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Lianas are a prominent growth form in tropical forests, and there is compelling evidence that they are increasing in abundance throughout the Neotropics. While recent evidence shows that soil resources limit tree growth even in deep shade, the degree to which soil resources limit lianas in forest understories, where they coexist with trees for decades, remains unknown. Regardless, the physiological underpinnings of soil resource limitation in deeply shaded tropical habitats remain largely unexplored for either trees or lianas. Theory predicts that lianas should be more limited by soil resources than trees because they occupy the quick-return end of the "leaf economic spectrum," characterized by high rates of photosynthesis, high specific leaf area, short leaf life span, affinity to high-nutrient sites, and greater foliar nutrient concentrations. To address these issues, we asked whether soil resources (nitrogen, phosphorus, and potassium), alone or in combination, applied experimentally for more than a decade would cause significant changes in the morphology or physiology of tree and liana seedlings in a lowland tropical forest. We found evidence for the first time that phosphorus limits the photosynthetic performance of both trees and lianas in deeply shaded understory habitats. More importantly, lianas always showed significantly greater photosynthetic capacity, quenching, and saturating light levels compared to trees across all treatments. We found little evidence for nutrient x growth form interactions, indicating that lianas were not disproportionately favored in nutrient-rich habitats. Tree and liana seedlings differed markedly for six key morphological traits, demonstrating that architectural differences occurred very early in ontogeny prior to lianas finding a trellis (all seedlings were self-supporting). Overall, our results do not support nutrient loading as a mechanism of increasing liana abundance in the Neotropics. Rather, our finding that lianas always outperform trees, in terms of photosynthetic processes and under contrasting rates of resource supply of macronutrients, will allow lianas to increase in abundance if disturbance and tree turnover rates are increasing in Neotropical forests as has been suggested.

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.

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 x 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.

Over the past two decades, liana density and basal area have been increasing in many tropical forests, which has profound consequences for forest diversity and functioning. One hypothesis to explain increasing lianas is elevated nutrient deposition in tropical forests resulting from fossil fuels, agricultural fertilizer, and biomass burning. We tested this hypothesis by surveying all lianas ≥1 cm in diameter (n = 3,967) in 32 plots in a fully factorial nitrogen (N), phosphorus (P), and potassium (K) addition experiment in a mature tropical forest in central Panama. We conducted the nutrient‐addition experiment from 1998 until present and we first censused lianas in 2013 and then again in 2018. After 20 yr of nutrient addition (1998–2018), liana density, basal area, and rarefied species richness did not differ significantly among any of the nutrient‐addition and control treatments. Moreover, nutrient addition in the most recent 5 yr of the experiment did not affect liana relative growth, recruitment, or mortality rates. From 2013 until 2018, liana density, basal area, and species richness increased annually by 1.6%, 1.4%, and 2.4%, respectively. Nutrient addition did not influence these increases. Our findings indicate that nutrient deposition does not explain increasing lianas in this tropical forest. Instead, increases in tree mortality and disturbance, atmospheric carbon dioxide, drought frequency and severity, and hunting pressure may be more likely explanations for the increase in lianas in tropical forests. [ABSTRACT FROM AUTHOR]

Abstract: 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. [ABSTRACT FROM AUTHOR]

Questions remain as to which soil nutrients limit primary production in tropical forests. Phosphorus (P) has long been considered the primary limiting element in lowland forests, but recent evidence demonstrates substantial heterogeneity in response to nutrient addition, highlighting a need to understand and diagnose nutrient limitation across diverse forests. Fine-root characteristics including their abundance, functional traits, and mycorrhizal symbionts can be highly responsive to changes in soil nutrients and may help to diagnose nutrient limitation. Here, we document the response of fine roots to long-term nitrogen (N), P, and potassium (K) fertilization in a lowland forest in Panama. Because this experiment has demonstrated that N and K together limit tree growth and P limits fine litter production, we hypothesized that fine roots would also respond to nutrient addition. Specifically we hypothesized that N, P, and K addition would reduce the biomass, diameter, tissue density, and mycorrhizal colonization of fine roots, and increase nutrient concentration in root tissue. Most morphological root traits responded to the single addition of K and the paired addition of N and P, with the greatest response to all three nutrients combined. The addition of N, P, and K together reduced fine-root biomass, length, and tissue density, and increased specific root length, whereas root diameter remained unchanged. Nitrogen addition did not alter root N concentration, but P and K addition increased root P and K concentration, respectively. Mycorrhizal colonization of fine roots declined with N, increased with P, and was unresponsive to K addition. Although plant species composition remains unchanged after 14 years of fertilization, fine-root characteristics responded to N, P, and K addition, providing some of the strongest stand-level responses in this experiment. Multiple soil nutrients regulate fine-root abundance, morphological and chemical traits, and their association with mycorrhizal fungi in a species-rich lowland tropical forest. [ABSTRACT FROM AUTHOR]

Foliar nitrogen to phosphorus (N:P) ratios are widely used to indicate soil nutrient availability and limitation, but the foliar ratios of woody plants have proven more complicated to interpret than ratios from whole biomass of herbaceous species. This may be related to tissues in woody species acting as nutrient reservoirs during active growth, allowing maintenance of optimal N:P ratios in recently produced, fully expanded leaves (i.e., "new" leaves, the most commonly sampled tissue). Here we address the hypothesis that N:P ratios of newly expanded leaves are less sensitive indicators of soil nutrient availability than are other tissue types in woody plants. Seedlings of five naturally established tree species were harvested from plots receiving two years of fertilizer treatments in a lowland tropical forest in the Republic of Panama. Nutrient concentrations were determined in new leaves, old leaves, stems, and roots. For stems and roots, N:P ratios increased after N addition and decreased after P addition, and trends were consistent across all five species. Older leaves also showed strong responses to N and P addition, and trends were consistent for four of five species. In comparison, overall N:P ratio responses in new leaves were more variable across species. These results indicate that the N:P ratios of stems, roots, and older leaves are more responsive indicators of soil nutrient availability than are those of new leaves. Testing the generality of this result could improve the use of tissue nutrient ratios as indices of soil nutrient availability in woody plants [ABSTRACT FROM AUTHOR]