Plant Chemical Defenses Research Articles

Gene expression plasticity across hosts of an invasive scale insect species.

For plant-eating insects, we still have only a nascent understanding of the genetic basis of host-use promiscuity. Here, to improve that situation, we investigated host-induced gene expression plasticity in the invasive lobate lac scale insect, Paratachardina pseudolobata (Hemiptera: Keriidae). We were particularly interested in the differential expression of detoxification and effector genes, which are thought to be critical for overcoming a plant’s chemical defenses. We collected RNA samples from P. pseudolobata on three different host plant species, assembled transcriptomes de novo, and identified transcripts with significant host-induced gene expression changes. Gene expression plasticity was pervasive, but the expression of most detoxification and effector genes was insensitive to the host environment. Nevertheless, some types of detoxification genes were more differentially expressed than expected by chance. Moreover, we found evidence of a trade-off between expression of genes involved in primary and secondary metabolism; hosts that induced lower expression of genes for detoxification induced higher expression of genes for growth. Our findings are largely consonant with those of several recently published studies of other plant-eating insect species. Thus, across plant-eating insect species, there may be a common set of gene expression changes that enable host-use promiscuity.

Plant chemical defense indirectly mediates aphid performance via interactions with tending ants.

The benefits of mutualistic interactions are often highly context dependent. We studied the interaction between the milkweed aphid Aphis asclepiadis and a tending ant, Formica podzolica. Although this interaction is generally considered beneficial, variation in plant genotype may alter it from mutualistic to antagonistic. Here we link the shift in strength and relative benefit of the ant-aphid interaction to plant genotypic variation in the production of cardenolides, a class of toxic defensive chemicals. In a field experiment with highly variable genotypes of the common milkweed (Asclepias syriaca), we show that plant cardenolides, especially polar forms, are ingested by aphids and excreted in honeydew proportionally to plant concentrations without directly affecting aphid performance. Ants consume honeydew, and aphids that excreted high amounts of cardenolides received fewer ant visits, which in turn reduced aphid survival. On at least some plant genotypes, aphid numbers per plant were reduced in the presence of ants to levels lower than in corresponding ant-exclusion treatments, suggesting antagonistic ant behavior. Although cardenolides appear ineffective as direct plant defenses against aphids, the multi-trophic context reveals an ant-mediated negative indirect effect on aphid performance and population dynamics.

How Herbivore Browsing Strategy Affects Whole-Plant Photosynthetic Capacity.

If a browse damage index indicates that a tree has been 50% browsed by herbivores, does this mean half the leaves are entirely eaten or are all the leaves half eaten? Were the affected leaves old or young? Large or small? In sunshine or shade? Understanding what effect browsing will have on the photosynthetic capacity and the plant's survival ability clearly requires a greater understanding of browsing strategy across the canopy than can be given by a single index value. We developed stochastic models of leaf production, growth and consumption using data from kamahi (Weinmannia racemosa) trees in New Zealand which have been browsed by possums (Trichosurus vulpecula), to ascertain which of six feasible browsing strategies possums are most likely to be employing. We compared the area distribution of real fallen leaves to model output in order to select the best model, and used the model to predict the age distribution of leaves on the tree and thus infer its photosynthetic capability. The most likely browsing strategy that possums employ on kamahi trees is a preference for virgin (i.e. previously unbrowsed) leaves, consistent with the idea that browsing increases the production of chemical plant defences. More generally, our results show that herbivore browsing strategy can significantly change the whole-plant photosynthetic capability of any plant and hence its ability to survive, and therefore, herbivore damage indices should be used in conjunction with more detailed information about herbivore browsing strategy.

