Intact amino acid uptake by northern hardwood and conifer trees

Empirical and modeling studies of the N cycle in temperate forests of eastern North America have focused on the mechanisms regulating the production of inorganic N, and assumed that only inorganic forms of N are available for plant growth. Recent isotope studies in field conditions suggest that amino acid capture is a widespread ecological phenomenon, although northern temperate forests have yet to be studied. We quantified fine root biomass and applied tracer-level quantities of U–¹³C₂–¹⁵N-glycine, ¹⁵NH₄ ⁺ and ¹⁵NO₃ ⁻ in two stands, one dominated by sugar maple and white ash, the other dominated by red oak, beech, and hemlock, to assess the importance of amino acids to the N nutrition of northeastern US forests. Significant enrichment of ¹³C in fine roots 2 and 5 h following tracer application indicated intact glycine uptake in both stands. Glycine accounted for up to 77% of total N uptake in the oak–beech–hemlock stand, a stand that produces recalcitrant litter, cycles N slowly and has a thick, amino acid-rich organic horizon. By contrast, glycine accounted for only 20% of total N uptake in the sugar maple and white ash stand, a stand characterized by labile litter and rapid rates of amino acid production and turnover resulting in high rates of mineralization and nitrification. This study shows that amino acid uptake is an important process occurring in two widespread, northeastern US temperate forest types with widely differing rates of N cycling.     

Differential effects of sugar maple, red oak, and hemlock tannins on carbon and nitrogen cycling in temperate forest soils

Tannins are abundant secondary chemicals in leaf litter that are hypothesized to slow the rate of soil-N cycling by binding protein into recalcitrant polyphenol–protein complexes (PPCs). We studied the effects of tannins purified from sugar maple, red oak, and eastern hemlock leaf litter on microbial activity and N cycling in soils from northern hardwood–conifer forests of the northeastern US. To create ecologically relevant conditions, we applied tannins to soil at a concentration (up to 2 mg g⁻¹ soil) typical of mineral soil horizons. Sugar maple tannins increased microbial respiration significantly more than red oak or hemlock tannins. The addition of sugar maple tannins also decreased gross N mineralization by 130% and, depending upon the rate of application, decreased net rates of N mineralization by 50–290%. At low concentrations, the decrease in mineralization appeared to be driven by greater microbial-N immobilization, while at higher concentrations the decrease in mineralization was consistent with the formation of recalcitrant PPCs. Low concentrations of red oak and hemlock tannins stimulated microbial respiration only slightly, and did not significantly affect fluxes of inorganic N in the soil. When applied to soils containing elevated levels of protein, red oak and hemlock tannins decreased N mineralization without affecting rates of microbial respiration, suggesting that PPC formation decreased substrate availability for microbial immobilization. Our results indicate that tannins from all three species form recalcitrant PPCs, but that the degree of PPC formation and its attendant effect on soil-N cycling depends on tannin concentration and the pool size of available protein in the soil.     

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

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

Detritus Production and Soil N Transformations in Old-Growth Eastern Hemlock and Sugar Maple Stands

To examine the linkage between forest cover type, litter inputs, and patterns of net N mineralization versus the turnover of N among soil microbes, we measured both the net and gross rates of N mineralization in replicated, adjacent old-growth eastern hemlock [Tsuga canadensis(L.) Carr.] or sugar maple (Acer saccharum Marsh.) stands in upper Michigan. Mean aboveground net primary production and annual litterfall mass were significantly higher (P < 0.01) in the maple forests (870 g·m^-2·y^-1 and 439 g·m^-2·y^-1, respectively) than in the hemlock forests (480 g·m^-2·y^-1 and 344 g·m^-2·y^-1, respectively). Forest floor and coarse woody debris mass, however, were significantly lower (P < 0.05) in the maple forests (2.2 and 0.1 kg·m^-2, respectively) than in the hemlock forests (2.9 and 0.2 kg·m^-2, respectively). Litterfall N concentration was not significantly different (P > 0.10) between the two forest types. In situ gross rates of N mineralization were higher (P < 0.06) in the maple forests than in the hemlock forests (7.5 and 6.1 mg N·kg soil^-1·d^-1 respectively), but in situ net N mineralization varied independently of forest type and stand-level litterfall N concentration. Cover type–dependent differences in detritus production and detritus C quality appear to result in different N turnover rates, but the balance between gross mineralization and immobilization of N is very sensitive to within stand variability and varies at a scale smaller than cover type alone can predict. Received 3 Feburary 1999; accepted 27 August 1999.     

