The nitrogen-sulfur ratio of acid rain modulates the leaf- and root-mediated co-allelopathy of Solidago canadensis.

The majority of allelopathic studies on invasive plants have focused primarily on their leaf-mediated allelopathy, with relatively little attention paid to their root-mediated allelopathy, especially co-allelopathy mediated by both leaves and roots. It is conceivable that the diversified composition of acid rain may influence the allelopathy of invasive plants. This study aimed to evaluate the leaf and root-mediated co-allelopathy of the invasive plant Solidago canadensis L. under acid rain with different nitrogen-sulfur ratios (N/S) on Lactuca sativa L. via a hydroponic incubation. The root-mediated allelopathy of S. canadensis was found to be more pronounced than the leaf-mediated allelopathy of S. canadensis with nitric acid at pH 4.5, but the leaf-mediated allelopathy of S. canadensis was observed to be more pronounced than the root-mediated allelopathy of S. canadensis with sulfuric-rich acid at pH 4.5. The leaf and root-mediated co-allelopathy of S. canadensis was more pronounced than that of either part alone with sulfuric acid at pH 5.6 and nitric acid at pH 4.5, but not with nitric-rich acid at pH 4.5 and sulfuric-rich acid at pH 4.5. Sulfuric acid and sulfuric-rich acid with stronger acidity intensified the leaf-mediated allelopathy of S. canadensis. Nitric acid and nitric-rich acid attenuated the leaf-mediated allelopathy of S. canadensis, and most types of acid rain (especially nitric acid and nitric-rich acid) also attenuated the root-mediated allelopathy of S. canadensis and the leaf and root-mediated co-allelopathy of S. canadensis. Sulfuric acid and sulfuric-rich acid produced a more pronounced effect than nitric acid and nitric-rich acid. Hence, the N/S ratio of acid rain influenced the allelopathy of S. canadensis under acid rain with multiple N/S ratios.

Integrated analysis of mRNA-seq and miRNA-seq reveals the advantage of polyploid Solidago canadensis in sexual reproduction.

BACKGROUND: The invasion of Solidago canadensis probably related to polyploidy, which may promotes its potential of sexual reproductive. S. canadensis as an invasive species which rapidly widespread through yield huge numbers of seed, but the mechanism remains unknown. To better understand the advantages of sexual reproduction in hexaploid S. canadensis, transcriptome and small RNA sequencing of diploid and hexaploid cytotypes in flower bud and fruit development stages were performed in this study. RESULTS: The transcriptome analysis showed that in the flower bud stage, 29 DEGs were MADS-box related genes with 14 up-regulated and 15 down-regulated in hexaploid S. canadensis; 12 SPL genes were detected differentially expressed with 5 up-regulated and 7 down-regulated. In the fruit development stage, 26 MADS-box related genes with 20 up-regulated and 6 down-regulated in hexaploid S. canadensis; 5 SPL genes were all up-regulated; 28 seed storage protein related genes with 18 were up-regulated and 10 down-regulated. The weighted gene co-expression network analysis (WGCNA) identified 19 modules which consisted of co-expressed DEGs with functions such as sexual reproduction, secondary metabolism and transcription factors. Furthermore, we discovered 326 miRNAs with 67 known miRNAs and 259 novel miRNAs. Some of miRNAs, such as miR156, miR156a and miR156f, which target the sexual reproduction related genes. CONCLUSION: Our study provides a global view of the advantages of sexual reproduction in hexaploid S. canadensis based on the molecular mechanisms, which may promote hexaploid S. canadensis owing higher yield and fruit quality in the process of sexual reproduction and higher germination rate of seeds, and finally conductive to diffusion, faster propagation process and enhanced invasiveness.

Polyploidization-driven differentiation of freezing tolerance in Solidago canadensis.