Borrowed plant defences: Deterring browsers using a forestry by-product

Mammalian browsers can cause ecologically and economically significant damage to important tree species, particularly at the vulnerable seedling stage. Some tree species or particular parts of trees are less preferred as a result of chemical defences – harnessing such defences could reduce the appeal of more preferred trees and thus their loss to browsing. We tested the potential of birch bark extract, a forestry by-product, to protect seedlings of Scots pine Pinus sylvestris from two globally significant browsers: moose Alces alces and red deer Cervus elaphus. We quantified the responses by captive animals to extract application as a function of both the distance between seedlings and whether the extract was applied to all or alternate seedlings. Both moose and red deer consumed pine seedlings less frequently when seedlings were treated with birch bark extract, but their responses differed when only alternate seedlings were treated. At inter-seedling distances equivalent to those used in forestry, red deer browsed untreated seedlings more frequently when alternate seedlings were treated than when none were treated (associational susceptibility), whereas moose browsed untreated seedlings less frequently with treated than untreated neighbours (associational refuge). These neighbourhood effects were not evident at inter-seedling distances three times larger or smaller than those used in forestry. There was also no significant difference in the frequency of browsing on treated seedlings in alternate compared to completely treated arrangements at any inter-seedling distance. Red deer removed significantly less foliage from treated than untreated seedlings once browsed. Browsing by moose followed a similar but non-significant trend. The relative number of untreated to treated seedlings browsed was 2:1 for red deer and 4:1 for moose. Red deer used a finer scale of selection than moose for choosing between seedlings, consistent with their smaller body mass. However, the coarser-scale of selection used by moose resulted in consumption of more untreated relative to treated seedlings. Our study highlights the potential of existing plant defensive chemistry, harvested from forestry by-products, to reduce herbivore damage to tree seedlings. We also demonstrate the need to consider both plant associational effects and the foraging strategies of the dominant herbivores in applying repellents.

Complete genome of Pieris rapae, a resilient alien, a cabbage pest, and a source of anti-cancer proteins

The Small Cabbage White ( Pieris rapae) is originally a Eurasian butterfly. Being accidentally introduced into North America, Australia, and New Zealand a century or more ago, it spread throughout the continents and rapidly established as one of the most abundant butterfly species. Although it is a serious pest of cabbage and other mustard family plants with its caterpillars reducing crops to stems, it is also a source of pierisin, a protein unique to the Whites that shows cytotoxicity to cancer cells. To better understand the unusual biology of this omnipresent agriculturally and medically important butterfly, we sequenced and annotated the complete genome from USA specimens. At 246 Mbp, it is among the smallest Lepidoptera genomes reported to date. While 1.5% positions in the genome are heterozygous, they are distributed highly non-randomly along the scaffolds, and nearly 20% of longer than 1000 base-pair segments are SNP-free (median length: 38000 bp). Computational simulations of population evolutionary history suggest that American populations started from a very small number of introduced individuals, possibly a single fertilized female, which is in agreement with historical literature. Comparison to other Lepidoptera genomes reveals several unique families of proteins that may contribute to the unusual resilience of Pieris. The nitrile-specifier proteins divert the plant defense chemicals to non-toxic products. The apoptosis-inducing pierisins could offer a defense mechanism against parasitic wasps. While only two pierisins from Pieris rapae were characterized before, the genome sequence revealed eight, offering additional candidates as anti-cancer drugs. The reference genome we obtained lays the foundation for future studies of the Cabbage White and other Pieridae species.

Community-level relaxation of plant defenses against herbivores at high elevation

Ecological gradients shape the evolution of plant functional traits through variation in climate, abiotic factors, and biological processes. While recent observations highlight elevation clines in functional trait variation, several traits vary independently of others and vary in species-specific manner, thereby hindering community-level generalization. In this study, we examined whether whole plant communities at different elevations independently converge to elevation-specific trait values. Using a combination of field-derived trait quantification and a literature survey, we analyzed eight functional traits relating to plant palatability [specific leaf area (SLA), leaf dry matter content, carbon-to-nitrogen ratio], and defenses (spines, trichomes, silica, and the relative importance of phenolic- or non-phenolic-based chemical defenses in plants) of plants growing along several mountain transects in Switzerland, and computed community-level means for each trait. We observed a general decline of most defense traits in communities at high elevations. The individual traits’ patterns underlined a concerted decline of the physico-chemical defense syndrome at high elevation, and low SLA values at high elevation also indicate investment in the production of leaves with slow turnover rates. In addition, we found that high-elevation plant trait values tend to converge, while at low elevation, to diverge within communities. Our data suggest that community-level defense syndromes along elevation gradients are influenced by ecological filtering. Lower herbivore pressures select for reduced plant defenses, while harsh environmental conditions select for investment in tolerance-related traits. Secondly, since species are highly variable in their functional trait space within communities, niche-partitioning theory might explain which variabilities of the defense functional space are selected within communities, particularly at low elevation.