The phosphorus status of northern hardwoods differs by species but is unaffected by nitrogen fertilization

Northern hardwood forests in the eastern US exhibit species-specific influences on nitrogen (N) cycling, suggesting that their phosphorus (P) cycling characteristics may also vary by species. These characteristics are increasingly important to understand in light of evidence suggesting that atmospheric N deposition has increased N availability in the region, potentially leading to phosphorus limitation. We examined how P characteristics differ among tree species and whether these characteristics respond to simulated N deposition (fertilization). We added NH₄NO₃ fertilizer (50 kg ha^?1 year^?1) to single-species plots of red oak (Quercus rubra L.), sugar maple (Acer saccharum Marsh.), eastern hemlock (Tsuga canadensis (L.) Carr.), American beech (Fagus grandifolia Ehrh.), and yellow birch (Betula alleghaniensis Britt.), in the Catskill Mountains, New York from 1997 to 2007. Species differences were observed in foliar, litter and root P concentrations, but all were unaffected by a cumulative N fertilization of 550 kg/ha. Similarly, measures of soil P availability and biotic P sufficiency differed by species but were unaffected by N fertilization. Results suggest species exhibit unique relationships to P as well as N cycles. We found little evidence that N fertilization leads to increased P limitation in these northern hardwood forests. However, species such as sugar maple and red oak may be sufficient in P, whereas beech and hemlock may be less sufficient and therefore potentially more sensitive to future N-stimulated P limitation.     

Influence of Tree Species on Forest Nitrogen Retention in the Catskill Mountains, New York, USA

This study examines the effect of four tree species on nitrogen (N) retention within forested catchments of the Catskill Mountains, New York (NY). We conducted a 300-day ¹⁵N field tracer experiment to determine how N moves through soil, microbial, and plant pools under different tree species and fertilization regimes. Samples were collected from single-species plots of American beech (Fagus grandifolia Ehrh.), eastern hemlock (Tsuga canadensis L.), red oak (Quercus rubra L.), and sugar maple (Acer saccharum Marsh). Using paired plots we compared the effects of ambient levels of N inputs (11 kg N/ha/y) to additions of 50 kg N/ha/y that began 1.5 years prior to and continued throughout this experiment. Total plot ¹⁵N recovery (litter layer, organic and mineral soil to 12 cm, fine roots, and aboveground biomass) did not vary significantly among tree species, but the distribution of sinks for ¹⁵N within the forest ecosystem did vary. Recovery in the forest floor was significantly lower in sugar maple stands compared to the other species. ¹⁵Nitrogen recovery was 22% lower in the fertilized plots compared to the ambient plots and red oak stands had the largest drop in ¹⁵N recovery as a result of N fertilization. Aboveground biomass became a significantly greater ¹⁵N sink with fertilization, although it retained less than 1% of the tracer addition. These results indicate that different forest types vary in the amount of N retention in the forest floor, and that forest N retention may change depending upon N inputs.     

Biomass and morphology of fine roots in temperate broad-leaved forests differing in tree species diversity: is there evidence of below-ground overyielding?

