Experimental evidence of nutrient limitations on primary productivity in Afrotropical forests is rare and globally underrepresented yet are crucial for understanding constraints to terrestrial carbon ...uptake. In an ecosystem‐scale nutrient manipulation experiment, we assessed the early responses of tree growth rates among different tree sizes, taxonomic species, and at a community level in a humid tropical forest in Uganda. Following a full factorial design, we established 32 (eight treatments × four replicates) experimental plots of 40 × 40 m each. We added nitrogen (N), phosphorus (P), potassium (K), their combinations (NP, NK, PK, and NPK), and control at the rates of 125 kg N ha−1 year−1, 50 kg P ha−1 year−1 and 50 kg K ha−1 year−1, split into four equal applications, and measured stem growth of more than 15,000 trees with diameter at breast height (dbh) ≥1 cm. After 2 years, the response of tree stem growth to nutrient additions was dependent on tree sizes, species and leaf habit but not community wide. First, tree stem growth increased under N additions, primarily among medium‐sized trees (10–30 cm dbh), and in trees of Lasiodiscus mildbraedii in the second year of the experiment. Second, K limitation was evident in semi‐deciduous trees, which increased stem growth by 46% in +K than –K treatments, following a strong, prolonged dry season during the first year of the experiment. This highlights the key role of K in stomatal regulation and maintenance of water balance in trees, particularly under water‐stressed conditions. Third, the role of P in promoting tree growth and carbon accumulation rates in this forest on highly weathered soils was rather not pronounced; nonetheless, mortality among saplings (1–5 cm dbh) was reduced by 30% in +P than in –P treatments. Although stem growth responses to nutrient interaction effects were positive or negative (likely depending on nutrient combinations and climate variability), our results underscore the fact that, in a highly diverse forest ecosystem, multiple nutrients and not one single nutrient regulate tree growth and aboveground carbon uptake due to varying nutrient requirements and acquisition strategies of different tree sizes, species, and leaf habits.
Estimates of asymbiotic biological N fixation (BNF) in temperate grasslands are few with large variations. In the past six decades, European grasslands have been subjected to intensive management ...practices and presently it is not known how asymbiotic BNF is influenced by these practices. Our objective was to assess the impact of fertilizer application and mowing frequency on asymbiotic BNF in a Central European grassland. In 2008, we established a three-factorial experiment with two fertilizer treatments (no fertilizer application and combined nitrogen (N), phosphorus (P) and potassium (K) fertilization at 180–30–100 kg ha−1 yr−1), two mowing frequencies (cut once and thrice per year) and three sward compositions through the application of herbicides (control, monocot- and dicot-enhanced swards). Three years after the initial sward manipulation, there was no more difference in functional group composition. Between June 2011 and May 2012, we measured in-situ asymbiotic BNF using the acetylene reduction assay, calibrated with 15N2-fixation method. Across treatments, asymbiotic BNF rates in the 0–5-cm soil depth ranged from 1.7 (±0.2 SE) kg ha−1 yr−1 for fertilized plots cut once a year to 5.7 (±2.3 SE) kg ha−1 yr−1 for unfertilized plots cut thrice a year. Fertilization decreased asymbiotic BNF, suggesting that the potential positive effect of increased soil P levels might have been overruled by the negative effect of increased soil mineral N levels. Intensive mowing stimulated asymbiotic BNF, which was probably due to an increase in rhizodeposition. Our calibration of the acetylene reduction assay with the 15N2-fixation method resulted in a conversion factor of 0.61, which largely deviates from the theoretical conversion factor of 3. Furthermore, laboratory incubations under increased soil moisture and temperature conditions overestimated BNF rates compared to in-situ measurements. Thus, laboratory measurements with altered soil moisture, temperature or disturbed soil may lead to strong biases in estimates of asymbiotic BNF. Our results suggest that input of N through BNF may be considerable in temperate grasslands. We conclude that BNF studies should be conducted in-situ and that the acetylene reduction assay should be calibrated against 15N2-fixation calibration for reliable estimates.
