Plant responses to fertilization experiments in lowland, species-rich, tropical forests.

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.

Temperature response of CO2 exchange in three tropical tree species.

Tropical forests play a critical role in the global carbon cycle, but our limited understanding of the physiological sensitivity of tropical forest trees to environmental factors complicates predictions of tropical carbon fluxes in a changing climate. We determined the short-term temperature response of leaf photosynthesis and respiration of seedlings of three tropical tree species from Panama. For one of the species net CO2 exchange was also measured in situ. Dark respiration of all species increased linearly - not exponentially - over a ~30°C temperature range. The early-successional species Ficus insipida Willd. and Ochroma pyramidale (Cav. ex Lam.) Urb. had higher temperature optima for photosynthesis (Topt) and higher photosynthesis rates at Topt than the late-successional species Calophyllum longifolium Willd. The decrease in photosynthesis above Topt could be assigned, in part, to observed temperature-stimulated photorespiration and decreasing stomatal conductance (gS), with unmeasured processes such as respiration in the light, Rubisco deactivation, and changing membrane properties probably playing important additional roles, particularly at very high temperatures. As temperature increased above Topt, gS of laboratory-measured leaves first decreased, followed by an increase at temperatures >40-45°C. In contrast, gS of canopy leaves of F. insipida in the field continued to decrease with increasing temperature, causing complete suppression of photosynthesis at ~45°C, whereas photosynthesis in the laboratory did not reach zero until leaf temperature was ~50°C. Models parameterised with laboratory-derived data should be validated against field observations when they are used to predict tropical forest carbon fluxes.

Pervasive effects of a dominant foliar endophytic fungus on host genetic and phenotypic expression in a tropical tree.

It is increasingly recognized that macro-organisms (corals, insects, plants, vertebrates) consist of both host tissues and multiple microbial symbionts that play essential roles in their host's ecological and evolutionary success. Consequently, identifying benefits and costs of symbioses, as well as mechanisms underlying them are research priorities. All plants surveyed under natural conditions harbor foliar endophytic fungi (FEF) in their leaf tissues, often at high densities. Despite producing no visible effects on their hosts, experiments have nonetheless shown that FEF reduce pathogen and herbivore damage. Here, combining results from three genomic, and two physiological experiments, we demonstrate pervasive genetic and phenotypic effects of the apparently asymptomatic endophytes on their hosts. Specifically, inoculation of endophyte-free (E-) Theobroma cacao leaves with Colletotrichum tropicale (E+), the dominant FEF species in healthy T. cacao, induces consistent changes in the expression of hundreds of host genes, including many with known defensive functions. Further, E+ plants exhibited increased lignin and cellulose content, reduced maximum rates of photosynthesis (Amax), and enrichment of nitrogen-15 and carbon-13 isotopes. These phenotypic changes observed in E+ plants correspond to changes in expression of specific functional genes in related pathways. Moreover, a cacao gene (Tc00g04254) highly up-regulated by C. tropicale also confers resistance to pathogen damage in the absence of endophytes or their products in host tissues. Thus, the benefits of increased pathogen resistance in E+ plants are derived in part from up-regulation of intrinsic host defense responses, and appear to be offset by potential costs including reduced photosynthesis, altered host nitrogen metabolism, and endophyte heterotrophy of host tissues. Similar effects are likely in most plant-endophyte interactions, and should be recognized in the design and interpretation of genetic and phenotypic studies of plants.

Potassium, phosphorus, or nitrogen limit root allocation, tree growth, or litter production in a 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.

Multiple nutrients limit litterfall and decomposition in a tropical forest.

To explore the importance of 12 elements in litter production and decomposition, we fertilized 36 1600 m(2)-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.