Clean rain promotes fine root growth and soil respiration in a Norway spruce forest

Soil acidification and N saturation are considered to affect the decomposition of soil organic matter as well as growth and mortality of fine roots in many forest soils. Here we report from a field experiment where ‘clean rain’ has been applied to the soil for about 10 years under a roofed plot of a 71‐year‐old Norway spruce plantation at Solling, Central Germany. Reduced amounts of protons (?78%), sulphate (?53%), ammonium (?86%), and nitrate (?49%) were sprayed on the soil surface of the clean rain plot between 1992 and 2001. In an adjacent roofed control plot, throughfall was collected and immediately re‐sprinkled below the roof construction without any chemical manipulation. One year before the clean rain treatment started, live and dead fine root masses (≤2 mm) were determined from undisturbed soil cores down to 40 cm mineral soil depth. Total live fine root mass was significantly lower in the clean rain plot than in the control plot. After the first sampling, the soil holes were refilled with quartz sand and repeatedly sampled in June 1992, June 1996, and October 2001. There were no differences in live and dead fine root masses between the plots in 1992 and 1996. In 2001, both live and dead fine root masses of the clean rain plot were about twice as high as in the control plot, indicating that fine root growth recovered in the mineral soil following 10 years of clean rain treatment. Moreover, the clean rain treatment significantly reduced the total N concentrations of live fine roots and 1‐year‐old needles. Our results suggest that the reduced N input promoted fine root growth to compensate N deficiency. Reduced Al concentration in soil solution may have contributed to the recovery of fine root growth, however, the toxicity of Al species is largely unknown. Mean annual soil respiration rate was 24% higher in the period from 2000 to 2001, indicating that the clean rain treatment increased respiration of roots and heterotrophic microorganisms within the rhizosphere. Laboratory incubation of samples from the organic horizon and the top mineral soil revealed no differences between the plots in the decay rate of soil organic matter. Our results suggest that strong reductions in atmospheric N deposition from about 30 to 10 kg N ha^?1 yr^?1 and decreasing acid stress can have beneficial effects on growth of fine roots in the mineral soil within a decade. We conclude that biological recovery under reduced atmospheric loads can affect the nutrient and carbon budget of spruce soils in the long run.     

Separation of root and heterotrophic respiration within soil respiration by trenching, root biomass regression, and root excising methods in a cool-temperate deciduous forest in Japan

Trenching (Tr), root biomass regression (RR), and root excising (RE) methods were used to estimate the contribution of root (RR) and heterotrophic (HR) respiration to soil respiration (SR) in a cool-temperate deciduous forest in central Japan. The contribution ratios of RR to SR were 23 % (?16 to 46 %), 11 % (?19 to 61 %), and 115 % (20 to 393 %), as estimated by the Tr, RR, and RE methods, respectively. The contribution ratio showed clear seasonal variation with high values in summer for the Tr method, while they were undetectable for the RR and RE methods because of some methodological problems. These results suggest the Tr method is the best of the three methods used to estimate the contribution ratio of RR and HR to SR in the forest. Annual SR, RR, and HR rates, estimated by the Tr method, were 479, 369, 110 gC m^?2 year^?1, respectively. The seasonal variation of SR was mainly influenced by HR (77 %) throughout the year, while the influence of RR on SR was strongest in summer (46 %). This effect occurred because RR (Q ₁₀ = 7.5) is more sensitive to temperature than HR (Q ₁₀ = 3.2). Also, the contribution of fine RR to total RR was higher than that of coarse RR because of high respiratory activity (Q ₁₀ and R ₁₀) as well as the large biomass of fine roots. These results suggest that each component of SR responds differently to the same environmental factors and their relative influence on SR changes across the seasons.     