•We tested grassland management effects (mowing/fertilization) on asymbiotic BNF.•Conversion factors (acetylene reduction: N2 fixation) were clearly below the theoretical value of three.•Intensive mowing increased and combined fertilization of N, P, and K decreased BNF.•Asymbiotic BNF is an important input of N to unfertilized temperate grasslands.•Previous BNF estimates from temperate grasslands may be substantially biased.
Intensive management practices in large-scale oil palm plantations can slow down nutrient cycling and alter other soil functions. Thus, there is a need to reduce management intensity without ...sacrificing productivity. The aim of our study was to investigate the effect of management practices on gross rates of soil N cycling and soil fertility. In Jambi province, Indonesia, we established a management experiment in a large-scale oil palm plantation to compare conventional practices (i.e. high fertilization rates and herbicide weeding) with reduced management intensity (i.e. reduced fertilization rates and mechanical weeding). Also, we compared the typical management zones characterizing large-scale plantations: palm circle, inter-row and frond-stacked area. After 1.5 years of this experiment, reduced and conventional management showed comparable gross soil N cycling rates; however, there were stark differences among management zones. The frond-stacked area had higher soil N cycling rates and soil fertility (high microbial biomass, extractable C, soil organic C, extractable organic N, total N and low bulk density) than inter-row and palm circle (all p ≤ 0.05). Microbial biomass was the main driver of the soil N cycle, attested by its high correlation with gross N-cycling rates (r = 0.93–0.95, p < 0.01). The correlations of microbial N with extractable C, extractable organic N, soil organic C and total N (r = 0.76–0.89, p < 0.01) suggest that microbial biomass was mainly regulated by the availability of organic matter. Mulching with senesced fronds enhanced soil microbial biomass, which promoted nutrient recycling and thereby can decrease dependency on chemical fertilizers.
Oil palm plantations are intensively managed agricultural systems that increasingly dominate certain tropical regions. Oil palm monocultures have been criticized because of their reduced biodiversity ...compared to the forests they historically replaced, and because of their negative impact on soils, water, and climate. We experimentally test whether less intensive management schemes may enhance biodiversity and lessen detrimental effects on the environment while maintaining high yields. We compare reduced vs. conventional fertilization, as well as mechanical vs. chemical weed control (with herbicides) in a long-term, full-factorial, multidisciplinary experiment. We conducted the experiment in an oil palm company estate in Sumatra, Indonesia, and report the results of the first 2 years. We measured soil nutrients and functions, surveyed above- and below-ground organisms, tracked oil palm condition and productivity, and calculated plantation gross margins. Plants, aboveground arthropods, and belowground animals were positively affected by mechanical vs. chemical weed control, but we could not detect effects on birds and bats. There were no detectable negative effects of reduced fertilization or mechanical weeding on oil palm yields, fine roots, or leaf area index. Also, we could not detect detrimental effects of the reduced fertilization and mechanical weeding on soil nutrients and functions (mineral nitrogen, bulk density, and litter decomposition), but water infiltration and base saturation tended to be higher under mechanical weeding, while soil moisture, and microbial biomass varied with treatment. Economic performance, measured as gross margins, was higher under reduced fertilization. There might be a delayed response of oil palm to the different management schemes applied, so results of future years may confirm whether this is a sustainable management strategy. Nevertheless, the initial effects of the experiment are encouraging to consider less intensive management practices as economically and ecologically viable options for oil palm plantations.
