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As a strategic cost-saving alternative to constitutive resistance, induction of resistance against herbivores in plants can be especially beneficial when enemies are scarce or variable in abundance. Although probably describing the two ends of a continuum, constitutive and induced resistance strategies have long been observed to trade off within species. Examining these traits among populations along a successional gradient can help explain how temporally variable environments can maintain genetic variation and how ecosystem processes are affected by shifting plant resistance trait expression over time. Here we leverage large experimental plots that represent a chronosequence of succession up to 15 yr in combination with common garden experiments to examine changes in the selective environment and genetic differences in tall goldenrod’s (Solidago altissima) constitutive and induced resistance. We show that resistance against a specialist herbivore Trirhabda virgata was inducible in the plants originating from midsuccession, which coincides with the largest loads of herbivores. The flavonoid compound content of the leaves varied with successional stage of the population of origin, which is indicative of constitutive differences in secondary metabolite production. Finally, there was a clear trade-off between constitutive and induced resistance. Our study indicates that selection for resistance traits within a population can be highly variable over time and likely result in genetically determined shifts of resistance strategies over relatively short time periods via genotype sorting.

Understanding the evolutionary reasons for patterns of chemical defense in plants is an ongoing theoretical and empirical challenge. The goal is to develop a model that can reliably predict how defenses are distributed within the plant over space and time. This is difficult given that evolutionary, ecological, and physiological processes and tradeoffs can operate over different spatial and temporal scales. We evaluated the major predictions of two leading defense theories, the growth‐differentiation balance hypothesis (GDBH) and optimal defense theory (ODT). To achieve this, enemies, fitness components, terpenoids, and protease inhibitors were measured in Solidago altissima and used to construct conventional univariate and structural equation models (SEMs). Leaf‐tissue value indices extracted from an SEM revealed a strong correlation between tissue value and terpenoid defense that supports ODT. A tradeoff between serine protease inhibition and growth as well as an indirect tradeoff between growth and terpenoids manifested through galling insects supported the GDBH. Interestingly, there was a strong direct effect of terpenoids on rhizome mass, suggesting service to both storage and defense. The results support established theories but unknown genotypic traits explained much of the variation in defense, confirming the need to integrate emerging theories such as pollination constraints, defense syndromes, tolerance, mutualisms, and facilitation.

Plant-induced responses to multiple herbivores can mediate ecological interactions among herbivore species, thereby influencing herbivore community composition in nature. Several studies have indicated high specificity of induced responses to different herbivore species. In addition, there may be genetic variation for plant response specificity that can have significant ecological implications, by altering the competitive strength and hierarchical relationships among interacting herbivore species. However, few studies have examined whether plant populations harbor genetic variation for induction specificity. Using three distinct genotypes of Solidago altissima plants, we examined whether specialist herbivore species Dichomeris leuconotella, Microrhopala vittata, and Trirhabda virgata elicit specific induction responses from plants (specificity of elicitation), and whether induction differentially affects these herbivore species (specificity of effect). Results from bioassays and secondary metabolite analyses suggest that there is specificity of both elicitation and effect in the induced responses: D. leuconotella and M. vittata preferred and performed better on leaves damaged by conspecifics than heterospecifics, and induced qualitatively different secondary metabolite profiles. In contrast, T. virgata equally avoided but physiologically tolerated all types of damage. These patterns of specificity suggest that plant-induced responses mediate asymmetric competitive interactions between herbivore species, which potentially intensifies inter-specific relative to intra-specific competition. Plant genotypes widely differed in overall susceptibility to the herbivores and secondary metabolite production, yet we found no genotype-by-treatment interactions in insect performance, preference and plant secondary metabolite production. This lack of genetic variation for induction specificity suggests that competitive interactions between herbivore species on S. altissima are homogeneous across plant genotypes.

