Adaptation to exploit nitrate in surface soils predisposes yellow-cedar to climate-induced decline while enhancing the survival of western redcedar: A new hypothesis

Yellow-cedar (Chamaecyparis nootkatensis (D. Don) Spach) and western redcedar (Thuja plicata Donn), two valuable tree species of Pacific Northwest forests, are competitive in low productivity forests on wet, nearly saturated soils with low nitrogen (N) availability and turnover. We propose a mechanism where cedar trees survive in marginal conditions through exploiting a coupled Ca–NO₃⁻ nutrient cycle where trees assimilate N as nitrate (NO₃⁻), but must accumulate a counter-ion to NO₃⁻ such as calcium (Ca⁺²) to control their internal cell pH and provide electrochemical balance. The availability of NO₃⁻ in cedar forests is favored by increased microbial activity and shifts in microbial community composition that is conducive to N mineralization and nitrification at higher pH. Cedars influence the soils under their canopy by enriching the forest floor with calcium compounds leading to increases in pH. Cedars are also prone to precocious dehardening in the spring when N is released from freeze–thaw events in the soils and conditions appear to favor nitrifying microbial communities. Cedars must concentrate fine-root biomass near the soil surface to access Ca and NO₃⁻, but this beneficial physiological adaptation also creates a vulnerability to periodic root freezing injury that is leading to the decline and mortality of at least one of them—yellow-cedar.     

Seasonal and spatial variation of nitrogen dynamics in the litter and surface soil layers on a tropical dry evergreen forest slope

Seasonal and spatial variability of litterfall and NO₃⁻ and NH₄⁺ leaching from the litter layer and 5-cm soil depth were investigated along a slope in a tropical dry evergreen forest in northeastern Thailand. Using ion exchange resin and buried bag methods, the vertical flux and transformation of inorganic nitrogen (N) were observed during four periods (dry, early wet, middle wet, and late wet seasons) at 15 subplots in a 180-m × 40-m rectangular plot on the slope. Annual N input via litterfall and inorganic N leached from the litter layer and from 5-cm depth soil were 12.5, 6.9, and 3.7 g N m⁻² year⁻¹, respectively, whereas net mineralization and the inorganic N pool in 0–5-cm soil were 7.1 g N m⁻² year⁻¹ and 1.4 g N m⁻², respectively. During the early wet season (90 days), we observed 82% and 74% of annual NO₃⁻ leaching from the litter layer and 5-cm soil depth, respectively. Higher N input via leaf litterfall in the dry season and via precipitation in the early wet season may have led to higher NO₃⁻ leaching rate from litter and surface soil layers during the early wet season. Large spatial variability in both NO₃⁻ vertical flux and litterfall was also observed within stands. Small-scale spatial patterns of total N input via litterfall were significantly correlated with NO₃⁻ leaching rate from the surface soil layer. In tropical dry evergreen forests, litterfall variability may be crucial to the remarkable seasonal changes and spatial variation in annual NO₃⁻ vertical flux in surface soil layers.     

Gross N transformations were little affected by 4 years of simulated N and S depositions in an aspen-white spruce dominated boreal forest in Alberta, Canada

The effects of 4 years of simulated nitrogen (N) and sulfur (S) depositions on gross N transformations in a boreal forest soil in the Athabasca oil sands region (AOSR) in Alberta, Canada, were investigated using the ¹⁵N pool dilution method. Gross NH₄⁺ transformation rates in the organic layer tended to decline (P < 0.10, marginal statistical significance, same below) in the order of control (CK, i.e., no N or S addition), +N (30 kg N ha⁻¹ yr⁻¹), +S (30 kg S ha⁻¹ yr⁻¹), and +NS treatments, with an opposite trend in the mineral soil. Gross NH₄⁺ immobilization rates were generally higher than gross N mineralization rates across the treatments, suggesting that the studied soil still had potential for microbial immobilization of NH₄⁺, even after 4 years of elevated levels of simulated N and S depositions. For both soil layers, N addition tended to increase (P < 0.10) the gross nitrification and NO₃⁻ immobilization rates. In contrast, S addition reduced (P < 0.001) and increased (P < 0.001) gross nitrification as well as tended (P < 0.10) to reduce and increase gross NO₃⁻ immobilization rates in the organic and mineral soils, respectively. Gross nitrification and gross NO₃⁻ immobilization rates were tightly coupled in both soil layers. The combination of rapid NH₄⁺ cycling, negligible net nitrification rates and the small NO₃⁻ pool size after 4 years of elevated N and S depositions observed here suggest that the risk of NO₃⁻ leaching would be low in the studied boreal forest soil, consistent with N leaching measurements in other concurrent studies at the site that are reported elsewhere.     

