Aquilegia, Vol. 30 No. 2, April-May 2006: Newsletter of the Colorado Native Plant Society

https://epublications.regis.edu/aquilegia/1107/thumbnail.jp

Growth–Defense Trade-Offs Shape Population Genetic Composition in an Iconic Forest Tree Species

All organisms experience fundamental conflicts between divergent metabolic processes. In plants, a pivotal conflict occurs between allocation to growth, which accelerates resource acquisition, and to defense, which protects existing tissue against herbivory. Trade-offs between growth and defense traits are not universally observed, and a central prediction of plant evolutionary ecology is that context-dependence of these trade-offs contributes to the maintenance of intraspecific variation in defense [Züst and Agrawal, Annu. Rev. Plant Biol., 68, 513–534 (2017)]. This prediction has rarely been tested, however, and the evolutionary consequences of growth–defense trade-offs in different environments are poorly understood, especially in long-lived species [Cipollini et al., Annual Plant Reviews (Wiley, 2014), pp. 263–307]. Here we show that intraspecific trait trade-offs, even when fixed across divergent environments, interact with competition to drive natural selection of tree genotypes corresponding to their growth–defense phenotypes. Our results show that a functional trait trade-off, when coupled with environmental variation, causes real-time divergence in the genetic architecture of tree populations in an experimental setting. Specifically, competitive selection for faster growth resulted in dominance by fast-growing tree genotypes that were poorly defended against natural enemies. This outcome is a signature example of eco-evolutionary dynamics: Competitive interactions affected microevolutionary trajectories on a timescale relevant to subsequent ecological interactions [Brunner et al., Funct. Ecol. 33, 7–12 (2019)]. Eco-evolutionary drivers of tree growth and defense are thus critical to stand-level trait variation, which structures communities and ecosystems over expansive spatiotemporal scales

Sexual and genotypic variation in terpene quantitative and qualitative profiles in the dioecious shrub Baccharis salicifolia.

Terpenoids are secondary metabolites produced in most plant tissues and are often considered toxic or repellent to plant enemies. Previous work has typically reported on intra-specific variation in terpene profiles, but the effects of plant sex, an important axis of genetic variation, have been less studied for chemical defences in general, and terpenes in particular. In a prior study, we found strong genetic variation (but not sexual dimorphism) in terpene amounts in leaves of the dioecious shrub Baccharis salicifolia. Here we build on these findings and provide a more in-depth analysis of terpene chemistry on these same plants from an experiment consisting of a common garden with male (N = 19) and female (N = 20) genotypes sourced from a single population. Our goal in the present study was to investigate quantitative and qualitative differences in terpene profiles associated with plant sex and genotypic variation. For this, we quantified leaf mono- and sesquiterpene amount, richness, and diversity (quantitative profile), as well as the composition of compounds (qualitative profile). We found no evidence of sexual dimorphism in monoterpene or sesquiterpene profiles. We did, however, find significant genotypic variation in amount, diversity, and composition of monoterpenes, but no effects on sesquiterpenes. These findings indicated that genotypic variation in terpene profiles largely surpassed variation due to sexual dimorphism for the studied population of this species

Bacteria contribute to plant secondary compound degradation in a generalist herbivore system.