There is growing realization that intraspecific polyploidy in plants has important implications for the evolution of plant diversity and for plant and plant–herbivore community dynamics. Nonetheless, the rate at which polyploid cytotypes arise and their fate in local mixed-cytotype populations are not well understood. Although within- and especially among-population cytotype variation has been documented in many plants, particularly at the largest (continental) spatial scales, local and regional distributions of cytotypes have been well characterized only for a handful of species. Furthermore, few if any studies have addressed both local and regional patterns in the same plant species with sufficient power to detect cytotype variation on both scales. We assessed the co-occurrence and frequency distributions of diploid, tetraploid, and hexaploid cytotypes of Solidago altissima (Asteraceae) at local and regional spatial scales, sampling across a zone of presumed ploidy-form overlap in the midwestern United States. Determining ploidy by flow cytometry, we found many local populations containing two or three cytotypes, with the relative frequencies of cytotypes varying dramatically even among neighboring populations. AFLP marker data suggest that polyploid cytotypes of S. altissima likely have multiple origins from different diploid lineages. Our results are consistent with recent perpectives on polyploidization as an evolutionarily dynamic phenomenon with frequent multiple origins of higher-ploidy cytotypes.

Plant populations frequently maintain submaximal levels of resistance to natural enemies, even in the presence of substantial genetic variation for resistance. Identifying constraints on the evolution of increased resistance has been a major goal of researchers of plant–herbivore interactions. In a glasshouse study, we measured relative costs and benefits of resistance of tall goldenrod (Solidago altissima) to the gall-inducing tephritid Eurosta solidaginis. We exposed multiple ramets of 26 goldenrod genets to nutrient or shade stress and to oviposition by E. solidaginis. The presence of a gall cost a ramet an average of 1743 seeds, but the cost differed 10-fold across environments. Plant genets varied widely for an induced ‘hypersensitive’ response in which meristem cells become necrotic and kill E. solidaginis hatchlings before gall induction. There was no evidence that this highly effective resistance trait carried an allocation cost. However, the response carried a risk of autotoxicity, as necrosis killed the apex of 37% of the ungalled ramets. On average, a damaged apex cost each ramet 5015 seeds. Autotoxicity may constrain the resistance of S. altissima to an intermediate level, and variation in environmental conditions may alter the relative costs and benefits of resistance and tolerance, thus maintaining genetic variation within goldenrod populations.

Background: In the emerging field of community and ecosystem genetics, genetic variation and diversity in dominant plant species have been shown to play fundamental roles in maintaining biodiversity and ecosystem function. However, the importance of intraspecific genetic variation and diversity to floral abundance and pollinator visitation has received little attention. Methodology/Principal Findings: Using an experimental common garden that manipulated genotypic diversity (the number of distinct genotypes per plot) of Solidago altissima, we document that genotypic diversity of a dominant plant can indirectly influence flower visitor abundance. Across two years, we found that 1) plant genotype explained 45% and 92% of the variation in flower visitor abundance in 2007 and 2008, respectively; and 2) plant genotypic diversity had a positive and non-additive effect on floral abundance and the abundance of flower visitors, as plots established with multiple genotypes produced 25% more flowers and received 45% more flower visits than would be expected under an additive model. Conclusions/Significance: These results provide evidence that declines in genotypic diversity may be an important but little considered factor for understanding plant-pollinator dynamics, with implications for the global decline in pollinators due to reduced plant diversity in both agricultural and natural ecosystems.

Ecosystem engineers affect ecological communities by physically modifying the environment. Understanding the factors determining the distribution of engineers offers a powerful predictive tool for community ecology. In this study, we examine whether the goldenrod bunch gall midge (Rhopalomyia solidaginis) functions as an ecosystem engineer in an old-field ecosystem by altering the composition of arthropod species associated with a dominant host plant, Solidago altissima. We also examine the suite of factors that could affect the distribution and abundance of this ecosystem engineer. The presence of bunch galls increased species richness and altered the structure of associated arthropod communities. The best predictors of gall abundance were host-plant genotype and plot-level genotypic diversity. We found positive, nonadditive effects of genotypic diversity on gall abundance. Our results indicate that incorporating a genetic component in studies of ecosystem engineers can help predict their distribution and abundance, and ultimately their effects on biodiversity.

