We maintained a factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment for 11 years in a humid lowland forest growing on a relatively fertile soil in Panama to evaluate ...potential nutrient limitation of tree growth rates, fine-litter production, and fine-root biomass. We replicated the eight factorial treatments four times using 32 plots of 40 ×× 40 m each. The addition of K was associated with significant decreases in stand-level fine-root biomass and, in a companion study of seedlings, decreases in allocation to roots and increases in height growth rates. The addition of K and N together was associated with significant increases in growth rates of saplings and poles (1-–10 cm in diameter at breast height) and a further marginally significant decrease in stand-level fine-root biomass. The addition of P was associated with a marginally significant (
P
== 0.058) increase in fine-litter production that was consistent across all litter fractions. Our experiment provides evidence that N, P, and K all limit forest plants growing on a relatively fertile soil in the lowland tropics, with the strongest evidence for limitation by K among seedlings, saplings, and poles.
We present a meta-analysis of plant responses to fertilization experiments conducted in lowland, species-rich, tropical forests. We also update a key result and present the first species-level ...analyses of tree growth rates for a 15-yr factorial nitrogen (N), phosphorus (P), and potassium (K) experiment conducted in central Panama. The update concerns community-level tree growth rates, which responded significantly to the addition of N and K together after 10 yr of fertilization but not after 15 yr. Our experimental soils are infertile for the region, and species whose regional distributions are strongly associated with low soil P availability dominate the local tree flora. Under these circumstances, we expect muted responses to fertilization, and we predicted species associated with low-P soils would respond most slowly. The data did not support this prediction, species-level tree growth responses to P addition were unrelated to species-level soil P associations. The meta-analysis demonstrated that nutrient limitation is widespread in lowland tropical forests and evaluated two directional hypotheses concerning plant responses to N addition and to P addition. The meta-analysis supported the hypothesis that tree (or biomass) growth rate responses to fertilization are weaker in old growth forests and stronger in secondary forests, where rapid biomass accumulation provides a nutrient sink. The meta-analysis found no support for the long-standing hypothesis that plant responses are stronger for P addition and weaker for N addition. We do not advocate discarding the latter hypothesis. There are only 14 fertilization experiments from lowland, species-rich, tropical forests, 13 of the 14 experiments added nutrients for five or fewer years, and responses vary widely among experiments. Potential fertilization responses should be muted when the species present are well adapted to nutrient-poor soils, as is the case in our experiment, and when pest pressure increases with fertilization, as it does in our experiment. The statistical power and especially the duration of fertilization experiments conducted in old growth, tropical forests might be insufficient to detect the slow, modest growth responses that are to be expected.
1. Nutrients are a critical resource for plant growth, but the elements limiting growth in tropical forests have rarely been determined. 2. We investigated the influence of nitrogen (N), phosphorus ...(P), potassium (K) and micronutrients on seedling biomass and nutrient allocation in a factorial nutrient fertilization experiment in lowland tropical forest at the Barro Colorado Nature Monument, Panama. We also measured 8 years of herbivory and growth for 1800 seedlings. We sought to determine the identity of limiting elements and possible nutrient interactions. 3. The five study species were Alseis blackiana, Desmopsis panamensis, Heisteria concinna, Sorocea affinis and Tetragastris panamensis. Plants grew in deeply shaded understorey with a mean canopy openness of 4.9% (±0.7%; 1 SE). 4. Tissue N concentration increased by 11% with N addition. Tissue P concentration increased by 16% with P addition. Tissue K increased by 4% with K addition. K addition reduced root‐to‐shoot biomass ratio. There was no significant effect of fertilization on specific leaf area or leaf area ratio. 5. The proportion of leaves damaged and the mean level of damage by herbivory increased with P and K addition and showed a significant P × K interaction. 6.7ensp;Across all species and years, relative growth rate of height increased with K addition and with N and P in combination. Relative growth rate of leaf count trended 8.5% higher with K addition (P = 0.076). 7. We also added micronutrients in a parallel experiment. There was no effect of micronutrient addition on any seedling parameter. 8. Synthesis. K addition affected seedlings by enhancing tissue nutrient concentration, increasing herbivory, reducing root‐to‐shoot biomass ratio and increasing height growth. Additional effects of N or P on tissue chemistry, herbivory and growth offer support for the multiple limiting resources hypothesis. Our results suggest that seedling growth is limited by nutrients, especially K, even under highly shaded conditions in this lowland tropical forest.