Soil nitrogen transformations varied with plant community under Nanchang urban forests in mid-subtropical zone of China

Soil N transformations using the polyvinyl chloride (PVC) closed-top tube in situ incubation method were studied in Nanchang urban forests of the mid-subtropical region of China in different months of 2007. Four plots of 20 m × 20 m were established in four different plant communities that represented typical successional stages of forest development including shrubs, coniferous forest, mixed forest and broad- leaved forest. Average concentrations of soil NH 4 + -N from January to December were not different among the four plant communities. The concentrations of soil NO 3 - -N and mineral N, and the annual rates of ammonification, nitrification and net N-mineralization under the early successional shrub community and coniferous forest were generally lower than that of the late successional mixed and broad-leaved forests (p<0.05). Similar differences among the plant communities were also shown in the relative nitrification index (NH 4 + -N/NO 3 - -N) and relative nitrification intensity (nitrification rate/net N-mineralization rate). The annual net N-mineralization rate was increased from younger to older plant communities, from 15.1 and 41.4 kg·ha -1 ·a -1 under the shrubs and coniferous forest communities to 98.0 and 112.9 kg·ha -1 ·a -1 under the mixed and broad-leaved forests, respectively. Moreover, the high annual nitrification rates (50-70 kg·ha -1 ·a -1 ) and its end product, NO 3 - -N (2.4-3.8 mg·kg -1 ), under older plant communities could increase the potential risk of N loss. Additionally, the temporal patterns of the different soil N variables mentioned above varied with different plant community due to the combined affects of natural biological processes associated withforest maturation and urbanization. Our results indicated that urban for- ests are moving towards a state of "N saturation" (extremely nitrification rate and NO 3 - -N content) as they mature.     

Gross nitrification rates in four Japanese forest soils: heterotrophic versus autotrophic and the regulation factors for the nitrification

Measurements of gross NH ₄ ⁺ and NO ₃ ^? production in forest soils were conducted using the ¹⁵N pool dilution method. Mineral topsoils (0?C10?cm depth) were collected from four forests from northern to southern Japan with a natural climate gradient to elucidate the mechanisms regulating gross nitrification rates in forest soils. Additionally, we attempted to evaluate the relative importance of heterotrophic nitrification in gross total nitrification using acetylene as a specific inhibitor of autotrophic nitrification. Distinct differences were found among sites in the gross rates of NH ₄ ⁺ production (3.1?C11.4?mg?N?kg^?1?day^?1) and gross total nitrification (0.0?C6.1?mg?N?kg^?1?day^?1). The rates of gross heterotrophic nitrification were low in this study, indicating that heterotrophic nitrification is of minor importance in most forest mineral topsoils in Japan. Significant relations were found between gross autotrophic nitrification and gross NH ₄ ⁺ production, soil N, and soil C concentrations, but none was found between gross autotrophic nitrification and soil pH. We determined the critical value of the gross NH ₄ ⁺ production rates for gross autotrophic nitrification under which no gross autotrophic nitrification occurred, as well as the critical soil C/N ratio above which gross autotrophic nitrification ceased. Results show that tight coupling of production and consumption of NH ₄ ⁺ prevents autotrophic nitrifiers from utilizing NH ₄ ⁺ as long as NH ₄ ⁺ availability is low.     