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

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

Effects of soil frost on soil respiration and its radiocarbon signature in a Norway spruce forest soil

Apart from a general increase of mean annual air temperature, climate models predict a regional increase of the frequency and intensity of soil frost with possibly strong effects on C cycling of soils. In this study, we induced mild soil frost (up to −5 °C in a depth of 5 cm below surface) in a Norway spruce forest soil by removing the natural snow cover in the winter of 2005/2006. Soil frost lasted from January to April 2006 and was detected down to 15 cm depth. Soil frost effectively reduced soil respiration in the snow removal plots in comparison to undisturbed control plots. On an annual basis 6.2 t C ha⁻¹ a⁻¹ were emitted in the control plots compared with 5.1 t C ha⁻¹ a⁻¹ in the snow removal plots. Only 14% of this difference was attributed to reduced soil respiration during the soil frost period itself, whereas 63% of this difference originated from differences during the summer of 2006. Radiocarbon (Δ¹⁴C) signature of CO₂ revealed a considerable reduction of heterotrophic respiration on the snow removal plots, only partly compensated for by a slight increase of rhizosphere respiration. Similar CO₂ concentrations in the uppermost mineral horizons of both treatments indicate that differences between the treatments originated from the organic horizons. Extremely low water contents between June and October of 2006 may have inhibited the recovery of the heterotrophic organisms from the frost period, thereby enhancing the differences between the control and snow removal plots. We conclude that soil frost triggered a change in the composition of the microbial community, leading to an increased sensitivity of heterotrophic respiration to summer drought. A CO₂ pulse during thawing, such as described for arable soils several times throughout the literature, with the potential to partly compensate for reduced soil respiration during soil frost, appears to be lacking for this soil. Our results from this experiment indicate that soil frost reduces C emission from forest soils, whereas mild winters may enhance C losses from forest soils.     

Autotrophic and heterotrophic respiration in needle fir and Quercus-dominated stands in a cool-temperate forest, central Korea

To investigate annual variation in soil respiration (R _S) and its components [autotrophic (R _A) and heterotrophic (R _H)] in relation to seasonal changes in soil temperature (ST) and soil water content (SWC) in an Abies holophylla stand (stand A) and a Quercus-dominated stand (stand Q), we set up trenched plots and measured R _S, ST and SWC for 2 years. The mean annual rate of R _S was 436 mg CO₂ m⁻² h⁻¹, ranging from 76 to 1,170 mg CO₂ m⁻² h⁻¹, in stand A and 376 mg CO₂ m⁻² h⁻¹, ranging from 82 to 1,133 mg CO₂ m⁻² h⁻¹, in stand Q. A significant relationship between R _S and its components and ST was observed over the 2 years in both stands, whereas a significant correlation between R _A and SWC was detected only in stand Q. On average over the 2 years, R _A accounted for approximately 34% (range 17–67%) and 31% (15–82%) of the variation in R _S in stands A and Q, respectively. Our results suggested that vegetation type did not significantly affect the annual mean contributions of R _A or R _H, but did affect the pattern of seasonal change in the contribution of R _A to R _S.     

Fertilization of boreal forest reduces both autotrophic and heterotrophic soil respiration

The boreal forest is expected to experience the greatest warming of all forest biomes, raising concerns that some of the large quantities of soil carbon in these systems may be added to the atmosphere as CO₂. However, nitrogen deposition or fertilization has the potential to increase boreal forest production and retard the decomposition of soil organic matter, hence increasing both tree stand and soil C storage. The major contributors to soil‐surface CO₂ effluxes are autotrophic and heterotrophic respiration. To evaluate the effect of nutrient additions on the relative contributions from autotrophic and heterotrophic respiration, a large‐scale girdling experiment was performed in a long‐term nutrient optimization experiment in a 40‐year‐old stand of Norway spruce in northern Sweden. Trees on three nonfertilized plots and three fertilized plots were girdled in early summer 2002, and three nonfertilized and three fertilized plots were used as control plots. Each plot was 0.1 ha and contained around 230 trees. Soil‐surface CO₂ fluxes, soil moisture, and soil temperature were monitored in both girdled and nongirdled plots. In late July, the time of the seasonal maximum in soil‐surface CO₂ efflux, the total soil‐CO₂ efflux in nongirdled plots was 40% lower in the fertilized than in the nonfertilized plots, while the efflux in girdled fertilized and nonfertilized plots was 50% and 60% lower, respectively, than in the corresponding nongirdled controls. We attribute these reductions to losses of the autotrophic component of the total soil‐surface CO₂ efflux. The estimates of autotrophic respiration are conservative as root starch reserves were depleted more rapidly in roots of girdled than in nongirdled trees. Thus, heterotrophic activity was overestimated. Calculated on a unit area basis, both the heterotrophic and autotrophic soil respiration was significantly lower in fertilized plots, which is especially noteworthy given that aboveground production was around three times higher in fertilized than in nonfertilized plots.     

