The chemical ecology of tropical forest diversity: Environmental variation, chemical similarity, herbivory, and richness.

Species richness in tropical forests is correlated with other dimensions of diversity, including the diversity of plant-herbivore interactions and the phytochemical diversity that influences those interactions. Understanding the complexity of plant chemistry and the importance of phytochemical diversity for plant-insect interactions and overall forest richness has been enhanced significantly by the application of metabolomics to natural systems. The present work used proton nuclear magnetic resonance spectroscopy (1 H-NMR) profiling of crude leaf extracts to study phytochemical similarity and diversity among Piper plants growing naturally in the Atlantic Rainforest of Brazil. Spectral profile similarity and chemical diversity were quantified to examine the relationship between metrics of phytochemical diversity, specialist and generalist herbivory, and understory plant richness. Herbivory increased with understory species richness, while generalist herbivory increased and specialist herbivory decreased with the diversity of Piper leaf material available. Specialist herbivory increased when conspecific host plants were more spectroscopically dissimilar. Spectral similarity was lower among individuals of common species, and they were also more spectrally diverse, indicating phytochemical diversity is beneficial to plants. Canopy openness and soil nutrients also influenced chemistry and herbivory. The complex relationships uncovered in this study add information to our growing understanding of the importance of phytochemical diversity for plant-insect interactions and tropical plant species richness.

Terpenoid profiles of resin in the genus Dracaena are species specific.

Dragon's blood is the colloquial name for the red resin produced by tree species in the genus Dracaena (Asparagaceae), and the resin is directly involved in plant defensive mechanisms against pathogen and herbivore attack. It is also widely used in traditional folk medicine due to its antiviral, antimicrobial and antitumor activities. In the present work, a method using solid phase microextraction combined with two-dimensional gas chromatography with time-of-flight mass spectrometric detection was developed for the analysis of resin from five Dracaena species, namely Dracaena cinnabari Balf. f., D. serrulata Baker, D. ombet Heuglin ex Kotschy & Peyr., D. draco subsp. draco, and D. draco subsp. ajgal. Twenty terpenoid components in the resins of the five species were identified after comparative study of the volatile metabolite profiles. Monoterpenes were found to be species specific, and the observed differences might be further investigated as a possible means of identifying chemotaxonomic markers. In addition, for the first time, we describe the terpenoid volatile profiles of D. ombet and D. serrulata resins.

Early recruitment responses to interactions between frequent fires, nutrients, and herbivory in the southern Amazon.

Understanding tropical forest diversity is a long-standing challenge in ecology. With global change, it has become increasingly important to understand how anthropogenic and natural factors interact to determine diversity. Anthropogenic increases in fire frequency are among the global change variables affecting forest diversity and functioning, and seasonally dry forest of the southern Amazon is among the ecosystems most affected by such pressures. Studying how fire will impact forests in this region is therefore important for understanding ecosystem functioning and for designing effective conservation action. We report the results of an experiment in which we manipulated fire, nutrient availability, and herbivory. We measured the effects of these interacting factors on the regenerative capacity of the ecotone between humid Amazon forest and Brazilian savanna. Regeneration density, diversity, and community composition were severely altered by fire. Additions of P and N + P reduced losses of density and richness in the first year post-fire. Herbivory was most important just after germination. Diversity was positively correlated with herbivory in unburned forest, likely because fire reduced the number of reproductive individuals. This contrasts with earlier results from the same study system in which herbivory was related to increased diversity after fire. We documented a significant effect of fire frequency; diversity in triennially burned forest was more similar to that in unburned than in annually burned forest, and the community composition of triennially burned forest was intermediate between unburned and annually burned areas. Preventing frequent fires will therefore help reduce losses in diversity in the southern Amazon's matrix of human-altered landscapes.

An optimal defense strategy for phenolic glycoside production in Populus trichocarpa--isotope labeling demonstrates secondary metabolite production in growing leaves.

Large amounts of carbon are required for plant growth, but young, growing tissues often also have high concentrations of defensive secondary metabolites. Plants' capacity to allocate resources to growth and defense is addressed by the growth-differentiation balance hypothesis and the optimal defense hypothesis, which make contrasting predictions. Isotope labeling can demonstrate whether defense compounds are synthesized from stored or newly fixed carbon, allowing a detailed examination of these hypotheses. Populus trichocarpa saplings were pulse-labeled with 13CO2 at the beginning and end of a growing season, and the 13C signatures of phenolic glycosides (salicinoids), sugars, bulk tissue, and respired CO2 were traced over time. Half of the saplings were also subjected to mechanical damage. Populus trichocarpa followed an optimal defense strategy, investing 13C in salicinoids in expanding leaves directly after labeling. Salicinoids turned over quickly, and their production continued throughout the season. Salicin was induced by early-season damage, further demonstrating optimal defense. Salicinoids appear to be of great value to P. trichocarpa, as they command new C both early and late in the growing season, but their fitness benefits require further study. Export of salicinoids between tissues and biochemical pathways enabling induction also needs research. Nonetheless, the investigation of defense production afforded by isotope labeling lends new insights into plants' ability to grow and defend simultaneously.