Soil N cycling in old-growth forests across an Andosol toposequence in Ecuador

Nitrogen (N) deposition in the tropics is predicted to increase drastically in the next decades. The sparse information on N cycling in tropical forests revealed that the soil N status of an ecosystem is the key to analyze its reactions to projected increase in N input. Our study was aimed at (1) comparing the soil N availability of forest sites across an Ecuadorian Andosol toposequence by quantifying gross rates of soil N cycling in situ, and (2) determining the factors controlling the differences in soil N cycling across sites. The toposequence was represented by five old-growth forest sites with elevations ranging from 300 m to 1500 m. Our results provide general insights into the role of elevation-mediated factors (i.e. degree of soil development and temperature) in driving patterns of soil N cycling. Gross rates of N transformations, microbial N turnover time, and δ¹⁵N signatures in soil and leaf litter decreased with increasing elevation, signifying a decreasing N availability across the toposequence. This was paralleled by a decreasing degree of soil development with increasing elevation, as indicated by declining clay contents, total C, total N, effective cation exchange capacity and increasing base saturation. Soil N-cycling rates and δ¹⁵N signatures were highly correlated with mean annual temperature but not with mean annual rainfall and soil moisture which did not systematically vary across the toposequence. Microbial immobilization was the largest fate of produced NH₄⁺ across all sites, and nitrification activity was only 5–11% of gross NH₄⁺ production. We observed a fast reaction of NO₃⁻ to organic N and its role for N retention deserves further attention. If projected increase in N deposition will occur, the timing and magnitude of gaseous N losses may follow the pattern of N availability across this Andosol toposequence.     

Influence of variable organic matter retention on nutrient availability in a 10-year-old loblolly pine plantation

The effects of varying forest floor and slash retention at time of regeneration were evaluated 10 years after the establishment of a loblolly pine plantation near Millport, Alabama. Treatments included removing, leaving unaltered, or doubling the forest floor and slash material. Forest floor and litterfall mass and nutrient concentrations, available soil N, foliar nutrient concentrations and stand yield were all impacted by the treatments. Forest floor mass and nutrient contents in the doubled treatment were significantly greater than the other two treatments. The doubled treatment accumulated 25, 45 and 350% more forest floor mass and 56, 56, and 310% more N than the control treatment in the Oi, Oe, and Oa layers, respectively. The other nutrients followed similar patterns. Potentially mineralized NO₃⁻-N in the mineral soil was also significantly higher in the doubled treatment. The positive effect of doubling the forest floor on soil N availability was reflected in greater foliage production, 30% more litterfall and 25% more stand yield for this treatment. This study shows that increasing the forest floor retention has resulted in increased nutrient availability and improved tree growth.     

Tracing the fate of mineral N compounds under high ambient N deposition in a Norway spruce forest at Solling/Germany

The fate of high and equally distributed ammonium and nitrate deposition was followed in a 72-year-old roofed Norway spruce forest at Solling in central Germany by separately adding ¹⁵NH₄⁺ and ¹⁵NO₃⁻ to throughfall water since November 2001. The objective was to quantify the retention of atmospheric ammonium and nitrate in different ecosystem compartments as well as the leaching loss from the forest ecosystem. δ¹⁵N excess in tree tissues (needles, twigs, branches and bole woods) decreased with increased tissue age. Clear ¹⁵N signals in old tree tissues indicated that the added ¹⁵N was not only assimilated to newly produced tree tissues but also retranslocated to old ones. During a period of over 3-year ¹⁵N addition, 30% of ¹⁵NH₄⁺ and 36% of ¹⁵NO₃⁻ were found in tree compartments. For both ¹⁵N tracers, 15% of added ¹⁵N was found in needles, followed by woody tissues (twigs, branches and boles, 7–13%) and live fine roots (7%). The recovery of ¹⁵NH₄⁺ and ¹⁵NO₃⁻ in the live fine roots differed with soil depth. The recovery of ¹⁵NH₄⁺ tended to be higher in the live fine roots in the organic layer than in the upper mineral soil. In the live fine roots in deeper soil, the recovery of ¹⁵NO₃⁻ tended to be higher than that of ¹⁵NH₄⁺. Soil retained the largest proportion of ¹⁵N, accounting for 71% of ¹⁵NH₄⁺ and 42% of ¹⁵NO₃⁻. Most of ¹⁵NH₄⁺ was recovered in the organic layer (65%) and the recovery decreased with soil depth. Conversely, only 8% of ¹⁵NO₃⁻ was found in the organic layer and 34% of ¹⁵NO₃⁻ was evenly distributed throughout the mineral soil layers. Nitrate leaching accounted for 3% of ¹⁵NH₄⁺ and 19% of ¹⁵NO₃⁻. Only less than 1% of the both added ¹⁵N was leached as DON. These results suggested that trees had a high contribution to the retention of atmospheric N and soil retention capacity determined the loss of atmospheric N by nitrate leaching.     

