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.

Aboveground plant-to-plant communication reduces root nodule symbiosis and soil nutrient concentrations.

Aboveground communication between plants is well known to change defense traits in leaves, but its effects on belowground plant traits and soil characteristics have not been elucidated. We hypothesized that aboveground plant-to-plant communication reduces root nodule symbiosis via induction of bactericidal chemical defense substances and changes the soil nutrient environment. Soybean plants were exposed to the volatile organic compounds (VOCs) from damaged shoots of Solidago canadensis var. scabra, and leaf defense traits (total phenolics, saponins), root saponins, and root nodule symbiosis traits (number and biomass of root nodules) were measured. Soil C/N ratios and mineral concentrations were also measured to estimate the effects of resource uptake by the plants. We found that total phenolics were not affected. However, plants that received VOCs had higher saponin concentrations in both leaves and roots, and fewer root nodules than untreated plants. Although the concentrations of soil minerals did not differ between treatments, soil C/N ratio was significantly higher in the soil of communicated plants. Thus, the aboveground plant-to-plant communication led to reductions in root nodule symbiosis and soil nutrient concentrations. Our results suggest that there are broader effects of induced chemical defenses in aboveground plant organs upon belowground microbial interactions and soil nutrients, and emphasize that plant response based on plant-to-plant communications are a bridge between above- and below-ground ecosystems.

Transcription-mediated tissue-specific lignification of vascular bundle causes trade-offs between growth and defence capacity during invasion of Solidago canadensis.

Allocation of more resources to growth but less to defense causing growth vigor of invasive alien plant populations contributes to successful invasion. However, few studies has addressed to relationship between vascular development variation and this mechanism. In this study, a common garden experimentwas established to compare the growth and vascular bundle development between native and introduced populations of Solidago canadensis, which is a wide-distributed invasive species in China. Our results suggested that the rapid growth of introduced populations could be explained by the well-developed and highly lignified xylem; while native populations present more developed and highly lignified phloem, which contributed more resistance to the infection of Sclerotiun rofsii compared with introduced populations. This difference was resulted from tissue-specific tradeoff distribution of lignification related gene expression between xylem and phloem, which is regulated by upstream MYB transcription factors. Our study gives a novel insight of mechanism that explain invasion success: lignin-related gene transcription-mediated tissue-specific lignification of vascular bundle contributes tradeoffs in resource allocation between growth and defence capacity during successful invasion of S. canadensis.

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.

Evidence that ERF transcriptional regulators serve as possible key molecules for natural variation in defense against herbivores in tall goldenrod.

We collected Solidago altissima clones to explore their leaf damage resistance, and as a result identified five accessions that exhibited variable defense abilities against the generalist herbivore Spodoptera litura. In order to characterize molecules involved in such natural variation, we focused on ethylene response factors (ERFs) that exhibited distinct transcription patterns in the leaves of the five accessions (e.g., S1 and S2) after wounding: the transcript of SaERF1 and SaERF2 was induced in wounded S1 and S2 leaves, respectively. Although transcription levels of SaERFs in leaves of the five accessions did not correlate with the accessions' phytohormone levels, these transcription levels accorded with the possibility that ethylene and jasmonate signaling play crucial roles in wound-induced transcription of SaERF1 in S1 leaves, and SaERF2 in S2 leaves, respectively. SaERF1 was found to be a positive regulator of the GCC box and DRE element in the upstream regions of promoters of defense genes, whereas SaERF2 served as a negative regulator of genes controlled through the GCC box. Transgenic Arabidopsis plants expressing SaERF1 or SaERF2 showed enhanced and suppressed transcript levels, respectively, of a defensin gene, indicating that ERFs may be partly responsible for herbivore resistance properties of S. altissima accessions.

Ecological rigidity and the hardness of selection in the wild.

Hutchinson's ecological theater and evolutionary play is a classical view of evolutionary ecology-ecology provides a template in which evolution occurs. An opposing view is that ecological and evolutionary changes are like two actors on a stage, intertwined by density and frequency dependence. These opposing views correspond to hard and soft selection, respectively. Although often presented as diametrically opposed, both types of selection can occur simultaneously, yet we largely lack knowledge of the relative importance of hard versus soft selection in the wild. I use a dataset of 3000 individual gall makers from 15 wild local populations over 5 years to investigate the hardness of selection. I show that enemy attack consistently favors some gall sizes over others (hard selection) but that these biases can be fine-tuned by density and frequency dependence (soft selection). As a result, selection is hard and soft in roughly equal measures, but the importance of each type varies as species interactions shift. I conclude that eco-evolutionary dynamics should occur when a mix of hard and soft selection acts on a population. This work contributes to the rapprochement of disparate views of evolutionary ecology-ecology is neither a rigid theater nor a flexible actor, but instead embodies components of both.

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.

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.

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.

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.

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.

No evidence for local adaptation to salt stress in the existing populations of invasive Solidago canadensis in China.

Local adaptation is an important mechanism underlying the adaptation of plants to environmental heterogeneity, and the toxicity of salt results in strong selection pressure on salt tolerance in plants and different ecotypes. Solidago canadensis, which is invasive in China, has spread widely and has recently colonized alkali sandy loams with a significant salt content. A common greenhouse experiment was conducted to test the role of local adaptation in the successful invasion of S. canadensis into salty habitats. Salt treatment significantly decreased the growth of S. canadensis, including rates of increase in the number of leaves and plant height; the root, shoot, and total biomass. Furthermore, salt stress significantly reduced the net photosynthetic rate, stomatal conductance, transpiration rate and relative chlorophyll content but significantly increased peroxidase activity and the proline content of S. canadensis and the root/shoot ratio. Two-way analysis of variance showed that salt treatment had a significant effect on the physiological traits of S. canadensis, except for the intercellular CO2 concentration, whereas the population and the salt × population interaction had no significant effect on any physiological traits. Most of the variation in plasticity existed within and not among populations, excep for the root/shoot ratio. S. canadensis populations from soil with moderate/high salt levels grew similarly to S. canadensis populations from soils with low salt levels. No significant correlation between salt tolerance indices and soil salinity levels was observed. The plasticity of the proline content, intercellular CO2 concentration and chlorophyll content had significant correlations with the salt tolerance index. These findings indicate a lack of evidence for local adaption in the existing populations of invasive S. canadensis in China; instead, plasticity might be more important than local adaptation in influencing the physiological traits and salt tolerance ability across the S. canadensis distribution.

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.

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.

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.

Seed Predators, not Herbivores, Exert Natural Selection on Solidago spp. in an Urban Archipelago.

The effects of urbanization on biodiversity are well established, as a growing city will reduce the size and diversity of patches of native plants. Recolonization of old patches and discovery of new ones by arthropod herbivores should occur as predicted by island biogeography theory. Although colonization represents an increase in biodiversity, such arrivals may exert new forms of natural selection on plants through herbivory and seed predation. Using a single species of old-field aster (Solidago altissima L.), we found that the level of natural selection by seed predators and herbivores follows patterns of island biogeography, with lower amounts of damage on smaller islands, where there are fewer species, and hypothetically smaller populations of arthropods. We also found that in an urban system, levels of herbivory are far below the tolerance levels of Solidago, and that seed predators are likely to be the only arthropod to cause reduced fitness. The pattern seen also implies that as a patch of Solidago grows through clonal expansion, it will come under higher selective pressure.

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.