Biodiversity effects on ecosystem functioning in forests have only recently attracted increasing attention. The vast majority of studies in forests have focused on above-ground responses to differences in tree species diversity, while systematic analyses of the effects of biodiversity on root systems are virtually non-existent. By investigating the fine root systems in 12 temperate deciduous forest stands in Central Europe, we tested the hypotheses that (1) stand fine root biomass increases with tree diversity, and (2) ‘below-ground overyielding’ of species-rich stands in terms of fine root biomass is the consequence of spatial niche segregation of the roots of different species. The selected stands represent a gradient in tree species diversity on similar bedrock from almost pure beech forests to medium-diverse forests built by beech, ash, and lime, and highly-diverse stands dominated by beech, ash, lime, maple, and hornbeam. We investigated fine root biomass and necromass at 24 profiles per stand and analyzed species differences in fine root morphology by microscopic analysis. Fine root biomass ranged from 440 to 480 g m⁻² in the species-poor to species-rich stands, with 63–77% being concentrated in the upper 20 cm of the soil. In contradiction to our two hypotheses, the differences in tree species diversity affected neither stand fine root biomass nor vertical root distribution patterns. Fine root morphology showed marked distinctions between species, but these root morphological differences did not lead to significant differences in fine root surface area or root tip number on a stand area basis. Moreover, differences in species composition of the stands did not alter fine root morphology of the species. We conclude that ‘below-ground overyielding’ in terms of fine root biomass does not occur in the species-rich stands, which is most likely caused by the absence of significant spatial segregation of the root systems of these late-successional species.     

Soil carbon storage, litterfall and CO2 efflux in fertilized and unfertilized larch (Larix leptolepis) plantations

This study evaluated the effects of forest fertilization on the forest carbon (C) dynamics in a 36-year-old larch (Larix leptolepis) plantation in Korea. Above- and below-ground C storage, litterfall, root decomposition and soil CO₂ efflux rates after fertilization were measured for 2 years. Fertilizers were applied to the forest floor at rates of 112 kg N ha⁻¹ year⁻¹, 75 kg P ha⁻¹ year⁻¹ and 37 kg K ha⁻¹ year⁻¹ for 2 years (May 2002, 2003). There was no significant difference in the above-ground C storage between fertilized (41.20 Mg C ha⁻¹) and unfertilized (42.25 Mg C ha⁻¹) plots, and the C increment was similar between the fertilized (1.65 Mg C ha⁻¹ year⁻¹) and unfertilized (1.52 Mg C ha⁻¹ year⁻¹) plots. There was no significant difference in the soil C storage between the fertilized and unfertilized plots at each soil depth (0–15, 15–30 and 30–50 cm). The organic C inputs due to litterfall ranged from 1.57 Mg C ha⁻¹ year⁻¹ for fertilized to 1.68 Mg C ha⁻¹ year⁻¹ for unfertilized plots. There was no significant difference in the needle litter decomposition rates between the fertilized and unfertilized plots, while the decomposition of roots with 1–2 mm diameters increased significantly with the fertilization relative to the unfertilized plots. The mean annual soil CO₂ efflux rates for the 2 years were similar between the fertilized (0.38 g CO₂ m⁻² h⁻¹) and unfertilized (0.40 g CO₂ m⁻² h⁻¹) plots, which corresponded with the similar fluctuation in the organic carbon (litterfall, needle and root decomposition) and soil environmental parameters (soil temperature and soil water content). These results indicate that little effect on the C dynamics of the larch plantation could be attributed to the 2-year short-term fertilization trials and/or the soil fertility in the mature coniferous plantation used in this study.     

Carbon, nitrogen and phosphorus in volcanic soils following afforestation with native birch (Betula pubescens) and introduced larch (Larix sibirica) in Iceland