Solidago canadensis, originating from the temperate region of North America, has expanded southward to subtropical regions through polyploidization. Here we investigated whether freezing tolerance of S. canadensis was weakened during expansion. Measurement of the temperature causing 50% ruptured cells (LT50 ) in 35 S. canadensis populations revealed ploidy-related differentiation in freezing tolerance. Freezing tolerance was found to decrease with increasing ploidy. The polyploid populations of S. canadensis had lower ScICE1 gene expression levels but more ScICE1 gene copies than the diploids. Furthermore, more DNA methylation sites in the ScICE1 gene promoter were detected in the polyploids than in the diploids. The results suggest that promoter methylation represses the expression of multi-copy ScICE1 genes, leading to weaker freezing tolerance in polyploid S. canadensis compared to the diploids. The study provides empirical evidence that DNA methylation regulates expression of the gene copies and supports polyploidization-driven adaptation to new environments.

The mixed silicon and cadmium synergistically impact the allelopathy of Solidago canadensis L. on native plant species Lactuca sativa L.

Several invasive alien plants (IAP) can trigger evidently allelopathy on the seed germination and seedling growth (SgSg) of native plant species (NPS). The getting worse condition with heavy metal pollution (e.g., cadmium) can significantly impact SgSg of plant species. Silicon can offset the adverse effects of environmental pressure on the growth and development of plant species. Thus, it is important to evaluate the influences of silicon on the allelopathy of IAP on SgSg of NPS under cadmium stress to better understand the mechanism driving the successful colonization of IAP. This study focuses on the allelopathy of the infamous IAP Solidago canadensis L. (Canada goldenrod; by using leaf extracts) on SgSg of NPS Lactuca sativa L. under the separated and mixed silicon and cadmium addition. S. canadensis triggers notably allelopathy on SgSg of L. sativa and gradually upsurges with increasing leaf extract concentration. Thus, the growth performance of NPS will be gradually reduced with an increasing degree of S. canadensis invasion. Cadmium evidently declines SgSg of L. sativa due to the broken balance of plant species for nutrient absorption. The mixed S. canadensis leaf extracts and cadmium synergistically impact seed germination of L. sativa but antagonistically affect seedling growth of L. sativa. The mixed silicon and cadmium intensify the allelopathy of S. canadensis on SgSg of L. sativa probably due to the increased effective content of cadmium in plant roots under silicon addition. Thus, the mixed silicon and cadmium will be advantageous to the following invasion process of IAP largely via the depressed SgSg of NPS.

Costs of plant defense priming: exposure to volatile cues from a specialist herbivore increases short-term growth but reduces rhizome production in tall goldenrod (Solidago altissima).

BACKGROUND: By sensing environmental cues indicative of pathogens or herbivores, plants can "prime" appropriate defenses and deploy faster, stronger responses to subsequent attack. Such priming presumably entails costs-else the primed state should be constitutively expressed-yet those costs remain poorly documented, in part due to a lack of studies conducted under realistic ecological conditions. We explored how defence priming in goldenrod (Solidago altissima) influenced growth and reproduction under semi-natural field conditions by manipulating exposure to priming cues (volatile emissions of a specialist herbivore, Eurosta solidaginis), competition between neighbouring plants, and herbivory (via insecticide application). RESULTS: We found that primed plants grew faster than unprimed plants, but produced fewer rhizomes, suggesting reduced capacity for clonal reproduction. Unexpectedly, this effect was apparent only in the absence of insecticide, prompting a follow-up experiment that revealed direct effects of the pesticide esfenvalerate on plant growth (contrary to previous reports from goldenrod). Meanwhile, even in the absence of pesticide, priming had little effect on herbivore damage levels, likely because herbivores susceptible to the primed defences were rare or absent due to seasonality. CONCLUSIONS: Reduced clonal reproduction in primed plants suggest that priming can entail significant costs for plants. These costs, however, may only become apparent when priming cues fail to provide accurate information about prevailing threats, as was the case in this study. Additionally, our insecticide data indicate that pesticides or their carrier compounds can subtly, but significantly, affect plant physiology and may interact with plant defences.

Constituents of a mixed-ploidy population of Solidago altissima differ in plasticity and predicted response to selection under simulated climate change.