NSP-Dependent Simple Nitrile Formation Dominates upon Breakdown of Major Aliphatic Glucosinolates in Roots, Seeds, and Seedlings of Arabidopsis thaliana Columbia-0.

One of the best-studied plant defense systems, the glucosinolate-myrosinase system of the Brassicales, is composed of thioglucosides known as glucosinolates and their hydrolytic enzymes, the myrosinases. Tissue disruption brings these components together, and bioactive products are formed as a consequence of myrosinase-catalyzed glucosinolate hydrolysis. Among these products, isothiocyanates have attracted most interest as chemical plant defenses against herbivores and pathogens and health-promoting compounds in the human diet. Previous research has identified specifier proteins whose presence results in the formation of alternative product types, e.g., nitriles, at the expense of isothiocyanates. The biological roles of specifier proteins and alternative breakdown products are poorly understood. Here, we assessed glucosinolate breakdown product profiles obtained upon maceration of roots, seedlings and seeds of Arabidopsis thaliana Columbia-0. We identified simple nitriles as the predominant breakdown products of the major endogenous aliphatic glucosinolates in root, seed, and seedling homogenates. In agreement with this finding, genes encoding nitrile-specifier proteins (NSPs) are expressed in roots, seeds, and seedlings. Analysis of glucosinolate breakdown in mutants with T-DNA insertions in any of the five NSP genes demonstrated, that simple nitrile formation upon tissue disruption depended almost entirely on NSP2 in seeds and mainly on NSP1 in seedlings. In roots, about 70–80% of the nitrile-forming activity was due to NSP1 and NSP3. Thus, glucosinolate breakdown product profiles are organ-specifically regulated in A. thaliana Col-0, and high proportions of simple nitriles are formed in some parts of the plant. This should be considered in future studies on biological roles of the glucosinolate-myrosinase system.

Complete genome of Pieris rapae, a resilient alien, a cabbage pest, and a source of anti-cancer proteins.

The Small Cabbage White ( Pieris rapae) is originally a Eurasian butterfly. Being accidentally introduced into North America, Australia, and New Zealand a century or more ago, it spread throughout the continents and rapidly established as one of the most abundant butterfly species. Although it is a serious pest of cabbage and other mustard family plants with its caterpillars reducing crops to stems, it is also a source of pierisin, a protein unique to the Whites that shows cytotoxicity to cancer cells. To better understand the unusual biology of this omnipresent agriculturally and medically important butterfly, we sequenced and annotated the complete genome from USA specimens. At 246 Mbp, it is among the smallest Lepidoptera genomes reported to date. While 1.5% positions in the genome are heterozygous, they are distributed highly non-randomly along the scaffolds, and nearly 20% of longer than 1000 base-pair segments are SNP-free (median length: 38000 bp). Computational simulations of population evolutionary history suggest that American populations started from a very small number of introduced individuals, possibly a single fertilized female, which is in agreement with historical literature. Comparison to other Lepidoptera genomes reveals several unique families of proteins that may contribute to the unusual resilience of Pieris. The nitrile-specifier proteins divert the plant defense chemicals to non-toxic products. The apoptosis-inducing pierisins could offer a defense mechanism against parasitic wasps. While only two pierisins from Pieris rapae were characterized before, the genome sequence revealed eight, offering additional candidates as anti-cancer drugs. The reference genome we obtained lays the foundation for future studies of the Cabbage White and other Pieridae species.

Pine Defensive Monoterpene α-Pinene Influences the Feeding Behavior of Dendroctonus valens and Its Gut Bacterial Community Structure.