Afforestation has become an important tool for soil protection and land reclamation in Iceland. Nevertheless, the harsh climate and degraded soils are growth-limiting for trees, and little is know about changes in soil nutrients in maturing forests planted on the volcanic soils. In the present chronosequence study, changes in C, N and total P in soil (0–10 and 10–20 cm depth) and C and N in foliar tissue were investigated in stands of native Downy birch (Betula pubescens Enrh.) and the in Iceland introduced Siberian larch (Larix sibirica Ledeb.). The forest stands were between 14 and 97 years old and were established on heath land that had been treeless for centuries. Soils were Andosols derived from basaltic material and rhyolitic volcanic ash. A significant effect of tree species was only found for the N content in foliar tissue. Foliar N concentrations were significantly higher and foliar C/N ratios significantly lower in larch needles than in birch leaves. There was no effect of stand age. Changes in soil C and the soil nutrient status with time after afforestation were little significant. Soil C concentrations in 0–10 cm depth in forest stands older than 30 years were significantly higher than in heath land and forest stands younger than 30 years. This was attributed to a slow accumulation of organic matter. Soil N concentrations and soil P_tot were not affected by stand age. Nutrient pools in the two soil layers were calculated for an average weight of soil material (400 Mg soil ha⁻¹ in 0–10 cm depth and 600 Mg soil ha⁻¹ in 10–20 cm depth, respectively). Soil nutrient pools did not change significantly with time. Soil C pools were in average 23.6 Mg ha⁻¹ in the upper soil layer and 16.9 Mg ha⁻¹ in the lower soil layer. The highest annual increase in soil C under forest compared to heath land was 0.23 Mg C ha⁻¹ year⁻¹ in 0–10 cm depth calculated for the 53-year-old larch stand. Soil N pools were in average 1.0 Mg N ha⁻¹ in both soil layers and did not decrease with time despite a low N deposition and the uptake and accumulation of N in biomass of the growing trees. Soil P_tot pools were in average 220 and 320 kg P ha⁻¹ in the upper and lower soil layer, respectively. It was assumed that mycorrhizal fungi present in the stands had an influence on the availability of N and P to the trees. Responsible Editor: Hans Lambers.     

Nitrogen deposition effects on soil organic matter chemistry are linked to variation in enzymes,ecosystems and size fractions

Recent research has dramatically advanced our understanding of soil organic matter chemistry and the role of N in some organic matter transformations, but the effects of N deposition on soil C dynamics remain difficult to anticipate. We examined soil organic matter chemistry and enzyme kinetics in three size fractions (>250 μm, 63–250 μm, and <63 μm) following 6 years of simulated atmospheric N deposition in two ecosystems with contrasting litter biochemistry (sugar maple, Acer saccharum—basswood, Tilia americana and black oak, Quercus velutina—white oak, Q. alba). Ambient and simulated (80-kg NO₃ ⁻–N ha⁻¹ year⁻¹) atmospheric N deposition were studied in three replicate stands in each ecosystem. We found striking, ecosystem-specific effects of N deposition on soil organic matter chemistry using pyrolysis gas chromatography/mass spectrometry. First, furfural, the dominant pyrolysis product of polysaccharides, was significantly decreased by simulated N deposition in the sugar maple–basswood ecosystem (15.9 vs. 5.0%) but was increased by N deposition in the black oak–white oak ecosystem (8.8 vs. 24.0%). Second, simulated atmospheric N deposition increased the ratio of total lignin derivatives to total polysaccharides in the >250 μm fraction of the sugar maple–basswood ecosystem from 0.9 to 3.3 but there were no changes in other size classes or in the black oak–white oak ecosystem. Third, simulated N deposition increased the ratio of lignin derivatives to N-bearing compounds in the 63–250 and >250 μm fractions in both ecosystems but not in the <63 μm fraction. Relationships between enzyme kinetics and organic matter chemistry were strongest in the particulate fractions (>63 μm) where there were multiple correlations between oxidative enzyme activities and concentrations of lignin derivatives and between glycanolytic enzyme activities and concentrations of carbohydrates. Within silt-clay fractions (<63 μm), these enzyme-substrate correlations were attenuated by interactions with particle surfaces. Our results demonstrate that variation in enzyme activity resulting from atmospheric N deposition is directly linked to changes in soil organic matter chemistry, particularly those that occur within coarse soil size fractions. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Differential Snowpack Accumulation and Water Dynamics in Aspen and Conifer Communities: Implications for Water Yield and Ecosystem Function