Post-fire soil nitrogen content and vegetation composition in Sub-Boreal spruce forests of British Columbia's central interior,Canada

Forest fires are known to influence nutrient cycling, particularly soil nitrogen (N), as well as plant succession in northern forest ecosystems. However, few studies have addressed the dynamics of soil N and its relationship to vegetation composition after fire in these forests. To investigate soil N content and vegetation establishment after wildfire, 13 sites of varying age class were selected in the Sub-Boreal spruce zone of the central interior of British Columbia, Canada. Sites varied in time since the last forest fire and were grouped into three seral age classes: (a) early-seral (<14 years), (b) mid-seral (50–80 years) and (c) late-seral (>140 years). At each site, we estimated the percent cover occupied by trees, shrubs, herbs and mosses. In addition, the soil samples collected from the forest floor and mineral horizons were analyzed for the concentrations of total N, mineralizable N, available NO₃⁻-N and available NH₄⁺-N. Results indicated that soil N in both the forest floor and mineral horizons varied between the three seral age classes following wildfire. Significant differences in mineralizable N, available NO₃⁻-N and available NH₄⁺-N levels with respect to time indicated that available soil N content changes after forest fire. Percent tree and shrub cover was significantly correlated to the amount of available NH₄⁺-N and mineralizable N contents in the forest floor. In the mineral horizons, percent tree cover was significantly correlated to the available NH₄⁺-N, while herb cover was significantly correlated with available NO₃⁻-N. Moss cover was significantly correlated with total N, available NO₃⁻-N and mineralizable N in the forest floor and available NO₃⁻-N in the mineral horizons. We identified several unique species of shrubs and herbs for each seral age class and suggest that plant species are most likely influencing the soil N levels by their contributions to the chemical composition and physical characteristics of the organic matter.     

Silvopastoral use of Nothofagus antarctica in Southern Patagonian forests, influence over net nitrogen soil mineralization

In most temperate forest, nitrogen (N) is considered a limiting factor. This becomes important in extreme environments, as Nothofagus antarctica forests, where the antecedents are scarce. Thinning practices in N. antarctica forests for silvopastoral uses may modify the soil N dynamics. Therefore, the objective of this work was to evaluate the temporal variation of soil N in these ecosystems. The mineral extractable soil N, net nitrification and net N mineralization were evaluated under different crown cover and two site quality stands. The mineral N extractable (NH₄ ⁺–N + NO₃ ^?–N) was measured periodically. Net nitrification and net N mineralization were estimated through the technique of incubation of intact samples with tubes. The total mineral extractable N concentration varied between crown cover and dates, with no differences among site classes. The lowest and highest values were found in the minimal and intermediate crown cover, respectively. In the higher site quality stand, the annual net N mineralization was lower in the minimal crown cover reaching 11 kg N ha^?1 year^?1, and higher in the maximal crown cover (54 kg N ha^?1 year^?1). In the lower site quality stand there was no differences among crown cover. The same pattern was found for net nitrification. Thinning practices for silvopastoral use of these forests, keeping intermediate crown cover values, did not affect both N mineralization and nitrification. However, the results suggest that total trees removal from the ecosystem may decrease N mineralization and nitrification.     

Gap formation in Danish beech (Fagus sylvatica) forests of low management intensity: soil moisture and nitrate in soil solution

Soil moisture content (0–90 cm depth) and nitrate-nitrogen (NO₃-N) concentrations in soil solution (90 cm depth) were monitored after gap formation (diameter 15–18 m) in three Danish beech-dominated forests on nutrient-rich till soils. NO₃-N drainage losses were estimated by the water balance model WATBAL for one of the sites. Two forests were non-intervention forests (semi-natural and unmanaged), the third was subject to nature-based management. The study was intended to assess the range of effects of gap formation in forests of low management intensity. In the unmanaged and the nature-based managed forest, soil solution was collected for 5 years and soil moisture measured in the fourth year after gap formation. Average NO₃-N concentrations were significantly higher in the gaps (9.9 and 8.1 mg NO₃-N l⁻¹, respectively) than under closed canopy (0.2 mg l⁻¹). In the semi-natural forest, measurements were carried out up to 29 months after gap formation. Average NO₃-N concentrations in the gap were 19.3 mg NO₃-N l⁻¹. Gap formation alone did not account for this high level, as concentrations were high also under closed canopy (average 12.4 mg NO₃-N l⁻¹). However, the gap had significantly higher N concentrations when trees were in full leaf, and NO₃-N drainage losses were significantly increased in the gap. No losses occurred under closed canopy in growing seasons. Soil moisture was close to field capacity in all three gaps, but decreased under closed canopy in growing seasons. In the semi-natural forest, advanced regeneration and lateral closure of the gap affected soil moisture levels in the gap in the last year of the study.     

