Effects of fire severity on plant nutrient uptake reinforce alternate pathways of succession in boreal forests

Fire activity in the North American boreal region is projected to increase under a warming climate and trigger changes in vegetation composition. In black spruce forests of interior Alaska, fire severity impacts residual organic layer depth which is strongly linked to the relative dominance of deciduous versus coniferous trees in early succession. These alternate successional pathways may be reinforced by biogeochemical processes that affect the relative ability of deciduous versus coniferous trees to acquire limiting nutrients. To test this hypothesis, we examined changes in soil inorganic nitrogen (N) supply and in situ ¹⁵N root uptake by aspen (Populus tremuloides) and black spruce (Picea mariana) saplings regenerating in lightly and severely burned sites, 16 years following fire. Fire severity did not impact the composition or magnitude of N supply, and nitrate represented nearly 40 % of total N supply. Both aspen and spruce took up more N in severely burned than in lightly burned sites. Spruce exhibited only a moderately lower rate of nitrate uptake, and a higher ammonium uptake rate than aspen in severely burned sites. At the stand level, differences in species nutrient uptake were magnified, with aspen taking up nearly an order-of-magnitude more N per m² in severely burned than in lightly burned sites. We suggest that differences in nutrient sinks (biomass) established early in succession and effects of post-fire organic layer depth on nutrient uptake, are key mechanisms reinforcing the opposing stand dominance patterns that have developed in response to variations in organic layer depth.     

Contribution of winter processes to soil nitrogen flux in taiga forest ecosystems

We measured annual net nitrogen (N) mineralization, nitrification, and amino acid production in situ across a primary successional sequence in interior Alaska, USA. Net N mineralization per gram dry soil increased across the successional sequence, but with a sharp decline in the oldest stage (black spruce). Net N mineralization expressed per gram soil organic matter exhibited the opposite pattern, suggesting that soil organic matter quality decreases significantly across succession. Net N mineralization rates during the growing season from green-up (early May) through freeze-up (late September–early October) accounted for approximately 60% of the annual inorganic N flux, whereas the remaining N was released during the apparent dormant season. Nitrogen release during winter occurred primarily during October–January with only negligible N mineralization during early spring in stands of willow, alder, balsam poplar and white spruce. By contrast, black spruce stands exhibited substantial mineralization after snow melt during early spring. The high rates of N mineralization in late autumn through early winter coincide with high turnover of fine root biomass in these stands, suggesting that labile substrate production, rather than temperature, is a major controlling factor over N release in these ecosystems. We suggest that the convention of restricting measurements of soil processes to the growing season greatly underestimate annual flux rates of inorganic nitrogen in these high-latitude ecosystems.     

Effects of long-term nitrogen addition on phosphorus cycling in organic soil horizons of temperate forests

High atmospheric nitrogen (N) deposition is expected to impair phosphorus (P) nutrition of temperate forest ecosystems. We examined N and P cycling in organic soil horizons of temperate forests exposed to long-term N addition in the northeastern USA and Scandinavia. We determined N and P concentrations, enzyme activities and net N and P mineralization rates in organic soil horizons of two deciduous (Harvard Forest, Bear Brook) and two coniferous (Klosterhede, Gårdsjön) forests which had received experimental inorganic N addition between 25 and 150 kg N ha^?1 year^?1 for more than 25 years. Long-term N addition increased the activity of phosphatase (+?180%) and the activity of carbon (C)- and N-acquiring enzymes (cellobiohydrolase: +?70%, chitinase: +?25%). Soil N enrichment increased the N:P ratio of organic soil horizons by up to 150%. In coniferous organic soil horizons, net N and P mineralization were small and unaffected by N addition. In deciduous organic soil horizons, net N and P mineralization rates were significantly higher than at the coniferous sites, and N addition increased net N mineralization by up to 290%. High phosphatase activities concomitant with a 40% decline in P stocks of deciduous organic soil horizons indicate increased plant P demand. In summary, projected future global increases in atmospheric N deposition may induce P limitation in deciduous forests, impairing temperate forest growth.     