Abstract
Northern peatlands are experiencing more frequent and severe fire events as a result of changing climate conditions. Recent studies show that such a fire-regime change imposes a direct ...climate-warming impact by emitting large amounts of carbon into the atmosphere. However, the fires also convert parts of the burnt biomass into pyrogenic carbon. Here, we show a potential climate-cooling impact induced by fire-derived pyrogenic carbon in laboratory incubations. We found that the accumulation of pyrogenic carbon reduced post-fire methane production from warm (32 °C) incubated peatland soils by 13–24%. The redox-cycling, capacitive, and conductive electron transfer mechanisms in pyrogenic carbon functioned as an electron snorkel, which facilitated extracellular electron transfer and stimulated soil alternative microbial respiration to suppress methane production. Our results highlight an important, but overlooked, function of pyrogenic carbon in neutralizing forest fire emissions and call for its consideration in the global carbon budget estimation.
To explore the importance of 12 elements in litter production and decomposition, we fertilized 36 1600 m²-plots with combinations of N, P, K, or micronutrients (i.e. B, Ca, Cu, Fe, Mg, Mn, Mo, S, Zn) ...for 6 years in a lowland Panamanian forest. The 90% of litter falling as leaves and twigs failed to increase with fertilization, but reproductive litter (fruits and flowers) increased by 43% with N. K enhanced cellulose decomposition; one or more micronutrients enhanced leaf-litter decomposition; P enhanced both. Our results suggest tropical forests are a non-Liebig world of multiple nutrient limitations, with at least four elements shaping rates of litterfall and decomposition. Multiple metallomic enzymes and cofactors likely create gradients in the break down of leaf litter. Selection favours individuals that make more propagules, and even in an N-rich forest, N is a non-substitutable resource for reproduction.
One of the principal inputs of organic matter to forest soils is turnover of tree fine roots, but the process of decomposition of fine root litter and its conversion into stable soil organic matter ...(SOM) has received limited study. We labeled fine roots of sugar maple (
Acer saccharum
Marsh.) with
13
C and traced the label for 7 years into four contrasting soils to improve understanding of this process. After 7 years we recovered an average of 8.9% of the
13
C label, with about two-thirds recovered as coarse particulate organic matter and one-third in microaggregates and on silt and clay particles. No differences in
13
C recovery were detected between 1–2 and 3–4 order fine roots. Most of the
13
C in microaggregates (53–250 µm, 58%) was occluded within macroaggregates, and the recovery in this fraction increased significantly from year 2 to 7, illustrating the role of fine root detritus in the formation of microaggregates. This process was most pronounced in the A horizon of a higher pH soil (pH = 5.5) with high iron oxide content. Conversely, the lowest
13
C recovery in this fraction was observed in the A horizon of an acidic, fine-textured Inceptisol (Cambisol—World Reference Base). We estimate that annual input into relatively stable fractions of SOM represents about 14% of the total annual accumulation in these fractions; thus, our results support recent evidence that fine root litter is only a moderate contributor to stable SOM in acid temperate forest soils.
The importance of niche vs. neutral assembly mechanisms in structuring tropical tree communities remains an important unsettled question in community ecology Bell G (2005) Ecology 86:1757-1770. There ...is ample evidence that species distributions are determined by soils and habitat factors at landscape (<10⁴ km²) and regional scales. At local scales (<1 km²), however, habitat factors and species distributions show comparable spatial aggregation, making it difficult to disentangle the importance of niche and dispersal processes. In this article, we test soil resource-based niche assembly at a local scale, using species and soil nutrient distributions obtained at high spatial resolution in three diverse neotropical forest plots in Colombia (La Planada), Ecuador (Yasuni), and Panama (Barro Colorado Island). Using spatial distribution maps of >0.5 million individual trees of 1,400 species and 10 essential plant nutrients, we used Monte Carlo simulations of species distributions to test plant-soil associations against null expectations based on dispersal assembly. We found that the spatial distributions of 36-51% of tree species at these sites show strong associations to soil nutrient distributions. Neutral dispersal assembly cannot account for these plant-soil associations or the observed niche breadths of these species. These results indicate that belowground resource availability plays an important role in the assembly of tropical tree communities at local scales and provide the basis for future investigations on the mechanisms of resource competition among tropical tree species.