Herbivores must overcome a variety of plant defenses, including coping with plant secondary compounds (PSCs). To help detoxify these defensive chemicals, several insect herbivores are known to harbor gut microbiota with the metabolic capacity to degrade PSCs. Leaf-cutter ants are generalist herbivores, obtaining sustenance from specialized fungus gardens that act as external digestive systems and which degrade the diverse collection of plants foraged by the ants. There is in vitro evidence that certain PSCs harm Leucoagaricus gongylophorus, the fungal cultivar of leaf-cutter ants, suggesting a role for the Proteobacteria-dominant bacterial community present within fungus gardens. In this study, we investigated the ability of symbiotic bacteria present within fungus gardens of leaf-cutter ants to degrade PSCs. We cultured fungus garden bacteria, sequenced the genomes of 42 isolates, and identified genes involved in PSC degradation, including genes encoding cytochrome P450 enzymes and genes in geraniol, cumate, cinnamate, and alfa-pinene/limonene degradation pathways. Using metatranscriptomic analysis, we showed that some of these degradation genes are expressed in situ. Most of the bacterial isolates grew unhindered in the presence of PSCs and, using gas chromatography-mass spectrometry (GC-MS), we determined that isolates from the genera Bacillus, Burkholderia, Enterobacter, Klebsiella, and Pseudomonas degrade alfa-pinene, beta-caryophyllene, or linalool. Using a headspace sampler, we show that subcolonies of fungus gardens reduced alfa-pinene and linalool over a 36-h period, while L. gongylophorus strains alone reduced only linalool. Overall, our results reveal that the bacterial communities in fungus gardens play a pivotal role in alleviating the effect of PSCs on the leaf-cutter ant system.Great Lakes Bioenergy Research Center/[DE-SC0018409]/GLBRC/Estados UnidosGreat Lakes Bioenergy Research Center/[DE-FC02- 07ER64494]/GLBRC/Estados UnidosNational Institutes of Health/[U19 TW009872]/NIH/Estados UnidosNational Institutes of Health/[U19 AI142720]/NIH/Estados UnidosNational Science Foundation/[DEB-1927155]/NSF/Estados UnidosUniversidad de Costa Rica/[810-B0-501]/UCR/Costa RicaMinisterio de Ciencia, Tecnología y Telecomunicaciones/[FI-290-09]/MICITT/Costa RicaUCR::Vicerrectoría de Investigación::Unidades de Investigación::Ciencias Básicas::Centro de Investigación en Estructuras Microscópicas (CIEMIC)UCR::Vicerrectoría de Investigación::Unidades de Investigación::Ciencias Básicas::Centro de Investigación en Biología Celular y Molecular (CIBCM)UCR::Vicerrectoría de Docencia::Salud::Facultad de Medicina::Escuela de Medicin

Genetic Variation in Aspen Phytochemical Patterns Structures Windows of Opportunity for Gypsy Moth Larvae

Empirical studies indicate that host-tree bud break will likely advance faster than spring-folivore egg hatch in response to predicted increases in temperature. How these phenological shifts will affect herbivory will depend on temporal patterns of foliar traits that occur during leaf expansion, and their effects on folivore performance. Through fine-scale time series sampling of newly flushed trembling aspen (Populus tremuloides) foliage, we observed a previously unknown peak in phenolic glycoside concentrations that coincides with the emergence of sensitive neonates of gypsy moths and rapidly declines soon after bud break. The magnitude and duration of the initial post-bud break peak in phenolic glycosides varied substantially among genotypes. In contrast, foliar nitrogen concentrations declined at a more uniform rate among genotypes throughout leaf expansion. In addition, leaf toughness remained uniformly low throughout these periods of phytochemical change, and did not rise or vary substantially among genotypes until after anticipated windows of climate change-induced shifts between bud break and egg hatch had elapsed. Controlled manipulation of intervals between gypsy moth egg hatch and aspen bud break generated differences in larval performance among hatch cohorts and host genotypes that corresponded with changes in foliar phenolic glycoside and nitrogen concentrations. These findings indicate that the effects of climate change-induced phenological shifts on herbivory will differ among host plant genotypes, and that genetic variation in foliar chemical patterns will strongly influence this heterogeneity

To Compete or Defend: Linking Functional Trait Variation With Life-History Tradeoffs in a Foundation Tree Species

Although chemical deterrents to herbivory often exact costs in terms of plant growth, the manner in which those costs arise, and their physiological relationship to other functional traits, remain unclear. In the absence of appreciable herbivory, we examined interrelationships among chemical defense levels and other foliar functional traits (e.g., light-saturated photosynthesis, specific leaf area, nitrogen concentration) as co-determinants of tree growth and, by extension, competitive ability in high-density populations comprising 16 genotypes of Populus tremuloides. Across genotypes, concentrations of chemical defenses were not significantly related to other leaf functional traits, but levels of the salicinoid phenolic glycosides (SPGs) salicin, salicortin and tremulacin were each negatively correlated with relative mass growth (RMG) of aboveground woody tissue (P ≤ 0.001). RMG, in turn, underpinned 77% of the genotypic variation in relative height growth (our index of competitive ability). RMG was also positively related to light-saturated photosynthesis (P ≤ 0.001), which, together with the three SPGs, explained 86% of genotypic RMG variation (P ≤ 0.001). Moreover, results of a carbon balance simulation indicated that costs of resource allocation to SPGs, reaching nearly a third of annual crown photosynthesis, were likely mediated by substantial metabolic turnover, particularly for salicin. The lack of discernible links between foliar defense allocation and other (measured) functional traits, and the illustrated potential of metabolic turnover to reconcile influences of SPG allocation on RMG, shed additional light on fundamental physiological mechanisms underlying evolutionary tradeoffs between chemical defense investment and competitive ability in a foundation tree species