Effects of Succession on Nitrogen Export in the West-Central Cascades, Oregon

This study examined impacts of succession on N export from 20 headwater stream systems in the west central Cascades of Oregon, a region of low anthropogenic N inputs. The seasonal and successional patterns of nitrate (NO₃−N) concentrations drove differences in total dissolved N concentrations because ammonium (NH₄−N) concentrations were very low (usually < 0.005 mg L⁻¹) and mean dissolved organic nitrogen (DON) concentrations were less variable than nitrate concentrations. In contrast to studies suggesting that DON levels strongly dominate in pristine watersheds, DON accounted for 24, 52, and 51% of the overall mean TDN concentration of our young (defined as predominantly in stand initiation and stem exclusion phases), middle-aged (defined as mixes of mostly understory reinitiation and older phases) and old-growth watersheds, respectively. Although other studies of cutting in unpolluted forests have suggested a harvest effect lasting 5 years or less, our young successional watersheds that were all older than 10 years still lost significantly more N, primarily as NO₃−N, than did watersheds containing more mature forests, even though all forest floor and mineral soil C:N ratios were well above levels reported in the literature for leaching of dissolved inorganic nitrogen. The influence of alder may contribute to these patterns, although hardwood cover was quite low in all watersheds; it is possible that in forested ecosystems with very low anthropogenic N inputs, even very low alder cover in riparian zones can cause elevated N exports. Only the youngest watersheds, with the highest nitrate losses, exhibited seasonal patterns of increased summer uptake by vegetation as well as flushing at the onset of fall freshets. Older watersheds with lower N losses did not exhibit seasonal patterns for any N species. The results, taken together, suggest a role for both vegetation and hydrology in N retention and loss, and add to our understanding of N cycling by successional forest ecosystems influenced by disturbance at various spatial and temporal scales in a region of relatively low anthropogenic N input.     

The Effect of Increased N Deposition on Nitrous oxide, Methane and Carbon dioxide Fluxes from Unmanaged Forest and Grassland Communities in Michigan

Atmospheric nitrogen deposition is anticipated to increase over the next decades with possible implications for future forest-atmosphere interactions. Increased soil N₂O emissions, depressed CH₄ uptake and depressed soil respiration CO₂ loss is considered a likely response to increased N deposition. This study examined fluxes of N₂O, CH₄ and CO₂ over two growing seasons from soils in unmanaged forest and grassland communities on abandoned agricultural areas in Michigan. All sites were subject to simulated increased N-deposition in the range of 1–3 g N m⁻² annually. Nitrous oxide fluxes and soil N concentrations in coniferous and grassland sites were on the whole unaffected by the increased N-inputs. It is noteworthy though that N₂O emissions increased three-fold in the coniferous sites in the first growing season in response to the low N treatment, although the response was barely significant (p<0.06). In deciduous forests, we observed increased levels of soil mineral N during the second year of N fertilization, however N₂O fluxes did not increase. Rates of methane oxidation were similar in all sites with no affect of field N application. Likewise, we did not observe any changes in soil CO₂ efflux in response to N additions. The combination of tillage history and vegetation type was important for the trace gas fluxes, i.e. soil CO₂ efflux was greater in successional grassland sites compared with the forested sites and CH₄ uptake was reduced in post-tillage coniferous- and successional sites compared with the old-growth deciduous site. Our results indicate that short-term increased N availability influenced individual processes linked to trace gas turnover in the soil independently from the ecosystem N status. However, changes in whole system fluxes were not evident and were very likely mediated by competitive N uptake processes.     

A Canopy Shift in Interior Alaskan Boreal Forests: Consequences for Above- and Belowground Carbon and Nitrogen Pools during Post-fire Succession

Global change models predict that high-latitude boreal forests will become increasingly susceptible to fire activity as climate warms, possibly causing a positive feedback to warming through fire-driven emissions of CO₂ into the atmosphere. However, fire-climate feedbacks depend on forest regrowth and carbon (C) accumulation over the post-fire successional interval, which is influenced by nitrogen (N) availability. To improve our understanding of post-fire C and N accumulation patterns in boreal forests, we evaluated above- and belowground C and N pools within 70 stands throughout interior Alaska, a region predicted to undergo a shift in canopy dominance as fire severity increases. Stands represented gradients in age and successional trajectory, from black spruce (Picea mariana) self-replacement to species replacement by deciduous species of trembling aspen (Populus tremuloides) and Alaska paper birch (Betula neoalaskana). Stands undergoing deciduous trajectories stored proportionally more of their C and N in aboveground stemwood and had 5–7 times faster rates of aboveground net primary productivity of trees compared to stands undergoing a black spruce trajectory, which stored more of their C and N in the soil organic layer (SOL), a thick layer of mostly undecomposed mosses. Thus, as successional trajectories shift, total C and N pool sizes will remain relatively unchanged, but there will be a trade-off in pool location and a potential increase in C and N longevity due to decreased flammability and decomposition rates of deciduous stemwood. Despite often warmer, drier conditions in deciduous compared to black spruce stands, deciduous stemwood has a C:N around 10 times higher than the black spruce SOL and often remains standing for many years with reduced exposure to fungal decomposers. Thus, a fire-driven shift in successional trajectories could cause a negative feedback to climate warming because of increased pool longevity in deciduous trajectories.     