Carbon,nitrogen and phosphorus leaching after site preparation at a boreal forest clear-cut area

Clear-cutting followed by mechanical site preparation is the major disturbance influencing nutrient and water fluxes in Fennoscandian boreal forests. The effects of soil harrowing on the fluxes of dissolved organic carbon (DOC), dissolved nitrogen compounds (organic N, NH₄⁺ and NO₃⁻) and water soluble phosphorus (PO₄³⁻) through a podzolic soil were studied in a clear-cut in eastern Finland for 5 years. The old, mixed coniferous stand was clear-cut and stem only harvested in 1996 followed by soil harrowing in 1998 and planting in June 1999. Zero-tension lysimeters were used to collect soil water from below different soil horizons in the three types of microsites that resulted from site preparation treatment: low ridges (25% of clear-cut area), shallow furrows (30%) and the undisturbed soil (45%). After soil harrowing, the leaching of DOC, N and P from below the B-horizon increased compared to pre-treatment levels. However, the increases were short-lasting; 1–2 years for inorganic N and P, and 5 years for DOC and organic N. The highest concentrations were associated with the ridges and lowest with the furrows, reflecting the differences in amount of organic matter present in each microsite type and, for N, to enhanced mineralization and nitrification. Leaching from below the B-horizon over the 5 years following soil harrowing for the whole clear-cut area was 36.5 kg ha⁻¹ for DOC, 0.88 kg ha⁻¹ for NH₄-N, 0.46 kg ha⁻¹ for NO₃-N, 1.24 kg ha⁻¹ for organic N and 0.09 kg ha⁻¹ for PO₄-P. Site preparation increased temporarily the risk for nutrient leaching into watercourses and groundwater from the clear-cut area but soil fertility was not affected since the leached amounts remained small. The main reasons for the observed low leaching values were the rapid recovery of ground vegetation and low N deposition loads.     

Soil NH4 +/NO3 − nitrogen characteristics in primary forests and the adaptability of some coniferous species

In terrestrial ecosystems, soil nutrient regimes at a plant’s living site generally represent the plant’s “nutrition habitat”. Plant species frequently well adapt to their original “nutrition habitat” during a long process of evolution, and the apparent preference for ammonium or nitrate nitrogen source (NH₄ ⁺ or NO₃ ⁻) might be an important aspect of the adaptation. Plants typically favor the nitrogen form most abundant in their natural habitats. Nitrate has been recognized as the dominant mineral nitrogen form in most agricultural soils and the main nitrogen source for crops, but it is not usually the case in forest ecosystems. A large number of studies show that the “nutrition habitats” associated with primary forest soils are typically dominated by NH₄ ⁺ rather than NO₃ ⁻, generally with NO₃ ⁻ content much lower than NH₄ ⁺. Low levels of NO₃ ⁻ in these forest soils generally correspond to low net rates of nitrification. The probable reasons for this phenomenon include: 1) nitrification limitations and/or inhibitions caused by lower pH, lower NH₄ ⁺ availability (autotrophic nitrifiers cannot successfully compete for NH₄ ⁺ with heterotrophic organisms and plants), or allelopathic inhibitors (tannins or higher-molecular-weight proanthocyanidins) in the soil; or 2) substantial microbial acquisition of nitrate in the soils, which makes net nitrification rates substantially less than gross nitrification rates even though the latter are relatively high. Many coniferous species (especially such late successional tree species as Tsuga heterophylla, Pinus banksiana, Picea glauca, Pseudotsuga meziesii, Picea abies, etc.) fully adapt to their original NH₄ ⁺-dominated “nutrition habitats” so that their capacities of absorbing and using non-reduced forms of nitrogen (e.g., NO₃ ⁻) substantially decrease. These conifers typically show distinct preference to NH₄ ⁺ and reduced growth due to nitrogen-metabolism disorder when NO₃ ⁻ is the main nitrogen source. The physiological and biochemical mechanisms that account for the adaptation to NH₄ ⁺-dominated systems (or limited ability to use NO₃ ⁻) for the coniferous species include: i) distribution and activity of enzymes for catalyzing nitrogen reduction and assimilation, generally characterized by lower nitrate reductase (NR); ii) greater tolerance to NH₄ ⁺ or rapid detoxification of ammonium nitrogen in the roots; iii) lower capacity of absorption to NO₃ ⁻ by roots that might be controlled by feedback regulations of certain N-transport compounds, such as glutamine; iv) relations and balance between nitrogen and other elements (such as Ca²⁺, Mg²⁺, and Zn²⁺ etc.). Some NH₄ ⁺-preferred conifers might be more adapted (tolerant) to lower base cation conditions; v) NO₃ ⁻ nutrition, rather than NH₄ ⁺, that may lead to the loss of considerable quantities of organic and inorganic carbon to the surrounding media and mycorrhizal symbiont and probably contribute to slower growth; and vi) the metabolic cost of reducing NO₃ ⁻ to NH₄ ⁺ that may make shade-tolerant conifers favor the uptake of reduced nitrogen (NH₄ ⁺). The adaptation of late successional conifers to NH₄ ⁺-dominated habitats has profound ecological implications. First, it might be an important prerequisite for the climax forest communities dominated by these conifers to maintain long-term stability. Second, primary coniferous or coniferous-broadleaved forests have been widely perturbed because of commercial exploitation, where the soil ammonium nitrogen pool tends to be largely transformed to nitrate after disturbance. In such a situation, the coniferous species that were dominant in undisturbed ecosystems may become poor competitors for nitrogen, and the site will be occupied by early successional (pioneer) plants better adapted to nitrate utilization. In other words, the implicit adaptation of many conifers dominant in undisturbed communities to ammonium nitrogen will cause difficulties in their regeneration on disturbed sites, which must be taken into account in the practical restoration of degraded temperate forest ecosystems. __________ Translated from Acta Ecologica Sinica, 2005, 25(11): 3,082–3,092 [译自: 生态学报]     