PREMISE OF THE STUDY: Polyploids possess unique attributes that influence their environmental tolerance and geographic distribution. It is often unknown, however, whether cytotypes within mixed-ploidy populations are also uniquely adapted and differ in their responses to environmental change. Here, we examine whether diploids and hexaploids from a single mixed-ploidy population of Solidago altissima differ in plasticity and potential response to natural selection under conditions simulating climate change. METHODS: Clonal replicates of diploid and hexaploid genotypes were grown in a randomized split-plot design under two temperature (+1.9°C) and two watering treatments (-13% soil moisture) implemented with open-top passive chambers placed under rainout shelters. Physiological, phenological, morphological traits, and a fitness correlate, reproductive biomass, were measured and compared among treatments. KEY RESULTS: Differences in traits suggest that diploids are currently better adapted to low- water availability than hexaploids. Both ploidy levels had adaptive plastic responses to treatments and are predicted to respond to selection, but often for different traits. Water availability generally had a stronger effect than temperature, but for some traits the effect of water depended on temperature. CONCLUSIONS: Diploid and hexaploid S. altissima may maintain fitness in the short term through adaptive plasticity and evolution depending on which traits are important in a warmer, drier environment. Hexaploids may be at a disadvantage compared to diploids because fewer traits were heritable. Our results underscore the importance of studying combinations of climate variables that are predicted to change simultaneously.

Impact of the invasive plant Solidago gigantea on soil nematodes in a semi-natural grassland and a temperate broadleaved mixed forest.

Relationships between alien plant species and their aboveground effects have been relatively well studied, but little is known about the effects of invasive plants on belowground faunal communities. Nematodes are abundant, ubiquitous and diverse soil biota, and alterations of their community compositions can illustrate changes in belowground ecosystems. In 2016 and 2017, we determined the response of species diversity, community composition and trophic composition of the soil nematode communities to invasion by the alien plant Solidago gigantea in two ecosystems, forest and grassland, where invasion takes place. Nematode abundance was higher and number of identified nematode species was lower at invaded than uninvaded sites, indicated by lower species diversity, regardless of ecosystem. Herbivorous nematodes were the most affected trophic group. Herbivore abundance was higher at invaded than uninvaded sites and in grassland than forest. The herbivorous species Boleodorus thylactus, Geocenamus sp., Helicotylenchus spp., Paratylenchus bukowinensis, Pratylenchoides crenicauda and Rotylenchus robustus were more abundant at the invaded sites. Abundances of nematodes in the other tropic groups were limited or not affected. The invasion did not significantly affect the ecological and functional indices, except for the Channel Index in 2016. Differences were observed in values of Enrichment Index (indicator of resource availability), Channel Index (indicator of ascendant bacterial/fungal decomposition channel) and Basal Index (indicator of depleted-perturbed soil food webs) between grassland and forests. We can thus conclude that invasion by S. gigantea significantly alters nematode community indicators (abundance, species diversity and specific trophic groups); however, this effect seems to be significantly influenced by the type of ecosystem where invasion takes place.

Comparative study of the morphological and phytochemical characterization of Romanian Solidago species.

In the present study, three indigenous species of Solidago genus (Solidago gigantea, Solidago virgaurea and Solidago canadensis) have been analyzed for the assessment of polyphenolic, phenyl propane derivates and essential oil contents. In addition, a comparative morphological study was also described. The leaves and the flowers of the three Solidago species were studied by scanning electron microscopy (SEM). The qualitative and quantitative characterizations of the main polyphenolic compounds from the hydrolyzed extracts were carried out by using high performance liquid chromatography with UV detection (HPLC-UV), high performance liquid chromatography coupled with mass spectrometry (HPLC-MS) and gas chromatography coupled with mass spectrometry (GC-MS) for the essential oil determination. The dominant flavonoidic aglycone found for all three species was quercetol with its highest concentration registered in Solidago canadensis. Four components, α-pinene, mircene, bornyl acetate and germacrene D, were detected in all the analyzed samples of essential oils. According to the comparative morphological analysis, morphoanatomical differences were observed for the tryhomes, stomata and flowers of the studied Romanian Solidago species.