In situ gross nitrogen transformations differ between temperate deciduous and coniferous forest soils

Despite long-term enhanced nitrogen (N) inputs, forests can retain considerable amounts of N. While rates of N inputs via throughfall and N leaching are increased in coniferous stands relative to deciduous stands at comparable sites, N leaching below coniferous stands is disproportionally enhanced relative to the N input. A better understanding of factors affecting N retention is needed to assess the impact of changing N deposition on N cycling and N loss of forests. Therefore, gross N transformation pathways were quantified in undisturbed well-drained sandy soils of adjacent equal-aged deciduous (pedunculate oak (Quercus robur L.)) and coniferous (Scots pine (Pinus sylvestris L.)) planted forest stands located in a region with high N deposition (north Belgium). In situ inorganic ¹⁵N labelling of the mineral topsoil (0–10?cm) combined with numerical data analysis demonstrated that (i) all gross N transformations differed significantly (p?<?0.05) between the two forest soils, (ii) gross N mineralization in the pine soil was less than half the rate in the oak soil, (iii) meaningful N immobilization was only observed for ammonium, (iv) nitrate production via oxidation of organic N occurred three times faster in the pine soil while ammonium oxidation was similar in both soils, and (v) dissimilatory nitrate reduction to ammonium was detected in both soils but was higher in the oak soil. We conclude that the higher gross nitrification (including oxidation of organic N) in the pine soil compared to the oak soil, combined with negligible nitrate immobilization, is in line with the observed higher nitrate leaching under the pine forest.     

Rapid Cycling of Organic Nitrogen in Taiga Forest Ecosystems

ABSTRACT We examined the dynamics of organic nitrogen (N) turnover in situ across a primary successional sequence in interior Alaska, USA, in an attempt to understand the magnitude of these fluxes in cold, seasonally frozen soils. Through a combination of soil extraction procedures and measurements of ¹³C-enriched CO₂ efflux from soils amended in the field with ¹³C-labeled amino acids, we were able to trace the fate of this N form. Amino acid turnover in situ at soil temperatures of 10°C or below show that amino acids represent a highly dynamic soil N pool with turnover times of approximately 3–6 h. The rapid turnover of free amino acids is associated with high soil proteolytic activity, which in turn is tightly correlated with soil protein concentration. Moreover, these estimates of soil amino acid turnover in the field correspond well with measurements of amino acid turnover under equivalent temperatures in the laboratory. The gross flux of amino acid-N over the growing season greatly exceeded the annual vegetation N requirement, suggesting that microbial biomass represent a significant sink for this organic N. Depending on the strength of this sink, N flow via free soil amino acids can potentially account for the entire N demand of vegetation in the absence of net N mineralization. These relationships underscore the important biogeochemical role of labile DON fractions in high-latitude forest ecosystems.     

Tree species mediated soil chemical changes in a Siberian artificial afforestation experiment

Natural and human-induced changes in the composition of boreal forests will likely alter soil properties, but predicting these effects requires a better understanding of how individual forest species alter soils. We show that 30 years of experimental afforestation in Siberia caused species-specific changes in soil chemical properties, including pH, DOC, DON, Na⁺, NH₄ ⁺, total C, C/N, Mn²⁺, and SO₄ ^2-. Some of these properties –- pH, total C, C/N, DOC, DON, Na⁺ –- also differed by soil depth, but we found no strong evidence for species-dependent effects on vertical differentiation of soil properties (i.e., no species × depth interaction). A number of soil properties –- NO₃ ^–, N, Al³⁺, Ca²⁺, Fe³⁺, K⁺, Mg²⁺ and Cl^– –- responded to neither species nor depth. The six studied species may be clustered into three groups based on their effects on the soil properties. Scots pine and spruce had the lowest pH, highest C/N ratio and intermediate C content in soil. The other two coniferous species, Arolla pine and larch, had the highest soil C contents, highest pH values, and intermediate C/N ratios. Finally, the two deciduous hardwood species, aspen and birch, had the lowest C/N ratio, intermediate pH values, and lowest C content. These tree-mediated soil chemical changes are important for their likely effects on soil microbiological activities, including C and N mineralization and the production and consumption of greenhouse gases.     