Early succession aspen and late succession conifer forests have different architecture and physiology affecting hydrologic transfer processes. An evaluation of water pools and fluxes was used to determine differences in the hydrologic dynamics between stands of quaking aspen (Populus tremuloides) and associated stands of mixed conifer consisting of white fir (Abies concolor), Douglas-fir (Pseudotsuga menziesii), and Engelmann spruce (Picea engelmannii). In 2005 and 2006, measurements of snow water accumulation, snow ablation (melt), soil water content, snowpack sublimation, and evapotranspiration (ET) were measured in adjacent aspen and conifer stands. Peak snow water equivalent (SWE) averaged 34–44% higher in aspen in 2005 (average snow fall) and 2006 (above average snow fall), respectively, whereas snow ablation rates were greater in aspen stands (21 mm day⁻¹) compared to conifer stands (11 mm day⁻¹). When changes in soil water content (due to over-winter snowmelt) were combined with peak snow accumulation in 2006, aspen had greater potential (42–83%) water yield for runoff and groundwater recharge. Snowpack sublimation during the ablation period was not significantly different between meadow, aspen, and conifer sites and comprised less than 5% of the winter precipitation. Extended conifer transpiration in spring and fall did not contribute to large differences in water yield (<28 mm y⁻¹). Summertime ET rates were higher in aspen plots (3.6 mm day⁻¹) than in conifer plots (2.7 mm day⁻¹), and differences in net ET largely reflected soil column porosity. This study shows that the largest differences in annual water yield between aspen and conifer stands result from differences in SWE and net summertime ET. Although SWE and accumulation of water in soil was greater in aspen, it was partly offset by greater net annual ET losses in aspen.     

Recent carbon and nitrogen accumulation and acidification in soils of two Scots pine ecosystems in Southern Germany

Changes of the soil chemical status during the recent 22–30 years at two historically degraded forest sites in southern Germany (Pfaffenwinkel, Pustert) stocked with mature Scots pine (Pinus sylvestris L.) stands were studied by repeated soil inventories conducted in 1974, 1982–1984, 1994, and 2004 on replicated control plots of fertilization experiments, allowing a statistical analysis. Additionally, the nutritional status of the stands at all plots was monitored from 1964 until 2004 by annual or bi-annual analysis of current-year foliage, and stand growth was assessed by repeated stand inventories carried out in 3- to 9-year intervals. For both sites, a statistically significant systematic decrease of the forest floor C/N ratio between 1974 and 2004 from 35.4 to 29.2 (Pfaffenwinkel) and from 36.5 to 23.0 (Pustert) was observed. The soils at both sites also showed a considerable accumulation of organic carbon (210 and 400 kg C ha⁻¹ year⁻¹ for Pfaffenwinkel and Pustert, respectively) and nitrogen (13 and 18 kg N ha⁻¹ year⁻¹). In addition, the mineral topsoil at both sites has acidified considerably, indicated by significantly decreased pH values (Pustert only; mean decrease 0.1 units per decade), base saturation, and base cation stocks. The trend of N enrichment and base cation loss in the soils is mirrored by the trends of stand nutrition at both sites, which are characterized by improved N nutrition and reduced supply with K, Mg (Pustert only), and Ca. The results confirm findings of other studies indicating a recent N eutrophication and acidification of forest soils in Central Europe and southern Scandinavia. Since soils with historic degradation due to earlier non-sustainable forest utilization are widespread in Central Europe, the results obtained on our study sites probably apply for large forested areas, suggesting a significant potential of Central European forests to sequester atmospheric carbon and nitrogen not only in stand biomass, but also in the soil.     