Spatial and temporal effects of soil temperature and moisture and the relation to fine root density on root and soil respiration in a mature apple orchard

We identified the role of various soil parameters and root density as drivers of soil respiration (R_s) in an apple orchard, measured during different periods of the year and at a range of distances from trees, in plots with a different history of nutrient supply. R_s was measured in April, May, August and December and studied in relation to soil temperature and moisture, total soil C and N, as well as to fine root density and medium-, and large-sized root density and root N concentration. The study also aimed to partition R_s by applying the root regression technique. R_s ranged from 0.06 in December to 1.49 g CO₂ m^?2 h^?1 in August. Average soil temperature alone explained up to 71% of the annual variability of R_s, while soil water content was negatively correlated to R_s. Fertilization, soil C and N concentration and root N had negligible effects on R_s. Fine root density, but not medium- and large-sized root density, contributed to explaining part of the yearly variability of R_s and proved to be a good predictor in December, when the statistical significance of the regression made it possible to estimate the autotrophic component of R_s as being about 35% of total soil respiration.     

Partitioning soil respiration: quantifying the artifacts of the trenching method

Sediment porewater nutrients often occur at concentrations that are orders of magnitude higher than nutrients in overlying waters, and accordingly may subsidise growth of benthic macroalgal mats in estuarine ecosystems. The relative contribution of porewater nutrients is expected to be particularly important for macroalgae entrained in intertidal mudflat sediments, where access to water column nutrients is tidally constrained. In this study, filamentous Gracilaria chilensis thalli were simultaneously exposed to sediment and overlying water nutrient sources, labelled using ¹⁵N tracers (¹⁵NH₄⁺ or ¹⁵NO₃^?) during a 5-day experiment. Dissolved inorganic N (DIN) uptake from porewater and overlying water accounted for 33 and 52%, respectively, of the N estimated as necessary to support the growth of G. chilensis, despite the two-fold lower DIN concentration of the overlying water and its periodic availability (8 h day^?1). Of the total N assimilated by the plants,?~?15% could not be accounted for, supporting the acquisition of other N forms in order to meet demand. We also found that regardless of background NH₄⁺:NO₃^? ratios (i.e. 1:3 in overlying water and 12:1 in porewater), plants accumulated ¹⁵NH₄⁺ significantly more readily than ¹⁵NO₃^?, indicating a preference for NH₄⁺. This ability to utilise multiple sources and species of N relatively rapidly may partly explain the competitive success of entrained macroalgae relative to non-entrained species and historically abundant seagrass beds in these environments. These results underscore the significance of both internal nutrient loading and external inputs as important in sustaining opportunistic macroalgal blooms in shallow estuaries.     

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.     

Partitioning sources of soil respiration in boreal black spruce forest using radiocarbon

Separating ecosystem and soil respiration into autotrophic and heterotrophic component sources is necessary for understanding how the net ecosystem exchange of carbon (C) will respond to current and future changes in climate and vegetation. Here, we use an isotope mass balance method based on radiocarbon to partition respiration sources in three mature black spruce forest stands in Alaska. Radiocarbon (Δ¹⁴C) signatures of respired C reflect the age of substrate C and can be used to differentiate source pools within ecosystems. Recently‐fixed C that fuels plant or microbial metabolism has Δ¹⁴C values close to that of current atmospheric CO₂, while C respired from litter and soil organic matter decomposition will reflect the longer residence time of C in plant and soil C pools. Contrary to our expectations, the Δ¹⁴C of C respired by recently excised black spruce roots averaged 14‰ greater than expected for recently fixed photosynthetic products, indicating that some portion of the C fueling root metabolism was derived from C storage pools with turnover times of at least several years. The Δ¹⁴C values of C respired by heterotrophs in laboratory incubations of soil organic matter averaged 60‰ higher than the contemporary atmosphere Δ¹⁴CO₂, indicating that the major contributors to decomposition are derived from a combination of sources consistent with a mean residence time of up to a decade. Comparing autotrophic and heterotrophic Δ¹⁴C end members with measurements of the Δ¹⁴C of total soil respiration, we calculated that 47–63% of soil CO₂ emissions were derived from heterotrophic respiration across all three sites. Our limited temporal sampling also observed no significant differences in the partitioning of soil respiration in the early season compared with the late season. Future work is needed to address the reasons for high Δ¹⁴C values in root respiration and issues of whether this method fully captures the contribution of rhizosphere respiration.     