Successional changes in soil nitrogen availability,non-symbiotic nitrogen fixation and carbon/nitrogen ratios in southern Chilean forest ecosystems

Vast areas of southern Chile are now covered by second-growth forests because of fire and logging. To study successional patterns after moderate-intensity, anthropogenic fire disturbance, we assessed differences in soil properties and N fluxes across a chronosequence of seven successional stands (2–130 years old). We examined current predictions of successional theory concerning changes in the N cycle in forest ecosystems. Seasonal fluctuations of net N mineralization (N_min) in surface soil and N availability (N_a; N_a=NH ₄ ⁺ –N+NO ₃ ^– –N) in upper and deep soil horizons were positively correlated with monthly precipitation. In accordance with theoretical predictions, stand age was positively, but weakly related to both N_a (r ²=0.282, P<0.001) and total N (N_tot; r ²=0.192, P<0.01), and negatively related to soil C/N ratios (r ²=0.187, P<0.01) in surface soils. A weak linear increase in soil N_min (upper plus deep soil horizons) was found across the chronosequence (r ²=0.124, P<0.022). N_min occurred at modest rates in early successional stands, suggesting that soil disturbance did not impair microbial processes. The relationship between N fixation (N_fix) in the litter layer and stand age best fitted a quadratic model (r ²=0.228, P<0.01). In contrast to documented successional trends for most temperate, tropical and Mediterranean forests, non-symbiotic N_fix in the litter layer is a steady N input to unpolluted southern temperate forests during mid and late succession, which may compensate for hydrological losses of organic N from old-growth ecosystems.     

Net aboveground biomass declines of four major forest types with forest ageing and climate change in western Canada's boreal forests

Biomass change of the world's forests is critical to the global carbon cycle. Despite storing nearly half of global forest carbon, the boreal biome of diverse forest types and ages is a poorly understood component of the carbon cycle. Using data from 871 permanent plots in the western boreal forest of Canada, we examined net annual aboveground biomass change (ΔAGB) of four major forest types between 1958 and 2011. We found that ΔAGB was higher for deciduous broadleaf (DEC) (1.44 Mg ha^?1 year^?1, 95% Bayesian confidence interval (CI), 1.22–1.68) and early‐successional coniferous forests (ESC) (1.42, CI, 1.30–1.56) than mixed forests (MIX) (0.80, CI, 0.50–1.11) and late‐successional coniferous (LSC) forests (0.62, CI, 0.39–0.88). ΔAGB declined with forest age as well as calendar year. After accounting for the effects of forest age, ΔAGB declined by 0.035, 0.021, 0.032 and 0.069 Mg ha^?1 year^?1 per calendar year in DEC, ESC, MIX and LSC forests, respectively. The ΔAGB declines resulted from increased tree mortality and reduced growth in all forest types except DEC, in which a large biomass loss from mortality was accompanied with a small increase in growth. With every degree of annual temperature increase, ΔAGB decreased by 1.00, 0.20, 0.55 and 1.07 Mg ha^?1 year^?1 in DEC, ESC, MIX and LSC forests, respectively. With every cm decrease of annual climatic moisture availability, ΔAGB decreased 0.030, 0.045 and 0.17 Mg ha^?1 year^?1 in ESC, MIX and LSC forests, but changed little in DEC forests. Our results suggest that persistent warming and decreasing water availability have profound negative effects on forest biomass in the boreal forests of western Canada. Furthermore, our results indicate that forest responses to climate change are strongly dependent on forest composition with late‐successional coniferous forests being most vulnerable to climate changes in terms of aboveground biomass.     