Reduced soil respiration in gaps in logged lowland dipterocarp forests

We studied the effects of forest composition and structure, and related biotic and abiotic factors on soil respiration rates in a tropical logged forest in Malaysian Borneo. Forest stands were classified into gap, pioneer, non-pioneer and mixed (pioneer, non-pioneer and unclassified trees) based on the species composition of trees >10 cm diameter breast height. Soil respiration rates did not differ significantly between non-gap sites (1290 ± 210 mg CO₂ m⁻² h⁻¹) but were double those in gap sites (640 ± 130 mg CO₂ m⁻² h⁻¹). Post hoc analyses found that an increase in soil temperature and a decrease in litterfall and fine root biomass explained 72% of the difference between gap and non-gap sites. The significant decrease of soil respiration rates in gaps, irrespective of day or night time, suggests that autotrophic respiration may be an important contributor to total soil respiration in logged forests. We conclude that biosphere-atmosphere carbon exchange models in tropical systems should incorporate gap frequency and that future research in tropical forest should emphasize the contribution of autotrophic respiration to total soil respiration.     

Does soil organic matter variation affect the retention of NH4 and NO3 in forest ecosystems?

Human activities have fundamentally changed global nitrogen (N) cycling, leading to elevated N deposition in most parts of the world. The fate of deposited N, whether being retained to sustain plant growth or causing ecosystem N saturation, is critical to the global carbon (C) cycling and local environment. In a short-term laboratory experiment, we used ¹⁵N-labeled NH₄⁺ and NO₃⁻ to study the fate of N inputs in forest soils and what regulates N retention. Soils with a wide range of organic matter content and other attributes were collected from a 70-year-old plantation containing monotypic stands of Norway spruce (Picea abies), red pine (Pinus resinosa), sugar maple (Acer saccharum), and red oak (Quercus rubra), and separated into 0-5 cm and 5-15 cm layers. Nitrogen added to the soil was either immediately extracted (Time 0: T0) with K₂SO₄ solution, or incubated for 7 d (T7) and then extracted. Retention of ¹⁵N into the non-extractable soil pool at T0 was limited; but after the 7-d incubation, between 20 and 70% of the ¹⁵NH₄⁺ was retained. Nitrification transformed on average 50% of the ¹⁵NH₄⁺ into ¹⁵NO₃⁻ during the incubation while retention of ¹⁵NO₃⁻ at T7 remained low (7.40 ± 1.08%). Retention of ¹⁵NH₄⁺ into non-extractable soil at T7 was positively correlated to the percentage of soil organic matter (SOM) (r² = 0.323, P < 0.001), and was significantly higher (P < 0.001) in the high-SOM 0-5 cm layer than in the low-SOM 5-15 cm layer. Conversion of ¹⁵NH₄⁺ to ¹⁵NO₃⁻ during incubation significantly reduced the ¹⁵NH₄⁺ retention (P < 0.001). Our results suggest that the variations of SOM and other soil attributes play strong roles in the retention of newly deposited inorganic N and could affect forest ecosystem responses to chronic N deposition.     