The exposure to plant defense chemicals has negative effects on insect feeding activity and modifies insect gut microbial community composition. Dendroctonus valens is a very destructive forest pest in China, and harbors a large diversity and abundance of gut microorganisms. Host pine defensive chemicals can protect the pines from attack by the holobiont. In this study, boring length of D. valens feeding on 0 mg/g α-pinene and 9 mg/g α-pinene concentration in phloem media for 6 and 48 h were recorded, and their gut bacterial communities were analyzed in parallel. Nine milligram per gram α-pinene concentration significantly inhibited boring length of D. valens and altered its gut microbial community structure after 6 h. The inhibition of boring length from 9 mg/g α-pinene in diets ceased after 48 h. No significant differences of the bacterial communities were observed between the beetles in 0 and 9 mg/g α-pinene concentration in phloem media after 48 h. Our results showed that the inhibition of the feeding behavior of D. valens and the disturbance to its gut bacterial communities in 9 mg/g α-pinene concentration in phloem media after 6 h were eliminated after 48 h. The resilience of gut bacterial community of D. valens may help the beetle catabolize pine defense chemical.

Choosy mothers pick challenging plants: maternal preference and larval performance of a specialist herbivore are not linked

1. Maternal preference is a dynamic process and interactions between preference and performance are fundamental for understanding evolutionary ecology and host association in insect–plant interactions. In the present study, the hypothesis of preference–performance was tested by offering solanaceous specialist Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae) larvae and adult females four plant congeners that ranged in suitability.2. Larval feeding, development, oviposition, plant glycoalkaloids, and headspace volatiles in the four plant species were analysed to examine the extent of variation, which might explain performance–preference differences.3. It was found that larval performance was mismatched with adult oviposition preferences. Adults laid more eggs on Solanum immite Dunal plants, which were poor hosts for larval development, feeding, and survival, compared to the other three Solanum species.4. Chemical plant defenses, in general, did not correlate with performance or preference, but some plant volatiles may have played a role in resolving female choice. Glycoalkaloids such as solanine and chaconine were detected in similar amounts in preferred and non‐preferred hosts, but there was significantly more limonene in the headspace of S. immite than in S. tuberosum L.5. The present findings suggest that we must consider the risk‐spreading hypothesis in cases where preference and performance are not positively correlated, particularly in specialist herbivores that can feed on a diversity of congener plants and may attempt to expand their exploits to other solanaceae species.

Characterization and expression profiling of ATP-binding cassette transporter genes in the diamondback moth, Plutella xylostella (L.).

BackgroundATP-binding cassette (ABC) transporters are one of the major transmembrane protein families found in all organisms and play important roles in transporting a variety of compounds across intra and extra cellular membranes. In some species, ABC transporters may be involved in the detoxification of substances such as insecticides. The diamondback moth, Plutella xylostella (L.), a destructive pest of cruciferous crops worldwide, is an important species to study as it is resistant to many types of insecticides as well as biological control Bacillus thuringiensis toxins.ResultsA total of 82 ABC genes were identified from our published P. xylostella genome, and grouped into eight subfamilies (ABCA-H) based on phylogenetic analysis. Genes of subfamilies ABCA, ABCC and ABCH were found to be expanded in P. xylostella compared with those in Bombyx mori, Manduca sexta, Heliconius melpomene, Danaus plexippus, Drosophila melanogaster, Tetranychus urticae and Homo sapiens. Phylogenetic analysis indicated that many of the ABC transporters in P. xylostella are orthologous to the well-studied ABC transporter genes in the seven other species. Transcriptome- and qRT-PCR-based analysis elucidated physiological effects of ABC gene expressions of P. xylostella which were developmental stage- and tissue-specific as well as being affected by whether or not the insects were from an insecticide-resistant strain. Two ABCC and one ABCA genes were preferentially expressed in midgut of the 4th-instar larvae of a susceptible strain (Fuzhou-S) suggesting their potential roles in metabolizing plant defensive chemicals. Most of the highly expressed genes in insecticide-resistant strains were also predominantly expressed in the tissues of Malpighian tubules and midgut.ConclusionsThis is the most comprehensive study on identification, characterization and expression profiling of ABC transporter genes in P. xylostella to date. The diversified features and expression patterns of this gene family may be associated with the evolutionary capacity of this species to develop resistance to a wide range of insecticides and biological toxins. Our findings provide a solid foundation for future functional studies on specific ABC transporter genes in P. xylostella, and for further understanding of their physiological roles and regulatory pathways in insecticide resistance.Electronic supplementary materialThe online version of this article (doi:10.1186/s12864-016-3096-1) contains supplementary material, which is available to authorized users.