Both habitat filtering and dispersal limitation influence the compositional structure of forest communities, but previous studies examining the relative contributions of these processes with ...variation partitioning have primarily used topography to represent the influence of the environment. Here, we bring together data on both topography and soil resource variation within eight large (24–50 ha) tropical forest plots, and use variation partitioning to decompose community compositional variation into fractions explained by spatial, soil resource and topographic variables. Both soil resources and topography account for significant and approximately equal variation in tree community composition (9–34% and 5–29%, respectively), and all environmental variables together explain 13–39% of compositional variation within a plot. A large fraction of variation (19–37%) was spatially structured, yet unexplained by the environment, suggesting an important role for dispersal processes and unmeasured environmental variables. For the majority of sites, adding soil resource variables to topography nearly doubled the inferred role of habitat filtering, accounting for variation in compositional structure that would previously have been attributable to dispersal. Our results, illustrated using a new graphical depiction of community structure within these plots, demonstrate the importance of small-scale environmental variation in shaping local community structure in diverse tropical forests around the globe.
Summary
Model denitrifiers convert NO3− to N2, but it appears that a significant fraction of natural populations are truncated, conducting only one or two steps of the pathway. To better understand ...the diversity of partial denitrifiers in soil and whether discrepancies arise between the presence of known N‐oxide reductase genes and phenotypic features, bacteria able to reduce
NO3− to
NO2− were isolated from soil, N‐oxide gas products were measured for eight isolates, and six were genome sequenced. Gas phase analyses revealed that two were complete denitrifiers, which genome sequencing corroborated. The remaining six accumulated NO and N2O to varying degrees and genome sequencing of four indicated that two isolates held genes encoding nitrate reductase as the only dissimilatory N‐oxide reductase, one contained genes for both nitrate and nitric oxide reductase, and one had nitrate and nitrite reductase. The results demonstrated that N‐oxide production was not always predicted by the genetic potential and suggested that partial denitrifiers could be readily isolated among soil bacteria. This supported the hypothesis that each N‐oxide reductase could provide a selectable benefit on its own, and therefore, reduction of nitrate to dinitrogen may not be obligatorily linked to complete denitrifiers but instead a consequence of a functionally diverse community.
Exotic earthworms can profoundly alter soil carbon (C) and nitrogen (N) dynamics in northern temperate forests, but the mechanisms explaining these responses are not well understood. We compared the ...soil microbial community (SMC) composition (measured as PLFAs) and enzyme activity between paired earthworm-invaded and earthworm-free plots in northern hardwood forests of New York, USA. We hypothesized that differences in SMCs and enzyme activity between plots would correspond with differences in soil C content and C:N ratios. Relative abundance of several bacterial (mostly gram-positive) PLFAs was higher and that of two fungal PLFAs was lower in earthworm compared to reference plots, largely because of earthworm incorporation of the organic horizon into mineral soil. In surface mineral soil earthworms increased arbuscular mycorrhizal fungi (AMF) and gram-positive bacterial PLFAs, and decreased fungal (mostly saprotrophic) and several bacterial (gram-negative and non-specific) PLFAs. Earthworms also increased the activities of cellulolytic relative to lignolytic enzymes in surface mineral soil, and the relationships between enzyme activities and components of the SMC suggest a substrate-mediated effect on the SMC and its metabolism of C. A highly significant relationship between components of the SMC and soil C:N also suggests that earthworms reduce soil C:N through functional and compositional shifts in the SMC. Finally, changes in AMF abundances were linked to phosphatase activity, suggesting that earthworms do not necessarily inhibit P-acquisition by AMF-associated plants in our study system. We conclude that the combined influence of earthworm-related changes in physical structure, accessibility and chemistry of organic matter, and relative abundance of certain groups of fungi and bacteria promote C metabolism, in particular by increasing the activities of cellulolytic vs. lignolytic enzymes.
•Earthworms increased abundance of many bacterial relative to most fungal indicators.•Abundance of arbuscular mycorrhizal fungi increased by earthworms.•Earthworms increased activities of cellulolytic compared to lignolytic enzymes.•Changes in enzyme activities contribute to more complete carbon metabolism.•Soil microbial community composition correlated with enzyme activity and soil C:N.
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