Response of the N and P cycles of an old-growth montane forest in Ecuador to experimental low-level N and P amendments

Atmospheric nitrogen (N) and phosphorus (P) depositions are expected to increase in the tropics as a consequence of increasing human activities in the next decades. In the literature, it is frequently assumed that tropical montane forests are N-limited, while tropical lowland forests are P-limited. In a low-level N and P addition experiment, we determined the short-term response of N and P cycles in a north Andean montane forest on Palaeozoic shists and metasandstones at an elevation of 2100 m a.s.l. to increased N and P inputs. We evaluated experimental N, P and N + P additions (50 kg ha⁻¹ yr⁻¹ of N, 10 kg ha⁻¹ yr⁻¹ of P and 50 kg + 10 kg ha⁻¹ yr⁻¹ of N and P, respectively) and an untreated control in a fourfold replicated randomized block design. We collected litter leachate, mineral soil solution (0.15 and 0.30 m depths), throughfall and litterfall before the treatment began (August 2007) until 16 months after the first nutrient application (April 2009). Less than 10 and 1% of the applied N and P, respectively, leached below the organic layer which contained almost all roots and no significant leaching losses of N and P occurred to below 0.15 m mineral soil depth. Deposited N and P from the atmosphere in dry and wet form were retained in the canopy of the control treatment using a canopy budget model. Nitrogen and P retention by the canopy were reduced and N and P fluxes in throughfall and litterfall increased in their respective treatments. The increase in N and P fluxes in throughfall after fertilization was equivalent to 2.5% of the applied N and 2% of the applied P. The fluxes of N and P in litterfall were up to 15% and 3%, respectively, higher in the N and N + P than in the control treatments. We conclude that the expected elevated N and P deposition in the tropics will be retained in the ecosystem, at least in the short term and hence, N and P concentrations in stream water will not increase. Our results suggest that in the studied tropical montane forest ecosystem on Palaeozoic bedrock, N and P are co-limiting the growth of organisms in the canopy and organic layer.     

Soil solution chemistry at different positions on slope in a conifer plantation forest

It is known that soil property varies along the slope. It suggests that soil solution chemistry also differs topographically. To determine the variation in soil solution chemistry within one watershed, soil solution chemistry at the different positions of the slope was investigated. Soil N transformation changed along the slope. NH₄ ⁺ ratio to inorganic N (NH₄ ⁺ + NO₃ ⁻) increased upslope. The tendency was verified by laboratory incubation. After incubation most of the mineralized N was nitrified at the lower part of the slope, while little nitrification occurred at the upper part of the slope. At the ridge and the backslope inorganic N form in soil solution was concomitant with inorganic N form by incubation. At the ridge NH₄ ⁺ was predominant form in soil solution, at that time major anion was sea salt originated Cl⁻. From this, soil solution chemistry seems to be regulated by the external nutrient source at the ridge. In the second year of lysimeter installation NO₃ ⁻ concentration increased in both sites and the ratio of NH₄ ⁺ to inorganic N decreased. It was considered due to the effect of lysimeter installation. The lag time and the magnitude of NO₃ ⁻ increase were different between the ridge and the backslope. It would be related with soil N transformation in pre-disturbance. The influence of disturbance were shown in other solute concentrations of soil solution.     

Does tree species composition control soil organic carbon pools in Mediterranean mountain forests?