Local root growth and death are mediated by contrasts in nutrient availability and root quantity between soil patches.

Plants are thought to be able to regulate local root growth according to its overall nutrient status as well as nutrient contents in a local substrate patch. Therefore, root plastic responses to environmental changes are probably co-determined by local responses of root modules and systematic control of the whole plant. Recent studies showed that the contrast in nutrient availability between different patches could significantly influence the growth and death of local roots. In this study, we further explored, beside nutrient contrast, whether root growth and death in a local patch are also affected by relative root quantity in the patch. We conducted a split-root experiment with different splitting ratios of roots of Canada goldenrod (Solidago canadensis) individuals, as well as high- (5× Hoagland solution versus water) or low- (1× Hoagland solution versus water) contrast nutrient conditions for the split roots. The results showed that root growth decreased in nutrient-rich patches but increased in nutrient-poor patches when more roots co-occurred in the same patches, irrespective of nutrient contrast condition. Root mortality depended on contrasts in both root quantity and nutrients: in the high-nutrient-contrast condition, it increased in nutrient-rich patches but decreased in nutrient-poor patches with increasing root proportion; while in the low-nutrient-contrast condition, it showed the opposite trend. These results demonstrated that root growth and death dynamics were affected by the contrast in both nutrient availability and root quantity between patches. Our study provided ecological evidence that local root growth and death are mediated by both the responses of root modules to a nutrient patch and the whole-plant nutrient status, suggesting that future work investigating root production and turnover should take into account the degree of heterogeneity in nutrient and root distribution.

Defense with benefits? Ducking plants outperformed erect plants in the goldenrod Solidago gigantea in the absence of herbivory.

PREMISE OF THE STUDY: Despite the fact that herbivores can be highly detrimental to their host plants' fitness, plant populations often maintain genetic variation for resistance to their natural enemies. Investigating the various costs (e.g., allocation tradeoffs, autotoxicity, and ecological costs) that may prevent plants from evolving to their fullest potential resistance has been a productive strategy for shedding insight into the eco-evolutionary dynamics of plant-herbivore communities. METHODS: Recent studies have shown that some individuals of goldenrod (Solidago spp.) evade apex-attacking herbivores by a temporary nodding of their stem (i.e., resistance-by-ducking). Although ducking provides an obvious fitness benefit to these individuals, nonducking (erect) morphs persist in goldenrod populations. In this study, I investigated potential costs of ducking in Solidago gigantea in terms of tradeoffs involving growth and reproduction in a common garden experiment using field-collected seeds. KEY RESULTS: The S. gigantea population contained substantial genetic variation for stem morph, with 28% erect and 72% ducking stems. In the absence of herbivory, ducking plants were taller, had thicker stems, and produced an average of 20% more seeds than erect plants. CONCLUSIONS: This study suggests that resistance-by-ducking, instead of being costly, actually comes with additional, nondefense-related benefits. These results support the conclusion that the factors that constrain the evolution of resistance in plant populations are likely to be more subtle and complex than simple tradeoffs in resource allocation.

Constraints on the evolution of resistance to gall flies in Solidago altissima: resistance sometimes costs more than it is worth.

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.

Rising atmospheric CO2 is reducing the protein concentration of a floral pollen source essential for North American bees.

At present, there is substantive evidence that the nutritional content of agriculturally important food crops will decrease in response to rising levels of atmospheric carbon dioxide, Ca However, whether Ca-induced declines in nutritional quality are also occurring for pollinator food sources is unknown. Flowering late in the season, goldenrod (Solidago spp.) pollen is a widely available autumnal food source commonly acknowledged by apiarists to be essential to native bee (e.g. Bombus spp.) and honeybee (Apis mellifera) health and winter survival. Using floral collections obtained from the Smithsonian Natural History Museum, we quantified Ca-induced temporal changes in pollen protein concentration of Canada goldenrod (Solidago canadensis), the most wide spread Solidago taxon, from hundreds of samples collected throughout the USA and southern Canada over the period 1842-2014 (i.e. a Ca from approx. 280 to 398 ppm). In addition, we conducted a 2 year in situtrial of S. Canadensis populations grown along a continuous Ca gradient from approximately 280 to 500 ppm. The historical data indicated a strong significant correlation between recent increases in Ca and reductions in pollen protein concentration (r(2)= 0.81). Experimental data confirmed this decrease in pollen protein concentration, and indicated that it would be ongoing as Ca continues to rise in the near term, i.e. to 500 ppm (r(2)= 0.88). While additional data are needed to quantify the subsequent effects of reduced protein concentration for Canada goldenrod on bee health and population stability, these results are the first to indicate that increasing Ca can reduce protein content of a floral pollen source widely used by North American bees.