The measurement of woody root decomposition using two methodologies in a Sitka spruce forest ecosystem

Background and aims

The decomposition of roots is an important process in the loss of carbon (C) and the mineralization of nitrogen (N) in forest ecosystems. The early stage decomposition rate of Sitka spruce (Picea sitchensis (Bong.) Carr.) roots was determined using trenched plots and decomposition bags.

Methods

Stumps of known age were trenched and quadrants (50?cm by 50?cm) excavated from randomly selected stumps every 6?months over 4?years, while the mass loss from buried roots in decomposition bags, divided among four diameter categories (ranging from fine roots <2?mm to large roots >50?mm), was monitored for 27?months. The C and N concentrations of excavated samples at different time points were analysed.

Results

The change in total root necromass per quadrant showed a higher decomposition rate-constant (k) of 0.24?±?0.068?year^?1 than the k-value of roots in decomposition bags (0.07?±?0.005?year^?1). The C concentration (47.24?±?0.609?%) did not significantly change with decomposition. There was a significant increase in the C:N ratio of roots in all diameter categories (fine: 48.92?%, small: 38.53?%, medium 11.71?%, large: 76.25?%) after 4?years of decomposition, driven by N loss. Root diameter accounted for 78?% of the variation in the N concentration of roots as decomposition progressed.

Conclusion

Though the trenched plot approach offered an alternative to the more common decomposition bag method for estimating root decomposition, high spatial variation and sampling difficulties may lead to an overestimation of the mass loss from trenched roots, thus, the decomposition bag method gives a more reliable decomposition rate-constant.     

Management intensification effects on autotrophic and heterotrophic soil respiration in subtropical grasslands

Sustainable management of grassland ecosystems for improved productivity can enhance their potential to sequester atmospheric CO₂ in the soil. However, land-use management influences the quantity and quality of carbon (C) inputs which may, in turn, affect microbial activity and soil C decomposition rates. Understanding the potential changes in magnitude of soil C loss through respiration is critical for a comprehensive assessment of land-use conversion and grassland management impacts on terrestrial C dynamics. Thus, this study was designed to assess the effect of land-use management intensification on soil respiration in subtropical grasslands. Experimental sites consisted of a gradient of management intensities ranging from native rangeland (lowest), silvopasture (intermediate), to sown pasture (highest). Increasing management intensity from native rangeland to sown pasture elevated soil respiration. There was a significant effect of ‘season vs. management’ interaction on total soil respiration (R_S), with greater increases in R_S from summer to winter in sown pasture (∼200%) compared to native rangeland and silvopasture (∼91%). The temperature sensitivity of R_S and heterotrophic soil respiration (R_H) increased with management intensification, with a highest Q₁₀ of 1.55 and 2.29, in sown pasture, compared to Q₁₀ values of 1.09 and 1.48 in native rangelands. These results suggested that potential increases in soil C stock with intensification may be susceptible to faster turnover under warming climate scenarios. Improved resilience (and longer residence) of additionally sequestered soil C after intensification may be crucial for long-term ecological resilience, especially with changing climatic conditions. These findings are relevant for sustainable grassland management, especially within subtropical ecoregions, and add to the understanding of changes that may occur in rates of soil C losses as native grasslands are converted to more productive grassland ecosystems.     

Variation of soil respiration and its components in hemiboreal Norway spruce stands of different ages

Aims

Understanding the linkage of soil respiration (Rs) with forest development is essential for long-term C cycle models. We estimated the variation and temperature sensitivity (Q₁₀ value) of Rs and its hetero-, (Rh) and autotrophic (Ra) components in relation to abiotic and biotic factors in Norway spruce stands of different ages, and the effect of trenching on microbial and soil characteristics.

Methods

Trenching method was used to partition Rs into Rh and Ra. Ingrowth core method was used to estimate fine root production. Soil microbial biomass was measured using manometric respirometers.