Large Loss of Dissolved Organic Nitrogen from Nitrogen-Saturated Forests in Subtropical China

Dissolved organic nitrogen (DON) has recently been recognized as an important component of terrestrial N cycling, especially under N-limited conditions; however, the effect of increased atmospheric N deposition on DON production and loss from forest soils remains controversial. Here we report DON and dissolved organic carbon (DOC) losses from forest soils receiving very high long-term ambient atmospheric N deposition with or without additional experimental N inputs, to investigate DON biogeochemistry under N-saturated conditions. We studied an old-growth forest, a young pine forest, and a young mixed pine/broadleaf forest in subtropical southern China. All three forests have previously been shown to have high nitrate (NO₃⁻) leaching losses, with the highest loss found in the old-growth forest. We hypothesized that DON leaching loss would be forest specific and that the strongest response to experimental N input would be in the N-saturated old-growth forest. Our results showed that under ambient deposition (35–50 kg N ha⁻¹ y⁻¹ as throughfall input), DON leaching below the major rooting zone in all three forests was high (6.5–16.9 kg N ha⁻¹ y⁻¹). DON leaching increased 35–162% following 2.5 years of experimental input of 50–150 kg N ha⁻¹ y⁻¹. The fertilizer-driven increase of DON leaching comprised 4–17% of the added N. A concurrent increase in DOC loss was observed only in the pine forest, even though DOC:DON ratios declined in all three forests. Our data showed that DON accounted for 23–38% of total dissolved N in leaching, highlighting that DON could be a significant pathway of N loss from forests moving toward N saturation. The most pronounced N treatment effect on DON fluxes was not found in the old-growth forest that had the highest DON loss under ambient conditions. DON leaching was highly correlated with NO₃⁻ leaching in all three forests. We hypothesize that abiotic incorporation of excess NO₃⁻ (through chemically reactive NO₂⁻) into soil organic matter and the consequent production of N-enriched dissolved organic matter is a major mechanism for the consistent and large DON loss in the N-saturated subtropical forests of southern China. Dr. YT Fang performed research, analyzed data, and wrote the paper; Prof. WX Zhu participated in the initial experimental design, analyzed data, and took part in writing the paper; Prof. P Gundersen conceived the study and took part in writing; Prof. JM Mo and Prof. GY Zhou conceived study; Prof. M Yoh analyzed part of the data and contributed to the development of DON model.     

N:P Ratio and the Nature of Nutrient Limitation in Calluna-Dominated Heathlands

There is growing evidence from different sources that prolonged high N deposition causes a shift from nitrogen (N) limitation to nitrogen and phosphorus (P) co-limitation or even P limitation in many terrestrial ecosystems. However, the number of ecosystems where the type of limitation has been directly tested by longer-term full-factorial field experiments is very limited. We conducted a 5-year fertilization experiment with N and P in the Lüneburger Heide (NW Germany) to test the hypothesis that, following decades of elevated atmospheric N inputs, plant growth in dry lowland heaths may have shifted from N to N–P co-limitation or P limitation. We also tested whether the plant tissue N:P ratio reflects the type of nutrient limitation in a continental lowland heathland. Experimental plots dominated by Calluna vulgaris received regular additions of N (50 kg N ha⁻¹ y⁻¹), P (20 kg P ha⁻¹ y⁻¹), a combination of both, or water only (control) from 2004 to 2008. Over the whole study period, a highly significant positive N effect on shoot length was found, thus indicating N limitation. We conclude that a clear shift from N limitation to N–P co-limitation or P limitation has not yet occurred. Tissue N:P ratios showed a high temporal variability and no relationship between tissue N:P ratio and the shoot length response of Calluna to nutrient addition was found. The N:P tool is thus of limited use at the local scale and within the range of N:P ratio observed in this study, and should only be used as a rough indicator for the prediction of the type of nutrient limitation in lowland heathland on a larger geographical scale with a broader interval of N:P ratio.     