Solidago gigantea plants from nonnative ranges compensate more in response to damage than plants from the native range.

Resistance and tolerance are two ways that plants cope with herbivory. Tolerance, the ability of a plant to regrow or reproduce after being consumed, has been studied less than resistance, but this trait varies widely among species and has considerable potential to affect the ecology of plant species. One particular aspect of tolerance, compensatory responses, can evolve rapidly in plant species; providing insight into interactions between consumers and plants. However, compensation by invasive species has rarely been explored. We compared compensatory responses to the effects of simulated herbivory expressed by plants from seven Solidago gigantea populations from the native North American range to that expressed by plants from nine populations from the nonnative European range. Populations were also collected along elevational gradients to compare ecotypic variation within and between ranges. Solidago plants from the nonnative range of Europe were more tolerant to herbivory than plants from the native range of North America. Furthermore, plants from European populations increased in total biomass and growth rate with elevation, but decreased in compensatory response. There were no relationships between elevation and growth or compensation for North American populations. Our results suggest that Solidago gigantea may have evolved to better compensate for herbivory damage in Europe, perhaps in response to a shift to greater proportion of attack from generalists. Our results also suggest a possible trade-off between rapid growth and compensation to damage in European populations but not in North American populations.

The allelopathic effects of invasive plant Solidago canadensis on seed germination and growth of Lactuca sativa enhanced by different types of acid deposition.

Invasive species can exhibit allelopathic effects on native species. Meanwhile, the types of acid deposition are gradually changing. Thus, the allelopathic effects of invasive species on seed germination and growth of native species may be altered or even enhanced under conditions with diversified acid deposition. This study aims to assess the allelopathic effects (using leaves extracts) of invasive plant Solidago canadensis on seed germination and growth of native species Lactuca sativa treated with five types of acid deposition with different SO4(2-) to NO3(-) ratios (1:0, sulfuric acid; 5:1, sulfuric-rich acid; 1:1, mixed acid; 1:5, nitric-rich acid; 0:1, nitric acid). Solidago canadensis leaf extracts exhibited significantly allelopathic effects on germination index, vigor index, and germination rate index of L. sativa. High concentration of S. canadensis leaf extracts also similarly exhibited significantly allelopathic effects on root length of L. sativa. This may be due to that S. canadensis could release allelochemicals and then trigger allelopathic effects on seed germination and growth of L. sativa. Acid deposition exhibited significantly negative effects on seedling biomass, root length, seedling height, germination index, vigor index, and germination rate index of L. sativa. This may be ascribed to the decreased soil pH values mediated by acid deposition which could produce toxic effects on seedling growth. Sulfuric acid deposition triggered more toxic effects on seedling biomass and vigor index of L. sativa than nitric acid deposition. This may be attributing to the difference in exchange capacity with hydroxyl groups (OH(-)) between SO4(2-) and NO3(-) as well as the fertilizing effects mediated by nitric deposition. All types of acid deposition significantly enhanced the allelopathic effects of S. canadensis on root length, germination index, vigor index, and germination rate index of L. sativa. This may be due to the negatively synergistic effects of acid deposition and S. canadensis on seed germination and growth of L. sativa. The ratio of SO4(2-) to NO3(-) in acid deposition was an important factor that profoundly affected the allelopathic effects of S. canadensis on the seed germination and growth of L. sativa possibly because the difference in exchange capacity with hydroxyl groups (OH(-)) between SO4(2-) and NO3(-) as well as the fertilizing effects triggered by nitric deposition. Thus, the allelopathic effects of invasive species on seed germination and growth of native plants might be enhanced under increased and diversified acid deposition.