Results

Rs varied in differently aged stands demonstrating non-linear response to development stage. The variation of Rs was explained by changes in biotic factors rather than by changes in soil microclimate. Rh was more sensitive to Ts than Rs or Ra. After 4 years of trenching soil pH, N, SOM and dehydrogenase activity were significantly changed in trenched plots compared to control plots.

Conclusions

Different Q₁₀ values of Rh and Ra in stands of different ages indicate the importance of Rs partitioning. Trenching should be used during a limited number of years because of the possible changes in chemical characteristics of soil and in the activity of soil microbial community.

     

Genetic host-tree effects on the ectomycorrhizal community and root characteristics of Norway spruce

A greenhouse experiment was used to study the effects of host genotype on short root formation and ectomycorrhizal (ECM) fungal community structure in Norway spruce (Picea abies (L.) Karst.). Rooted cuttings representing 55 clones were inoculated with a mix of vegetative hyphae of five ECM fungal species (Laccaria sp., Amphinema byssoides, Piloderma sp., Cadophora finlandia, Paxillus involutus). After one growing season, the ECM fungal community structure was determined by amplifying the fungal internal transcribed spacer (ITS) of ribosomal DNA directly from ECM root tips. Restriction profiles of obtained amplicons were then compared to those of the inoculated strains. Spruce clones differed in their ECM fungal community composition; we found a statistically significant clone-specific effect on ECM fungal diversity and dominating fungal species. Nevertheless, the broad sense heritabilities of the levels of Laccaria sp., Piloderma sp. and A. byssoides colonisations as well as the ECM fungal community structure were low (H ²?=?0.04?0.11), owing to the high within-clone variation. As nitrogen concentration of needles correlated negatively with ECM fungal richness, our results imply that in the experimental conditions nutrient acquisition of young trees may benefit from colonisation with only one or two ECM fungal species. The heritability of short root density was moderate (H ²?=?0.41) and highest among all the measured shoot and root growth characteristics of Norway spruce cuttings. We suggest that the genetic component determining root growth and short root formation is significant for the performance of young trees in natural environments as these traits drive the formation of the below-ground symbiotic interactions.     

The effects of clipping and soil moisture on leaf and root morphology and root respiration in two temperate and two tropical grasses

Numerous studies have explored the effect of environmental conditions on a number of plant physiological and structural traits, such as photosynthetic rate, shoot versus root biomass allocation, and leaf and root morphology. In contrast, there have been a few investigations of how those conditions may influence root respiration, even though this flux can represent a major component of carbon (C) pathway in plants. In this study, we examined the response of mass-specific root respiration (μmol CO₂ g⁻¹ s⁻¹), shoot and root biomass, and leaf photosynthesis to clipping and variable soil moisture in two C₃ (Festuca idahoensis Elmer., Poa pratensis L.) and two C₄ (Andropogon greenwayi Napper, and Sporobolus kentrophyllus K. Schum.) grass species. The C₃ and C₄ grasses were collected in Yellowstone National Park, USA and the Serengeti ecosystem, Africa, respectively, where they evolved under temporally variable soil moisture conditions and were exposed to frequent, often intense grazing. We also measured the influence of clipping and soil moisture on specific leaf area (SLA), a trait associated with moisture conservation, and specific root length (SRL), a trait associated with efficiency per unit mass of soil resource uptake. Clipping did not influence any plant trait, with the exception that it reduced the root to shoot ratio (R:S) and increased SRL in P. pratensis. In contrast to the null effect of clipping on specific root respiration, reduced soil moisture lowered specific root respiration in all four species. In addition, species differed in how leaf and root structural traits responded to lower available soil moisture. P. pratensis and A. greenwayi increased SLA, by 23% and 33%, respectively, and did not alter SRL. Conversely, S. kentrophyllus increased SRL by 42% and did not alter SLA. F. idahoensis responded to lower available soil moisture by increasing both SLA and SRL by 38% and 33%, respectively. These responses were species-specific strategies that did not coincide with photosynthetic pathway (C₃/C₄) or growth form. Thus, mass-specific root respiration responded uniformly among these four grass species to clipping (no effect) and increased soil moisture stress (decline), whereas the responses of other traits (i.e., R:S ratio, SLA, SRL) to the treatments, especially moisture availability, were species-specific. Consequently, the effects of either clipping or variation in soil moisture on the C budget of these four different grasses species were driven primarily by the plasticity of R:S ratios and the structural leaf and root traits of individual species, rather than variation in the response of mass-specific root respiration.     