The effect of forest type on throughfall deposition and seepage flux: a review

Converting deciduous forests to coniferous plantations and vice versa causes environmental changes, but till now insight into the overall effect is lacking. This review, based on 38 case studies, aims to find out how coniferous and deciduous forests differ in terms of throughfall (+stemflow) deposition and seepage flux to groundwater. From the comparison of coniferous and deciduous stands at comparable sites, it can be inferred that deciduous forests receive less N and S via throughfall (+stemflow) deposition on the forest floor. In regions with relatively low open field deposition of atmospheric N (<10 kg N ha⁻¹ year⁻¹), lower NH₄⁺ mean throughfall (+stemflow) deposition was, however, reported under conifers compared to deciduous forest, while in regions with high atmospheric N pollution (>10 kg N ha⁻¹ year⁻¹), the opposite could be concluded. The higher the open field deposition of NH₄⁺, the bigger the difference between the coniferous and deciduous throughfall (+stemflow) deposition. Furthermore, it can be concluded that canopy exchange of K⁺, Ca²⁺ and Mg²⁺ is on average higher in deciduous stands. The significantly higher stand deposition flux of N and S in coniferous forests is reflected in a higher soil seepage flux of NO₃⁻, SO₄²⁻, K⁺, Ca²⁺, Mg²⁺ and Al(III). Considering a subset of papers for which all necessary data were available, a close relationship between throughfall (+stemflow) deposition and seepage was found for N, irrespective of the forest type, while this was not the case for S. This review shows that the higher input flux of N and S in coniferous forests clearly involves a higher seepage of NO₃⁻ and SO₄²⁻ and accompanying cations K⁺, Ca²⁺, Mg²⁺ and Al(III) into the groundwater, making this forest type more vulnerable to acidification and eutrophication compared to the deciduous forest type.     

Global Within-Site Variance in Soil Solution Nitrogen and Hydraulic Conductivity Are Correlated with Clay Content

Nutrient fluxes in terrestrial ecosystems are governed by complex biological and physical interactions. Ecologists’ mechanistic understanding of these interactions has focused on biological controls including plant uptake and microbial processing. However, ecologists and hydrologists have recently demonstrated that physical controls are also important. Here, we show that within-site spatial variation in soil solution N concentrations is a function of soil clay content across a globally diverse array of field sites. Clay content explained 35 and 53% of the coefficient of variation (CV) in soil solution nitrate (NO₃ ^?) and dissolved organic nitrogen (DON), respectively. The CV of soil hydraulic conductivity is a similar function of clay content, suggesting that soil hydrology may be a significant mechanism affecting variation in soil solution N. Although vegetation physiognomy and soil C/N ratios are known to affect soil solution N concentrations, neither was significantly related to within-site spatial variation in NO₃ ^? or DON. However, the spatial variation of NO₃ ^? and DON was greater in younger forests than in paired older forests. Our data show that the heterogeneity of an important resource, soil solution N, is a predictable function of clay content. Resource heterogeneity, such as that described here for soil solution N, can affect population, community, and ecosystem processes.     

Nitrogen Isotope Patterns in Alaskan Black Spruce Reflect Organic Nitrogen Sources and the Activity of Ectomycorrhizal Fungi

Global patterns in soil, plant, and fungal stable isotopes of N (δ¹⁵N) show promise as integrated metrics of N cycling, particularly the activity of ectomycorrhizal (ECM) fungi. At small spatial scales, however, it remains difficult to differentiate the underlying causes of plant δ¹⁵N variability and this limits the application of such measurements to better understand N cycling. We conducted a landscape-scale analysis of δ¹⁵N values from 31 putatively N-limited monospecific black spruce (Picea mariana) stands in central Alaska to assess the two main hypothesized sources of plant δ¹⁵N variation: differing sources and ECM fractionation. We found roughly 20% of the variability in black spruce foliar N and δ¹⁵N values to be correlated with the concentration and δ¹⁵N values of soil NH₄ ⁺ and dissolved organic N (DON) pools, respectively. However, ¹⁵N-based mixing models from 24 of the stands suggested that fractionation by ECM fungi obscures the ¹⁵N signature of soil N pools. Models, regressions, and N abundance data all suggested that increasing dependence on soil DON to meet black spruce growth demands predicates increasing reliance on ECM-derived N and that black spruce, on average, received 53% of its N from ECM fungi. Future research should partition the δ¹⁵N values within the soil DON pool to determine how choice of soil δ¹⁵N values influence modeled ECM activity. The C balance of boreal forests is tightly linked to N cycling and δ¹⁵N values may be useful metrics of changes to these connections.     