We compared soil organic carbon (SOC) stocks and stability under two widely distributed tree species in the Mediterranean region: Scots pine (Pinus sylvestris L.) and Pyrenean oak (Quercus pyrenaica Willd.) at their ecotone. We hypothesised that soils under Scots pine store more SOC and that tree species composition controls the amount and biochemical composition of organic matter inputs, but does not influence physico-chemical stabilization of SOC. At three locations in Central Spain, we assessed SOC stocks in the forest floor and down to 50 cm in the mineral in pure and mixed stands of Pyrenean oak and Scots pine, as well as litterfall inputs over approximately 3 years at two sites. The relative SOC stability in the topsoil (0-10 cm) was determined through size-fractionation (53 μm) into mineral-associated and particulate organic matter and through KMnO₄-reactive C and soil C:N ratio.Scots pine soils stored 95-140 Mg ha⁻¹ of C (forest floor plus 50 cm mineral soil), roughly the double than Pyrenean oak soils (40-80 Mg ha⁻¹ of C), with stocks closely correlated to litterfall rates. Differences were most pronounced in the forest floor and uppermost 10 cm of the mineral soil, but remained evident in the deeper layers. Biochemical indicators of soil organic matter suggested that biochemical recalcitrance of soil organic matter was higher under pine than under oak, contributing as well to a greater SOC storage under pine. Differences in SOC stocks between tree species were mainly due to the particulate organic matter (not associated to mineral particles). Forest conversion from Pyrenean oak to Scots pine may contribute to enhance soil C sequestration, but only in form of mineral-unprotected soil organic matter.     

Nitrogen in soils beneath 18–65 year old stands of subtropical evergreen broad-leaved forests in Laoshan Mountains in Eastern China

Monitoring of soil nitrogen (N) cycling is useful to assess soil quality and to gauge the sustainability of management practices. We studied net N mineralization, nitrification, and soil N availability in the 0 10 cm and 11 30 cm soil horizons in east China during 2006 2007 using an in situ incubation method in four subtropical evergreen broad-leaved forest stands aged 18-, 36-, 48-, and 65-years. The proper- ties of surface soil and forest floor varied between stand age classes. C:N ratios of surface soil and forest floor decreased, whereas soil total N and total organic C, available P, and soil microbial biomass N increased with stand age. The mineral N pool was small for the young stand and large for the older stands. NO 3 - -N was less than 30% in all stands. Net rates of N mineralization and nitrification were higher in old stands than in younger stands, and higher in the 0 10 cm than in the 11 30 cm horizon. The differences were significant between old and young stands (p < 0.031) and between soil horizons (p < 0.005). Relative nitrification was somewhat low in all forest stands and declined with stand age. N trans- formation seemed to be controlled by soil moisture, soil microbial bio- mass N, and forest floor C:N ratio. Our results demonstrate that analyses of N cycling can provide insight into the effects of management distur- bances on forest ecosystems.     

Decomposing stumps influence carbon and nitrogen pools and fine-root distribution in soils

Decomposing stumps could significantly increase soil resource heterogeneity in forest ecosystems. However, the impact of these microsites on nutrient retention and cycling is relatively unknown. Stump soil was defined as the soil fraction directly altered by the decomposition of the primary rooting system (e.g. taproots) and aboveground stumps. Study sites were located in mature hardwood stands within the Jefferson National Forest in the Ridge and Valley Physiographic region of southwest Virginia. The objectives of this study were to: (i) determine the total soil volume altered by the decomposition of stumps and underlying root system, (ii) compare and contrast total C and N, extractable ammonium (NH₄⁺) and nitrate (NO₃⁻), potentially mineralizable N, microbial biomass C (MBC), root length and root surface area between the bulk soil (i.e. O, A, B and C horizons) and stump soil and (iii) evaluate how nutrient concentrations and fine-root dynamics change as stumps decompose over time using a categorical decay class system for stumps. Potentially mineralizable N was 2.5 times greater in stump soil than the A horizon (103 mg kg⁻¹ vs. 39 mg kg⁻¹), 2.7 times greater for extractable NH₄⁺ (16 mg kg⁻¹ vs. 6 mg kg⁻¹) and almost 4 times greater for MBC (1528 mg kg⁻¹ vs. 397 mg kg⁻¹). Approximately 19% of the total fine-root length and 14% of fine-root surface area occurred in the stump soil. Significant differences occurred in C and N concentrations between all four decay classes and the mineral soil. This validated the use of this system and the need to calculate weighted averages based on the frequency and soil volume influenced by each decay class. In this forest ecosystem, approximately 1.2% of the total soil volume was classified as stump soil and contained 10% and 4% of soil C and N. This study illustrates that including stump soil in soil nutrient budgets by decay class will increase the accuracy of ecosystem nutrient budgets.