Effects of soil phosphorus availability,temperature and moisture on soil respiration in Eucalyptus pauciflora forest

Rates of soil respiration (CO₂ efflux) were measured for a year in a mature Eucalyptus pauciflora forest in unfertilized and phosphorus-fertilized plots. Soil CO₂ efflux showed a distinct seasonal trend, and average daily rates ranged from 124 to 574 mg CO₂ m^–2 hr^–1. Temperature and moisture are the main variables that cause variation in soil CO₂ efflux; hence their effects were investigated over a year so as to then differentiate the treatment effect of phosphorus (P) nutrition.Soil temperature had the greatest effect on CO₂ efflux and exhibited a highly significant logarithmic relationship (r² = 0.81). Periods of low soil and litter moisture occurred during summer when temperatures were greater than 10 °C, and this resulted in depression of soil CO₂ efflux. During winter, when temperatures were less than 10 °C, soil and litter moisture were consistently high and thus their variation had little effect on soil CO₂ efflux. A multiple regression model including soil temperature, and soil and litter moisture accounted for 97% of the variance in rates of CO₂ efflux, and thus can be used to predict soil CO₂ efflux at this site with high accuracy. Total annual efflux of carbon from soil was estimated to be 7.11 t C ha^–1 yr^–1. The model was used to predict changes in this annual flux if temperature and moisture conditions were altered. The extent to which coefficients of the model differ among sites and forest types requires testing.Increased soil P availability resulted in a large increase in stem growth of trees but a reduction in the rate of soil CO₂ efflux by approximately 8%. This reduction is suggested to be due to lower root activity resulting from reduced allocation of assimilate belowground. Root activity changed when P was added to microsites within plots, and via the whole tree root system at the plot level. These relationships of belowground carbon fluxes with temperature, moisture and nutrient availability provide essential information for understanding and predicting potential changes in forest ecosystems in response to land use management or climate change.     

Influence of Norway spruce seedlings on the nutrient availability in mineral soil and forest floor material

A greenhouse experiment was performed to evaluate the effect of Norway spruce (Picea abies (L.) Karst.) seedlings on net nutrient availability in five different growing media containing F- or H-layer and mineral soil originating from a haplic podzol in northern Sweden. The initial total amounts of eight nutrient elements (N, K, P, Ca, Mg, Mn, Fe, Zn) and exchangeable amounts of the same elements were analyzed in pots with or without spruce seedlings. In the planted pots seedling nutrient uptake was also estimated. After 26 weeks, higher net nutrient availability with seedlings was found in 25 out of the 40 (62%) growing media and nutrient element combinations. A positive seedling effect on net nutrient availability might be explained by rhizodeposition stimulating the soil microorganism activity and accelerating the weathering of minerals or by seedling roots promoting the nutrient providing processes through changes in soil chemical and physical properties. Nitrogen availability was primarily affected by what part of the forest floor the growing medium contained although the positive response to seedling presence was apparent. The positive net availability response of P, Ca, Mg, Mn, Fe and Zn to seedling presence was on the other hand relatively strong. In the case of P, K, and Zn the growing medium composition (if the F- and H-layer was pure or mixed with mineral soil) was also an important factor for the estimated net availability. Pure F-and H-layer provided greater P- and K-availability while the availability of Zn increased when mineral soil was added. The influence of growing plants ought to be considered when soil samples are used for assessing the nutrient availability.     