Immune modulation enables a specialist insect to benefit from antibacterial withanolides in its host plant.

The development of novel plant chemical defenses and counter adaptations by herbivorous insect could continually drive speciation, producing more insect specialists than generalists. One approach to test this hypothesis is to compare closely related generalist and specialist species to reveal the associated costs and benefits of these different adaptive strategies. We use the specialized moth Heliothis subflexa, which feeds exclusively on plants in the genus Physalis, and its close generalist relative H. virescens. Specialization on Physalis plants necessitates the ability to tolerate withanolides, the secondary metabolites of Physalis species that are known to have feeding deterrent and immune inhibiting properties for other insects. Here we find that only H. subflexa benefits from the antibacterial properties of withanolides, and thereby gains a higher tolerance of the pathogen Bacillus thuringiensis. We argue that the specialization in H. subflexa has been guided to a large extent by a unique role of plant chemistry on ecological immunology.

When host-plant resistance to a pest leads to higher plant damage

The effects of the association between grasses and fungal endophytes on orthopterans are very poorly studied although they are important grassland pests. Here, the endemic New Zealand weta, Hemiandrus sp. ‘promontorius’, and Festulolium loliaceum infected with Epichloe uncinata, were used to study the effect of endophyte-mediated resistance in grasses on this large orthopteran insect in the laboratory, and the effects of this interaction on the grass. The insect was presented with F. loliaceum with and without E. uncinata infection in no-choice and paired choice experiments. Other controls were Epichloe festucae-infected Festuca rubra and endophyte-free Lolium perenne. In no-choice experiments, persistent attempts by the insect to graze the endophyte-infected grasses (but promptly abandoning them) resulted in a significantly higher number of plants lost due to excision at their stems after the first bite (P = 0.004). The inability of affected grasses to compensate for the lost biomass resulted in a lack of significant difference between the dry biomass of endophyte-infected and endophyte-free controls (P = 0.206). However, in choice experiments, there was a preference for the endophyte-free controls when they were paired with the endophyte-infected grasses (P < 0.05). The current work shows that endophyte-infected grasses can sustain high plant losses when attacked by an orthopteran insect in the absence of an alternative food source. This contrasts other endophyte/herbivory experiments in which high herbivory occurs because chemical plant defences are at a low concentration or the endophytes have other non-toxin roles in the plant.

Behavioral and morphological responses of an insect herbivore to low nutrient quality are inhibited by plant chemical defenses

Animals have several strategies to contend with nutritionally poor diets, including compensatory consumption and enhanced food utilization efficiencies. Plants produce a diversity of defense compounds that affect the ability of herbivores to utilize these strategies in response to variation in food nutritional quality. Little is known, however, about effects of allelochemicals on herbivores utilizing integrated behavioral and morphological responses to reduced food quality. Our objectives were to (1) examine how variation in diet nutritional quality influences compensatory responses of a generalist insect herbivore, and (2) determine how plant defenses affect these processes. Gypsy moth (Lymantria dispar) larvae were administered one of nine combinations of diet having low, moderate, or high nutritional quality and 0, 2, or 4 % purified aspen (Populus tremuloides) salicinoids. We quantified larval growth, consumption, frass production, and biomass allocation to midgut tissue over a 4-day bioassay. In the absence of salicinoids, larvae compensated for reduced nutritional quality and maintained similar growth across all diets through increased consumption, altered midgut biomass allocation, and improved processing efficiencies. Dietary salicinoids reduced larval consumption, midgut biomass allocation, digestive efficiencies, and growth at all nutritional levels, but the effect size was more pronounced when larvae were fed nutritionally suboptimal diets. Our findings demonstrate that integrated behavioral and morphological compensatory responses to reduced food quality are affected by plant defenses, ultimately limiting compensatory responses and reducing larval performance.