Controls on Soil Carbon Dioxide and Methane Fluxes in a Variety of Taiga Forest Stands in Interior Alaska

CO₂ and CH₄ fluxes were monitored over 4 years in a range of taiga forests along the Tanana River in interior Alaska. Floodplain alder and white spruce sites and upland birch/aspen and white spruce sites were examined. Each site had control, fertilized, and sawdust amended plots; flux measurements began during the second treatment year. CO₂ emissions decreased with successional age across the sites (alder, birch/aspen, and white spruce, in order of succession) regardless of landscape position. Although CO₂ fluxes showed an exponential relationship with soil temperature, the response of CO₂ production to moisture fit an asymptotic model. Of the manipulations, only N fertilization had an effect on CO₂ flux, decreasing flux in the floodplain sites but increasing it in the birch/aspen site. Landscape position was the best predictor of CH₄ flux. The two upland sites consumed CH₄ at similar rates (approximately 0.5 mg C m⁻² d⁻¹), whereas the floodplain sites had lower consumption rates (0–0.3 mg C m⁻² d⁻¹). N fertilization and sawdust both inhibited CH₄ consumption in the upland birch/aspen and floodplain spruce sites but not in the upland spruce site. The biological processes driving CO₂ fluxes were sensitive to temperature, moisture, and vegetation, whereas CH₄ fluxes were sensitive primarily to landscape position and biogeochemical disturbances. Hence, climate change effects on C-gas flux in taiga forest soils will depend on the relationship between soil temperature and moisture and the concomitant changes in soil nutrient pools and cycles. Received 10 March 1998; accepted 29 December 1999.     

Degradability, molecular weight and adsorption properties of dissolved organic carbon and nitrogen leached from different types of decomposing litter

Aims

We characterized dissolved organic matter (DOM) leached during decomposition of deciduous silver birch litter (Betula pendula Roth.), coniferous Norway spruce litter (Picea abies (L.) Karst.) and a mixture of these litters in order to find out whether the properties of DOM would explain the earlier observed signs for higher microbial activity in soil under birch than spruce.

Methods

DOM leached from decomposing litters was collected in a litter-column experiment in the laboratory. Adsorption properties (XAD-8 resin fractionation) and molecular weight as well as the degradability of dissolved organic carbon (DOC) and nitrogen (DON) were measured three times during decomposition: 1) in the early stages, 2) after the mass loss reached 20–30 % and 3) when the mass loss reached 30–40 %.

Results

The leaching of DOC hydrophilic neutrals and bases, regarded easily degradable, decreased during decomposition. The leaching of DOC in hydrophobic acids, regarded refractory, increased from spruce and especially from the mixture litter during decomposition and may be connected to the degree of litter decomposition that was highest for the mixture. Unexpectedly, the degradability of DOC differed only slightly between the litters but the degradability of DON was substantially higher for spruce than birch. Spruce DOM seemed to be more N-rich than birch DOM in the early stages of decomposition and it seemed that labile DON was mobilized earlier from spruce than birch litter.

Conclusions

We conclude that the decomposition degree of litter determines largely the properties of DOM. The observed differences in the properties of DOM sampled during the litter decomposition cannot explain differences in C and N cycling between birch and spruce.     

Persistent and pervasive compositional shifts of western boreal forest plots in Canada

Species compositional shifts have important consequences to biodiversity and ecosystem function and services to humanity. In boreal forests, compositional shifts from late‐successional conifers to early‐successional conifers and deciduous broadleaves have been postulated based on increased fire frequency associated with climate change truncating stand age‐dependent succession. However, little is known about how climate change has affected forest composition in the background between successive catastrophic fires in boreal forests. Using 1797 permanent sample plots from western boreal forests of Canada measured from 1958 to 2013, we show that after accounting for stand age‐dependent succession, the relative abundances of early‐successional deciduous broadleaves and early‐successional conifers have increased at the expense of late‐successional conifers with climate change. These background compositional shifts are persistent temporally, consistent across all forest stand ages and pervasive spatially across the region. Rising atmospheric CO₂ promoted early‐successional conifers and deciduous broadleaves, and warming increased early‐successional conifers at the expense of late‐successional conifers, but compositional shifts were not associated with climate moisture index. Our results emphasize the importance of climate change on background compositional shifts in the boreal forest and suggest further compositional shifts as rising CO₂ and warming will continue in the 21st century.     