Plant neighborhood effects on herbivory: damage is both density and frequency dependent.

Neighboring plants can affect the likelihood that a focal plant is attacked by herbivores. Both the density of conspecific neighbors (resource concentration or dilution effects) and the relative density of heterospecific neighbors (associational effects or effects of neighbor frequency) within the local neighborhood can affect herbivore load and plant damage. Understanding how these neighborhood effects influence processes such as plant competition or natural selection on plant resistance traits will require knowing how both plant density and frequency affect damage, but previous studies have generally confounded density and frequency effects. In this study, we independently manipulated the absolute density and frequency (i.e., relative density) of two plant species (Solanum carolinense and Solidago altissima) to characterize neighborhood composition effects on S. carolinense damage by herbivores, providing the first picture of how both density and frequency of neighbors influence damage in a single system. We found both a positive effect of S. carolinense density on S. carolinense damage (a resource concentration effect) and a nonlinear effect of S. altissima frequency on S. carolinense damage (associational susceptibility). If these types of patterns are common in nature, future studies seeking to understand neighborhood effects on damage need to incorporate both density and frequency effects and capture any nonlinear effects by selecting a range of values rather than focusing on only a pair of densities or frequencies. This type of data on neighborhood effects will allow us to understand the contribution of neighborhood effects to population-level processes such as competition, the evolution of plant resistance to herbivores, and yield gains in agricultural crop mixtures.

Informed herbivore movement and interplant communication determine the effects of induced resistance in an individual-based model.

1. Plant induced resistance to herbivory affects the spatial distribution of herbivores, as well as their performance. In recent years, theories regarding the benefit to plants of induced resistance have shifted from ideas of optimal resource allocation towards a more eclectic set of theories that consider spatial and temporal plant variability and the spatial distribution of herbivores among plants. However, consensus is lacking on whether induced resistance causes increased herbivore aggregation or increased evenness, as both trends have been experimentally documented. 2. We created a spatial individual-based model that can describe many plant-herbivore systems with induced resistance, in order to analyse how different aspects of induced resistance might affect herbivore distribution, and the total damage to a plant population, during a growing season. 3. We analyse the specific effects on herbivore aggregation of informed herbivore movement (preferential movement to less-damaged plants) and of information transfer between plants about herbivore attacks, in order to identify mechanisms driving both aggregation and evenness. We also investigate how the resulting herbivore distributions affect the total damage to plants and aggregation of damage. 4. Even, random and aggregated herbivore distributions can all occur in our model with induced resistance. Highest levels of aggregation occurred in the models with informed herbivore movement, and the most even distributions occurred when the average number of herbivores per plant was low. With constitutive resistance, only random distributions occur. Damage to plants was spatially correlated, unless plants recover very quickly from damage; herbivore spatial autocorrelation was always weak. 5. Our model and results provide a simple explanation for the apparent conflict between experimental results, indicating that both increased aggregation and increased evenness of herbivores can result from induced resistance. We demonstrate that information transfer from plants to herbivores, and from plants to neighbouring plants, can both be major factors in determining non-random herbivore distributions.

Rapid plant evolution in the presence of an introduced species alters community composition.

Because introduced species may strongly interact with native species and thus affect their fitness, it is important to examine how these interactions can cascade to have ecological and evolutionary consequences for whole communities. Here, we examine the interactions among introduced Rocky Mountain elk, Cervus canadensis nelsoni, a common native plant, Solidago velutina, and the diverse plant-associated community of arthropods. While introduced species are recognized as one of the biggest threats to native ecosystems, relatively few studies have investigated an evolutionary mechanism by which introduced species alter native communities. Here, we use a common garden design that addresses and supports two hypotheses. First, native S. velutina has rapidly evolved in the presence of introduced elk. We found that plants originating from sites with introduced elk flowered nearly 3 weeks before plants originating from sites without elk. Second, evolution of S. velutina results in a change to the plant-associated arthropod community. We found that plants originating from sites with introduced elk supported an arthropod community that had ~35 % fewer total individuals and a different species composition. Our results show that the impacts of introduced species can have both ecological and evolutionary consequences for strongly interacting species that subsequently cascade to affect a much larger community. Such evolutionary consequences are likely to be long-term and difficult to remediate.