Total nitrogen and phosphorus balance sheet in a typic ustochrept soil under intensive cropping and fertilizer use

Summary Balance sheets were computed for total nitrogen and phosphorus in plough layer (0–15 cm) of a Typic Ustochrept soil under continuous multiple cropping for seven years (1971–72 to 1977–78) with a fixed rotation of pearl millet (Pennisetum typhoideum L.) wheat (Triticum aestivum L.) (Vigna sinensis Savi.) The treatments considered of soil test-based rates of N, P and K, applied both singly and in combinations together with farm yard manure, sulphur and zinc superimposed over optimum rates (100%) of NPK. Heavy, losses of N (762–899 kg ha⁻¹) occurred in the plots which received high rates of Nviz. 150% of recommended NPK and 100% NPK plus FYM. Application of N alone accelerated N losses whereas addition of P, PK, PKS to N minimised such losses. Enrichment of P (66 to 198 kg ha⁻¹) occurred in all phosphate-treated plots. A marginal net decrease (29–54 kg ha⁻¹) in P levels was observed in control and N alone treatments.     

Soil Inorganic N Leaching in Edges of Different Forest Types Subject to High N Deposition Loads

We report on soil leaching of dissolved inorganic nitrogen (DIN) along transects across exposed edges of four coniferous and four deciduous forest stands. In a 64-m edge zone, DIN leaching below the main rooting zone was enhanced relative to the interior (at 128 m from the edge) by 21 and 14 kg N ha⁻¹ y⁻¹ in the coniferous and deciduous forest stands, respectively. However, the patterns of DIN leaching did not univocally reflect those of DIN throughfall deposition. DIN leaching in the first 20 m of the edges was lower than at 32–64 m from the edge (17 vs. 36 kg N ha⁻¹ y⁻¹ and 15 vs. 24 kg N ha⁻¹ y⁻¹ in the coniferous and deciduous forests, respectively). Nitrogen stocks in the mineral topsoil (0–30 cm) were, on average, 943 kg N ha⁻¹ higher at the outer edges than in the interior, indicating that N retention in the soil is probably one of the processes involved in the relatively low DIN leaching in the outer edges. We suggest that a complex of edge effects on biogeochemical processes occurs at the forest edges as a result of the interaction between microclimate, tree dynamics (growth and litterfall), and atmospheric deposition of N and base cations.     

Base-cation Cycling by Individual Tree Species in Old-growth Forests of Upper Michigan,USA

The influence of individual tree species on base-cation (Ca, Mg, K, Na) distribution and cycling was examined in sugar maple (Acer saccharum Marsh.), basswood (Tilia americana L.), and hemlock (Tsuga canadensis L.) in old-growth northern hardwood – hemlock forests on a sandy, mixed, frigid, Typic Haplorthod over two growing seasons in northwestern Michigan. Base cations in biomass, forest floor, and mineral soil (0–15 cm and 15–40 cm) pools were estimated for five replicated trees of each species; measured fluxes included bulk precipitation, throughfall, stemflow, litterfall, forest-floor leachate, mineralization + weathering, shallow-soil leachate, and deep-soil leachate. The three species differed in where base cations had accumulated within the single-tree ecosystems. Within these three single-tree ecosystems, the greatest quantity of base cations in woody biomass was found in sugar maple, whereas hemlock and basswood displayed the greatest amount in the upper 40 cm of mineral soil. Base-cation pools were ranked: sugar maple > basswood, hemlock in woody biomass; sugar maple, basswood > hemlock in foliage; hemlock > sugar maple, basswood in the forest floor, and basswood > sugar maple, hemlock in the mineral soil. Base-cation fluxes in throughfall, stemflow, the forest-floor leachate, and the deep-soil leachate (2000 only) were ranked: basswood > sugar maple > hemlock. Our measurements suggest that species-related differences in nutrient cycling are sufficient to produce significant differences in base-cation contents of the soil over short time intervals (<65 years). Moreover, these species-mediated differences may be important controls over the spatial pattern and edaphic processes of northern hardwood-hemlock ecosystems in the upper Great Lakes region.     