Slow response of soil organic matter to the reduction in atmospheric nitrogen deposition in a Norway spruce forest

Global nitrogen (N) deposition rates in terrestrial environments have quadrupled since preindustrial times, causing structural and functional changes of ecosystems. Different emission reduction policies were therefore devised. The aim of our study was to investigate if, and over what timescale, processes of soil organic matter (OM) transformation respond to a decline in atmospheric N deposition. A N‐saturated spruce forest (current N deposition: 34 kg ha^?1 yr^?1; critical N load: 14 kg ha^?1 yr^?1), where N deposition has been reduced to 11.5 kg ha^?1 yr^?1 since 1991, was studied. Besides organic C and organic and inorganic N, noncellulosic carbohydrates, amino sugars and amino acids were determined. A decline in organic N in litter indicated initial effects at plant level. However, there were no changes in biomarkers upon the reduction in N deposition. In addition, inorganic N was not affected by reduced N deposition. The results showed that OM cycling and transformation processes have not responded so far. It was concluded that no direct N deposition effects have occurred due to the large amount of stored organic N, which seems to compensate for the reduction in deposited N. Obviously, the time span of atmospheric N reduction (about 14.5 years) is too short compared with the mean turnover time of litter to cause indirect effects on the composition of organic C and N compounds. It is assumed that ecological processes, such as microbial decomposition or recycling of organic N and C, react slowly, but may start within the next decade with the incorporation of the new litter.     

Interactive effects of soil warming and fertilization on root production, mortality, and longevity in a Norway spruce stand in Northern Sweden

The effects of soil warming and nitrogen availability on root production, longevity and mortality were studied using minirhizotrons in irrigation (C), fertilized (F), heated (H), and heated‐fertilized (HF) plots in a Norway spruce stand in northern Sweden from October 1996 to October 1997. Irrigation was included in all treatment plots. Heating cables were used to maintain the soil temperature in heated plots at 5°C above that in unheated plots during the growing season. A Kaplan–Meier approach was used to estimate the longevity of fine roots and Cox proportional hazards regression to analyze the effects of the H, F, and HF treatments on the risk of root mortality. The proportion of annual root length production contributed by winter–spring production amounted to 52% and 49% in heated plots and heated‐fertilized plots, respectively. The annual root length production in C plots was significantly higher than in other treatments, while the HF treatment gave significantly greater production compared with the F treatment. The risk of mortality (hazard ratio) relative to C plots was higher in H plots (358%) and F plots (191%). The interaction between heating and fertilizing was strongly significant. The increase in the risk of root mortality in combined fertilization and heating (103%) was lower than that in the H or F plots. The results show that nitrogen addition combined with warmer temperatures decreases the risk of root mortality, and fine root production is a function of the length of the growing season. In the future, fertilization combined with the warmer temperatures expected to follow predicted climatic change may increase root production in boreal forests at low fertility sites.     

Contribution of newly grown extramatrical ectomycorrhizal mycelium and fine roots to soil respiration in a young Norway spruce site

Background and aims

Partitioning of soil respiration is a challenging task when resolving the C cycling in forest ecosystems. Our aim was to partition the respiration of newly grown extramatrical ectomycorrhizal mycelium (ECM) and fine roots (and their associated microorganisms) in a young Norway spruce forest.

Methods

Ingrowth mesh bags of 16 cm diameter and 12 cm height were placed in the upper soil and left for 12–16 months in 2010 and 2011. The 2 mm mesh size allowed the ingrowth of ECM and fine roots whereas a 45 μm mesh size allowed only the ingrowth of ECM. The mesh bags were filled with either homogenized EA horizon soil, pure quartz sand (QS) or crushed granite (CG, only 2011), each with five replicates. Controls without any ingrowth were established for each substrate by solid plastic tubes (2010) and by 1 μm mesh bags (2011). Fluxes of CO₂ from the mesh bags and controls were measured biweekly during the growing season by the closed chamber method.

Results

The contribution of ECM to soil respiration was largest in the QS treatments, reaching cumulatively 1.2 and 2.2 Mg C ha^?1 6 months^?1 in 2010 and 2011, respectively. For EA and CG treatments, the cumulative respiration from ECM was larger than from controls, however the differences being not statistically significant. The respiration of newly grown fine roots in QS amounted to 1.0 Mg C ha^?1 in 2010, but could not be identified in 2011 since fluxes from 2 mm and 45 μm mesh bags were similar. The correlation of total root length in single QS mesh bags to CO₂ fluxes was poor. The contribution of fine root respiration was also not detectable in the EA and CG treatment. No correlation was found between the autumnal biomass of newly grown ECM and its cumulative respiration.

Conclusion

Our results suggest a substantial contribution of newly grown ECM to soil respiration. Respiration of ECM might be larger than respiration of fine roots.