The volatile emission of Eurosta solidaginis primes herbivore-induced volatile production in Solidago altissima and does not directly deter insect feeding.

BACKGROUND: The induction of plant defenses in response to herbivory is well documented. In addition, many plants prime their anti-herbivore defenses following exposure to environmental cues associated with increased risk of subsequent attack, including induced volatile emissions from herbivore-damaged plant tissues. Recently, we showed in both field and laboratory settings that tall goldenrod plants (Solidago altissima) exposed to the putative sex attractant of a specialist gall-inducing fly (Eurosta solidaginis) experienced less herbivory than unexposed plants. Furthermore, we observed stronger induction of the defense phytohormone jasmonic acid in exposed plants compared to controls. These findings document a novel class of plant-insect interactions mediated by the direct perception, by plants, of insect-derived olfactory cues. However, our previous study did not exclude the possibility that the fly emission (or its residue) might also deter insect feeding via direct effects on the herbivores. RESULTS: Here we show that the E. solidaginis emission does not (directly) deter herbivore feeding on Cucurbita pepo or Symphyotrichum lateriflorum plants--which have no co-evolutionary relationship with E. solidaginis and thus are not expected to exhibit priming responses to the fly emission. We also document stronger induction of herbivore-induced plant volatiles (HIPV) in S. altissima plants given previous exposure to the fly emission relative to unexposed controls. No similar effect was observed in maize plants (Zea mays), which have no co-evolutionary relationship with E. solidaginis. CONCLUSIONS: Together with our previous findings, these results provide compelling evidence that reduced herbivory on S. altissima plants exposed to the emission of male E. solidaginis reflects an evolved plant response to olfactory cues associated with its specialist herbivore and does not involve direct effects of the fly emission on herbivore feeding behavior. We further discuss mechanisms by which the priming of HIPV responses documented here might contribute to enhanced S. altissima defense against galling.

Exploring plant defense theory in tall goldenrod, Solidago altissima.

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.

Physiological acclimation to drought stress in Solidago canadensis.

Solidago canadensis is an invasive species from North America that is spreading across Europe, Australia and temperate Asia. We hypothesized that the species' wide ecological amplitude is also based on its potential in hydraulic acclimation, and analyzed hydraulic and anatomical properties along a transect with decreasing soil humidity. Stem hydraulic conductivity, vulnerability to drought-induced embolism, stomatal closure during dehydration and xylem-anatomical parameters were quantified at three sites. At the humid site, specific hydraulic conductivity of stems (1.0 ± 0.2 kg m(-1) MPa(-1) s(-1)) was about twofold higher, and leaf-specific conductivity about 1.5 times higher (3.1 ± 0.5 kg m(-1) MPa(-1) s(-1)) than at the dry site. Water potential (Ψ) at 50% loss of conductivity was -3.7 ± 0.1 MPa at the dry site and -3.1 ± 0.2 MPa at the humid site (September). Vulnerability to drought-induced embolism decreased along the transect and over the vegetation period. At drier sites, stomata started closing at lower Ψ while complete stomatal closure was reached at less negative Ψ (12% of maximum stomatal conductance: -2.5 ± 0.0 and -3.0 ± 0.2 MPa at the dry and humid site). The safety margin between stomatal closure and 50% loss of conductivity was 1.2 and 0.2 MPa at the dry and humid sites. The observed variability indicated an efficient acclimation in hydraulic conductivity and safety: plants at dry sites exhibited lower specific hydraulic conductivity, higher embolism resistance and broader safety margins, signifying a trade-off between the hydraulic safety and efficiency. The observed intraspecific plasticity in hydraulic and anatomical traits may help to explain the invasive potential of this species.