Aims Our goals were (1) to determine whether tree species diversity affects nutrient (N, P and K) cycling, and (2) to assess whether there is competition for these nutrients between microbial biomass ...and trees. Methods We measured nutrient resorption efficiency by trees, nutrient contents in leaf litterfall, decomposition rates of leaf litter, nutrient turnover in decomposing leaf litter, and plant-available nutrients in the soil in monospecies stands of beech, oak, hornbeam and lime and in mixed-species stands, each consisting of three of these species. Results Cycling of nutrients through leaf litter input and decomposition were influenced by the types of tree species and not simply by tree species diversity. Trees and microbial biomass were competing strongly for P, less for K and only marginally for N. Such competition was most pronounced in mono-species stands of beech and oak, which had low nutrient turnover in their slow decomposing leaf litter, and less in mono-species stands of hornbeam and lime, which had high nutrient turnover in their fast decomposing leaf litter. Conclusions The low soil P and K availability in beech stands, which limit the growth of beech at Hainich, Germany, were alleviated by mixing beech with hornbeam and lime. These species-specific effects on nutrient cycling and soil nutrient availability can aid forest management in improving productivity and soil fertility.
Although the canopy can play an important role in forest nutrient cycles, canopy‐based processes are often overlooked in studies on nutrient deposition. In areas of nitrogen (N) and phosphorus (P) ...deposition, canopy soils may retain a significant proportion of atmospheric inputs, and also receive indirect enrichment through root uptake followed by throughfall or recycling of plant litter in the canopy. We measured net and gross rates of N cycling in canopy soils of tropical montane forests along an elevation gradient and assessed indirect effects of elevated nutrient inputs to the forest floor. Net N cycling rates were measured using the buried bag method. Gross N cycling rates were measured using ¹⁵N pool dilution techniques. Measurements took place in the field, in the wet and dry season, using intact cores of canopy soil from three elevations (1000, 2000 and 3000 m). The forest floor had been fertilized biannually with moderate amounts of N and P for 4 years; treatments included control, N, P, and N + P. In control plots, gross rates of NH₄ ⁺ transformations decreased with increasing elevation; gross rates of NO₃ ⁻ transformations did not exhibit a clear elevation trend, but were significantly affected by season. Nutrient‐addition effects were different at each elevation, but combined N + P generally increased N cycling rates at all elevations. Results showed that canopy soils could be a significant N source for epiphytes as well as contributing up to 23% of total (canopy + forest floor) mineral N production in our forests. In contrast to theories that canopy soils are decoupled from nutrient cycling in forest floor soil, N cycling in our canopy soils was sensitive to slight changes in forest floor nutrient availability. Long‐term atmospheric N and P deposition may lead to increased N cycling, but also increased mineral N losses from the canopy soil system.
Tropical montane forests are commonly limited by N or co-limited by N and P. Projected increases in N deposition in tropical montane regions are thought to be insufficient for vegetation demand and ...are not therefore expected to affect soil N availability and N₂O emissions. We established a factorial N- and P-addition experiment (i.e., N, P, N + P, and control) across an elevation gradient of montane forests in Ecuador to test these hypotheses: (1) moderate rates of N and P additions are able to stimulate soil-N cycling rates and N₂O fluxes, and (2) the magnitude and timing of soil N₂O-flux responses depend on the initial nutrient status of the forest soils. Moderate rates of nutrients were added: 50 kg N ha⁻¹ year⁻¹ (in the form of urea) and 10 kg P ha⁻¹ year⁻¹ (in the form of NaH₂PO ₄ . 2H₂O) split in two equal applications. We tested the hypotheses by measuring changes in net rates of soil–N cycling and N₂O fluxes during the first 2 years (2008–2009) of nutrient manipulation in an old-growth premontane forest at 1,000 m, growing on a Cambisol soil with no organic layer, in an old-growth lower montane forest at 2,000 m, growing on a Cambisol soil with an organic layer, and an old-growth upper montane rainforest at 3,000 m, growing on a Histosol soil with a thick organic layer. Among the control plots, net nitrification rates were largest at the 1,000-m site whereas net nitrification was not detectable at the 2,000- and 3,000-m sites. The already large net nitrification at the 1,000-m site was not affected by nutrient additions, but net nitrification became detectable at the 2,000- and 3000-m sites after the second year of N and N + P additions. N₂O emissions increased rapidly following N and N + P additions at the 1,000-m site whereas only smaller increases occurred at the 2,000- and 3,000-m sites during the second year of N and N + P additions. Addition of P alone had no effect on net rates of soil N cycling and N₂O fluxes at any elevation. Our results showed that the initial soil N status, which may also be influenced by presence or absence of organic layer, soil moisture and temperature as encompassed by the elevation gradient, is a good indicator of how soil N cycling and N₂O fluxes may respond to future increases in nutrient additions.