The effect of soil freezing on N cycling: comparison of two headwater subcatchments with different vegetation and snowpack conditions in the northern Hokkaido Island of Japan

Climate change models predict that the snowpacks of temperate forests will develop later and be shallower resulting in a higher propensity for soil freezing. In the northern most island of Japan, Hokkaido, snowpack depth decreases from west to east. This snowpack depth gradient provided a unique opportunity to test the effects of variable snowpack and soil freezing on N biogeochemistry. The Shibecha Northern Catchment in Shibecha Experimental Forest, eastern Hokkaido had deciduous trees and a mean annual snowpack of 0.7 m while the M3 catchment in Uryu Experimental Forest, western Hokkaido had mixed deciduous and coniferous tree species and a mean annual snowpack of 2.0 m. We conducted a field study (October 2004–April 2005) to determine if differences in Shibecha and Uryu soil extractable N, N mineralization, and nitrification were controlled by the variability in soil freezing during winter or tree species composition that affected the quality of the forest floor. The mixed deciduous and coniferous trees forming the Uryu forest floor had a higher C:N ratio (25.0 vs. 22.4 at Shibecha), higher lignin:N ratio (15 vs. 8.8), and higher lignin concentrations (0.28 vs. 0.18 g lignin g⁻¹). These differences in forest floor quality contributed to higher net N mineralization and nitrification in Shibecha compared to Uryu. In Shibecha, soil remained frozen for the entire study. For Uryu, except for an early period with cold temperatures and no snow, the soil generally remained unfrozen. As a result of the early winter cold period and soil freezing, extractable soil NH₄⁺ did not change but NO₃⁻ increased. Reciprocal 0–5 cm mineral soil transplants made between Shibecha and Uryu and incubated during winter at 0, 5, and 30 cm suggested that soil freezing resulted in greater net N mineralization yet lower nitrification regardless of the soil origin. The effect of soil freezing should be considered when evaluating differences in N dynamics between temperate ecosystems having a propensity for soil freezing.     

Neutron radiography as a tool for revealing root development in soil: capabilities and limitations

Chronic atmospheric nitrogen deposition affects the cycling of carbon (C) and nitrogen (N) in forest ecosystems, and thereby alters the stable C isotopic abundance of plant and soil. Three successional stages, disturbed, rehabilitated and mature forests were studied for their responses to different nitrogen input levels. N-addition manipulative experiments were conducted at low, medium and high N levels. To study the responses of C cycling to N addition, the C concentration and ¹³C natural abundances for leaf, litter and soil were measured. Labile organic carbon fractions in mineral soils were measured to quantify the dynamics of soil organic C (SOC). Results showed that three-year continuous N addition did not significantly increase foliar C and N concentration, but decreased C/N ratio and enriched ¹³C in N-rich forests. In addition, N addition significantly decreased microbial biomass C, and increased water soluble organic C in surface soils of N-rich forests. This study suggests that N addition enhances the water consumption per unit C assimilation of dominant plant species, restricts SOC turnover in N-poor forests at early and medium successional stages (thus favored SOC sequestration), and vice versa for N-rich mature forests.     

Effects of above-ground browsing by mammals on mycorrhizal infection in an early successional taiga ecosystem

Using an exclosure experiment in the willow stage of primary succession on the floodplain of the Tanana River, we tested the hypothesis that browsing can reduce mycorrhizal infection. We measured the effects winter browsing by moose (Alcesalces) and snowshoe hare (Lepusamericanus) had on mycorrhizal infection and fine root biomass of willow (Salix spp.) and balsam poplar (Populusbalsamifera). We found that protection from winter browsing increased ectomycorrhizal infection by 10% in the top 5 cm of the soil profile, by 23% at 5–10 cm, and by 42% at the 10–15 cm depth. Mammal browsing in taiga forests is now recognized as a major cause of the shift from palatable deciduous species such as willow and balsam poplar to less palatable species such as alder and spruce. We suggest that browsing-induced reduction in ectomycorrhizal infection of salicaceous species plays a central role in this shift in plant community composition. Received: 26 March 1996 / Accepted: 26 September 1996