Ontogenetic differences of herbivory on woody and herbaceous plants: a meta-analysis demonstrating unique effects of herbivory on the young and the old, the slow and the fast.

The effect of herbivory on plant performance is the subject of a large number of ecological studies, and plant responses to herbivory range from reduced reproduction to overcompensation. Because plant defenses, stored resources, and allocation demands change throughout a plant's lifetime, it can be hypothesized the effects of herbivory also vary with development. The present work extends previous analyses to incorporate hundreds of studies in a new meta-analysis addressing this topic. Herbivores had an overall negative effect on plant growth and reproduction, and, in contrast to a previous meta-analysis, this work shows the timing of herbivory is relevant. Differences in the effects of herbivory between life stages existed for woody plant reproduction and perennial herb growth. In addition, tree and shrub growth was reduced by herbivore damage at early ontogenetic stages, and perennial herb reproduction was limited by adult stage herbivory. These results partially support the continuum of an ontogenetic response model. Finally, consideration of this synthesis in conjunction with other work led to the conclusion that different plant groups optimize their defense investments in unique ways. Slow-growing plants may strongly chemically defend young tissues, supporting the plant-age hypothesis, because early herbivory is detrimental to growth. Faster-growing herbs may invest more in antiherbivore defense when they are older, supporting the growth-differentiation balance hypothesis, because later herbivory limits their reproduction.

Interactions between repeated fire, nutrients, and insect herbivores affect the recovery of diversity in the southern Amazon.

Surface fires burn extensive areas of tropical forests each year, altering resource availability, biotic interactions, and, ultimately, plant diversity. In transitional forest between the Brazilian cerrado (savanna) and high stature Amazon forest, we took advantage of a long-term fire experiment to establish a factorial study of the interactions between fire, nutrient availability, and herbivory on early plant regeneration. Overall, five annual burns reduced the number and diversity of regenerating stems. Community composition changed substantially after repeated fires, and species common in the cerrado became more abundant. The number of recruits and their diversity were reduced in the burned area, but burned plots closed to herbivores with nitrogen additions had a 14 % increase in recruitment. Diversity of recruits also increased up to 50 % in burned plots when nitrogen was added. Phosphorus additions were related to an increase in species evenness in burned plots open to herbivores. Herbivory reduced seedling survival overall and increased diversity in burned plots when nutrients were added. This last result supports our hypothesis that positive relationships between herbivore presence and diversity would be strongest in treatments that favor herbivory--in this case herbivory was higher in burned plots which were initially lower in diversity. Regenerating seedlings in less diverse plots were likely more apparent to herbivores, enabling increased herbivory and a stronger signal of negative density dependence. In contrast, herbivores generally decreased diversity in more species rich unburned plots. Although this study documents complex interactions between repeated burns, nutrients, and herbivory, it is clear that fire initiates a shift in the factors that are most important in determining the diversity and number of recruits. This change may have long-lasting effects as the forest progresses through succession.

Costs of defense and a test of the carbon-nutrient balance and growth-differentiation balance hypotheses for two co-occurring classes of plant defense.

One of the goals of chemical ecology is to assess costs of plant defenses. Intraspecific trade-offs between growth and defense are traditionally viewed in the context of the carbon-nutrient balance hypothesis (CNBH) and the growth-differentiation balance hypothesis (GDBH). Broadly, these hypotheses suggest that growth is limited by deficiencies in carbon or nitrogen while rates of photosynthesis remain unchanged, and the subsequent reduced growth results in the more abundant resource being invested in increased defense (mass-balance based allocation). The GDBH further predicts trade-offs in growth and defense should only be observed when resources are abundant. Most support for these hypotheses comes from work with phenolics. We examined trade-offs related to production of two classes of defenses, saponins (triterpenoids) and flavans (phenolics), in Pentaclethra macroloba (Fabaceae), an abundant tree in Costa Rican wet forests. We quantified physiological costs of plant defenses by measuring photosynthetic parameters (which are often assumed to be stable) in addition to biomass. Pentaclethra macroloba were grown in full sunlight or shade under three levels of nitrogen alone or with conspecific neighbors that could potentially alter nutrient availability via competition or facilitation. Biomass and photosynthesis were not affected by nitrogen or competition for seedlings in full sunlight, but they responded positively to nitrogen in shade-grown plants. The trade-off predicted by the GDBH between growth and metabolite production was only present between flavans and biomass in sun-grown plants (abundant resource conditions). Support was also only partial for the CNBH as flavans declined with nitrogen but saponins increased. This suggests saponin production should be considered in terms of detailed biosynthetic pathway models while phenolic production fits mass-balance based allocation models (such as the CNBH). Contrary to expectations based on the two defense hypotheses, trade-offs were found between defenses and photosynthesis, indicating that studies of plant defenses should include direct measures of physiological responses.