1. Ecological and physiological costs of resistance to herbivory are core concepts in the study of plant–insect interactions, but identifying them remains challenging. These costs are most obvious when trade‐offs in resource allocation occur between different growth and resistance traits. 2. We used plant genotypes collected from long‐term herbivore exclusion plots and from plots with natural herbivory. We evaluated putative trade‐offs between resistance to two different herbivore species (the larvae of the beetle Trirhabda virgata and the moth Spodoptera exigua), and between resistance and growth as a potential outcome of differential natural selection. 3. We hypothesized that long‐term exclusion of herbivores would result in differential natural selection on plant resistance traits that are expressed by marked differences in mean resistance between plants from populations with and without herbivores. The results supported this hypothesis. Genotypes from herbivore exclusion plots were less resistant to the more common herbivore in the system, T. virgata, than genotypes from control plots. In contrast, the genotypes from the two herbivory regimes were equally resistant to the rarer S. exigua caterpillars. As a consequence, resistance to one herbivore species did not correlate with resistance to another, suggesting independent evolution of resistance to multiple herbivores. 4. Plant growth correlated positively with resistance to Trirhabda, but not to Spodoptera, and only in plants from herbivore exclusion plots, leading us to reject the hypothesis of a growth resistance trade‐off. 5. Synthesis. Our results suggest that correlations between growth and resistance are context dependent and may only be apparent in populations relieved from certain natural pressures, such as in natural populations relieved from natural selection by herbivores.

Nitrogen use efficiency (NUE) has been widely used to study the relationship between nitrogen uptake and dry mass production in the plant. As a subsystem of plant nitrogen use efficiency (NUE), I have defined leaf-level NUE as the surplus production (gross production minus leaf respiration) per unit amount of nitrogen allocated to the leaf, with factorization into leaf nitrogen productivity (NP) and mean residence time of leaf nitrogen (MRT). These concepts were applied to two herbaceous stands: a perennial Solidago altissima stand and an annual Amaranthus patulus stand. S. altissima had more than three times higher leaf NUE than A. patulus due to nearly three times longer MRT of leaf N. In both species, NUE and NP were higher at the leaf level than at the plant level, because most leaf N is involved directly in the photosynthetic activity and because leaf surplus production is higher than the plant net production. MRT was longer at the plant level. The more than twice as long MRT at the plant level as at the leaf level in S. altissima was due to a large contribution of nitrogen storage belowground in the winter in this species. Thus, comparisons between a perennial and an annual system and between plant- and leaf-level NUE with their components revealed the importance of N allocation, storage, recycling, and turnover of organs for leaf photosynthetic production and plant dry mass growth.

Premise of the Study: Polyploidy in plants can result in genetic isolation, ecological differences among cytotypes, and, ultimately, speciation. Cytotypes should be sympatric only if they are segregated in an ecological niche or through prezygotic isolation. We tested whether sympatric diploid, tetraploid, and hexaploid ramets of Solidago altissima L. (Asteraceae) differ in their ecological niche. Methods: We measured how cytotypes were distributed within habitats, their morphology, and the composition of their communities of herbivorous insects at 10 natural field sites. We also conducted a common garden experiment to confirm whether observed differences in morphology or communities of herbivores were due to cytotype or environmental effects. Key Results: Diploid ramets often grew in open areas, relatively far from woody plants, and were associated with a high species richness of herbaceous plants, especially grasses. Hexaploids often grew in heavy shading under woody plants where grasses were scarce. Finally, tetraploids usually grew in transition areas between diploids and hexaploids. Hexaploid ramets also were taller than ramets of the other cytotypes and had larger leaves. Two species of insects, the leaf-galling fly Asteromyia carbonifera and the phloem-tapping aphid Uroleucon nigrotuberculatum, were more abundant on hexaploid ramets than on ramets of other cytotypes in the field. When grown in a common garden, however, cytotypes were similar in morphology and communities of herbivores. Conclusions: We conclude that cytotypes of S. altissima differ in their spatial distribution within habitats and that spatial variation in environmental factors influence plant morphology and communities of herbivorous insects.