Root Growth and Recovery in Temperate Broad-Leaved Forest Stands Differing in Tree Species Diversity

In contrast to studies on aboveground processes, the effect of species diversity on belowground productivity and fine-root regrowth after disturbance is still poorly studied in forests. In 12 old-growth broad-leaved forest stands, we tested the hypotheses that (i) the productivity and recovery rate (regrowth per standing biomass) of the fine-root system (root diameter < 2 mm) increase with increasing tree species diversity, and that (ii) the seasonality of fine-root biomass and necromass is more pronounced in pure than in tree species-rich stands as a consequence of non-synchronous root biomass peaks of the different species. We investigated stands with 1, 3, and 5 dominant tree species growing under similar soil and climate conditions for changes in fine-root biomass and necromass during a 12-month period and estimated fine-root productivity with two independent approaches (ingrowth cores, sequential coring). According to the analysis of 360 ingrowth cores, fine-root growth into the root-free soil increased with tree species diversity from 72 g m⁻² y⁻¹ in the monospecific plots to 166 g m⁻² y⁻¹ in the 5-species plots, indicating an enhanced recovery rate of the root system after soil disturbance with increasing species diversity (0.26, 0.34, and 0.51 y⁻¹ in 1-, 3-, and 5-species plots, respectively). Fine-root productivity as approximated by the sequential coring data also indicated a roughly threefold increase from the monospecific to the 5-species stand. We found no indication of a more pronounced seasonality of fine-root mass in species-poor as compared to species-rich stands. We conclude that species identification on the fine root level, as conducted here, may open new perspectives on tree species effects on root system dynamics. Our study produced first evidence in support of the hypothesis that the fine-root systems of more diverse forest stands are more productive and recover more rapidly after soil disturbance than that of species-poor forests.     

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

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

Impact of long-term nitrogen addition on carbon stocks in trees and soils in northern Europe

The aim of this study was to quantify the effects of fertiliser N on C stocks in trees (stems, stumps, branches, needles, and coarse roots) and soils (organic layer +0–10 cm mineral soil) by analysing data from 15 long-term (14–30 years) experiments in Picea abies and Pinus sylvestris stands in Sweden and Finland. Low application rates (30–50 kg N ha⁻¹ year⁻¹) were always more efficient per unit of N than high application rates (50–200 kg N ha⁻¹ year⁻¹). Addition of a cumulative amount of N of 600–1800 kg N ha⁻¹ resulted in a mean increase in tree and soil C stock of 25 and 11 kg (C sequestered) kg⁻¹ (N added) (“N-use efficiency”), respectively. The corresponding estimates for NPK addition were 38 and 11 kg (C) kg⁻¹ (N). N-use efficiency for C sequestration in trees strongly depended on soil N status and increased from close to zero at C/N 25 in the humus layer up to 40 kg (C) kg⁻¹ (N) at C/N 35 and decreased again to about 20 kg (C) kg⁻¹ (N) at C/N 50 when N only was added. In contrast, addition of NPK resulted in high (40–50 kg (C) kg⁻¹ (N)) N-use efficiency also at N-rich (C/N 25) sites. The great difference in N-use efficiency between addition of NPK and N at N-rich sites reflects a limitation of P and K for tree growth at these sites. N-use efficiency for soil organic carbon (SOC) sequestration was, on average, 3–4 times lower than for tree C sequestration. However, SOC sequestration was about twice as high at P. abies as at P. sylvestris sites and averaged 13 and 7 kg (C) kg⁻¹ (N), respectively. The strong relation between N-use efficiency and humus C/N ratio was used to evaluate the impact of N deposition on C sequestration. The data imply that the 10 kg N ha⁻¹ year⁻¹ higher deposition in southern Sweden than in northern Sweden for a whole century should have resulted in 2.0 ± 1.0 (95% confidence interval) kg m⁻² more tree C and 1.3 ± 0.5 kg m⁻² more SOC at P. abies sites in the south than in the north for a 100-year period. These estimates are consistent with differences between south and north in tree C and SOC found by other studies, and 70–80% of the difference in SOC can be explained by different N deposition.