Soil respiration is the second largest flux in the global carbon cycle, yet the underlying below-ground process, carbon dioxide (CO2 ) production, is not well understood because it can not be ...measured in the field. CO2 production has frequently been calculated from the vertical CO2 diffusive flux divergence, known as "soil-CO2 profile method". This relatively simple model requires knowledge of soil CO2 concentration profiles and soil diffusive properties. Application of the method for a tropical lowland forest soil in Panama gave inconsistent results when using diffusion coefficients (D) calculated based on relationships with soil porosity and moisture ("physically modeled" D). Our objective was to investigate whether these inconsistencies were related to (1) the applied interpolation and solution methods and/or (2) uncertainties in the physically modeled profile of D. First, we show that the calculated CO2 production strongly depends on the function used to interpolate between measured CO2 concentrations. Secondly, using an inverse analysis of the soil-CO2 profile method, we deduce which D would be required to explain the observed CO2 concentrations, assuming the model perception is valid. In the top soil, this inversely modeled D closely resembled the physically modeled D. In the deep soil, however, the inversely modeled D increased sharply while the physically modeled D did not. When imposing a constraint during the fit parameter optimization, a solution could be found where this deviation between the physically and inversely modeled D disappeared. A radon (Rn) mass balance model, in which diffusion was calculated based on the physically modeled or constrained inversely modeled D, simulated observed Rn profiles reasonably well. However, the CO2 concentrations which corresponded to the constrained inversely modeled D were too small compared to the measurements. We suggest that, in well-structured soils, a missing description of steady state CO2 exchange fluxes across water-filled pores causes the soil-CO2 profile method to fail. These fluxes are driven by the different diffusivities in inter- vs. intra-aggregate pores which create permanent CO2 gradients if separated by a "diffusive water barrier". These results corroborate other studies which have shown that the theory to treat gas diffusion as homogeneous process, a precondition for use of the soil-CO2 profile method, is inaccurate for pore networks which exhibit spatial separation between CO2 production and diffusion out of the soil.
Degraded lands are common in human-influenced tropical semiarid areas, and the potential for C sequestration through rehabilitation of these areas is substantial. In this study, we investigated ...changes in ecosystem C stocks (ECS) after establishing exclosures on degraded communal grazing lands, and identified easily measurable biophysical and management-related factors that can be used to predict ECS restoration in the highlands of Tigray, Ethiopia. We selected replicated (n = 3) 5-, 10-, 15-, and 20-yr-old exclosures and paired each exclosure with an adjacent communal grazing land. All exclosures displayed higher ECS than the communal grazing lands. Differences in ECS between exclosures and grazing lands varied between 29 (±4.9) and 61 (±6.7) Mg C ha−1 and increased with exclosure duration. In exclosures, much of the variability in ECS was explained by a combination of the following variables: precipitation, clay content, vegetation canopy cover, woody biomass, and exclosure duration (R 2 = 0.77–0.90). Precipitation and vegetation canopy cover also explained much of the variability of ECS in communal grazing lands (R 2 = 0.48–0.55). Our results help to establish baseline information for C sequestration projects and to predict the expected ecosystem C sequestration under exclosures. Expansion of exclosures would increase grazing pressure on the remaining communal grazing area. Therefore, the decision to establish additional exclosures should also include an economic analysis and an evaluation of the social consequences.
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