most important factors in explaining floristic gradients in the forest. Soil moisture (dry to mesic) was not strongly correlated with DCA axes for neither tree nor field layer, but had a significant effect on the distribution of a number of herb species. Forest structure was not important in explaining species distributions. Field layer species richness showed a positive relationship with the pH-gradient. At the scale of 1m² plots we also found a highly significant negative relationship between field layer species richness and the plot-wise Ellenberg indicator value for nitrogen availability. Structure of the tree layer had little effect on field layer species richness. The mixed composition, long continuity, and presence of a high proportion of the regional species pool assigns a high conservation value to Bolderslev Skov and makes it an important site for future studies of the dynamic properties, niche preferences, and inter-specific competition of temperate deciduous forest species. This revised version was published online in June 2006 with corrections to the Cover Date.     

Factors influencing sapling composition in canopy gaps of a temperate deciduous forest

To detect the factors that affect sapling species composition in gaps, we investigated 55 gaps in an old-growth temperate deciduous forest in Ogawa Forest Reserve, central Japan. Gap size, gap age, gap maker species, topographic location, adult tree composition around gaps, and saplings of tree species growing in the gaps were censused. For gaps 5 m², mean gap size was 70 m² and the maximum was 330 m². Estimated ages of gaps had a tendency to be concentrated in particular periods relating to strong wind records in the past. The sapling composition in gaps was highly and significantly correlated to that under closed canopy, indicating the importance of advance regeneration in this forest. However, some species showed significant occurrence biases in gaps or under closed canopy, suggesting differences in shade tolerance. The result of MANOVA showed that gap size and topography were important factors in determining the sapling composition in gaps. Species of gap makers affected the sapling composition indirectly by influencing gap size. The existence of parent trees around gaps had effects on sapling densities of several species. Gap age did not have clear influences on sapling composition. Variations in gap size and topography were considered as important factors that contribute to maintenance of species diversity in this forest.