Folivory with leaf folding by giant flying squirrels: its patterns and possible function

AbstractJapanese giant flying squirrels, Petaurista leucogenys, are entirely arboreal folivores. From spring to summer, their principal food source consists of leaves of the deciduous oak Quercus acutissima in western Tokyo, Japan. Before consuming the central part of each leaf, the squirrels fold the leaf two or more times with their forelegs. In the wild, leaf folding is a very rare behavior, even in primates. In our study, this peculiar feeding behavior was not observed across all study sites or even in some local populations. Herbivores generally try to maximize their intake of nutritious foods (e.g., sugars) containing low levels of plant defensive chemicals (e.g., phenols). We found that total phenolic contents in individual leaves of Q. acutissima were lower in the center than at the margins, whereas sugar (glucose) was homogenously distributed. Consequently, consumption of only leaf centers appears to be an adaptation by Japanese giant flying squirrels to avoid leaf margin defensive chemicals and structures. These leaf margin characteristics, which may have developed as defenses against herbivorous insects that generally feed from the margins, have contributed to the skillful and complex feeding behavior of this generalist folivore.

The influence of plant defensive chemicals, diet composition, and winter severity on the nutritional condition of a free‐ranging, generalist herbivore

When consuming plants, herbivores must deal with both low nutritional quality from cell wall constituents and potentially toxic plant secondary metabolites, which are often inversely related. Herbivores that consume a highly nutritious, but chemically defended plant, may consume high levels of toxins that require energy for detoxification. Alternatively, herbivores may avoid consuming high levels of toxins by consuming a diverse diet that may be lower in overall nutritional quality. In this study, we assessed the relationship among nutritional restriction, detoxification and diet diversity in a free‐ranging wild herbivore. We collected urine deposited in the snow (hereafter, snow‐urine) and feces by free‐ranging moose Alces americanus, a generalist browser, during winter. We used the ratio of urinary urea nitrogen to creatinine (UN:C), measured in snow‐urine samples, as an indicator of nutritional restriction, and the ratio of glucuronic acid to creatinine (GA:C), as an indicator of investment in detoxification. We used microhistology to determine diet composition from fecal pellets. GA:C and UN:C were positively associated, suggesting that nutritional condition tends to be worse for individuals investing more in detoxification. We found, after accounting for the influence of winter severity, diet diversity and UN:C to be negatively related, suggesting that increasingly diverse diets were associated with improved nutritional condition. Overall, the most important predictor of UN:C was winter severity and proportion of diet comprised of balsam fir Abies balsamea. Physiological indicators of nutritional restriction tended to be worse during severe winters and among individuals that had consumed more balsam fir. These results highlight complex relationships among environmental conditions, foraging decisions, and costs of detoxification that can influence nutritional condition of herbivores.

Effects of plant defenses and water availability on seasonal foraging preferences of the Nubian Ibex (Capra nubiana)

The study of herbivore patch use has implications for herbivore habitat quality assessment, foraging behaviors, species interactions, and coexistence in patchy environments. This research focuses on the comparison of the effects of two qualitatively different plant defenses, mechanical (thorns) and chemical (tannins), on ibex foraging preferences during different seasons of the year. The occurrence of both chemical and mechanical plant defenses were experimentally manipulated in artificial resource patches, in addition to water availability. Ibex foraging preferences were quantified using giving-up densities during four separate fieldwork sessions in each of the seasons of the year at cliff sites overlooking the Zin Valley of the Negev Highlands. Both mechanical and chemical plant defenses significantly hindered ibex food intake overall. Mechanical and chemical defenses acted as substitutable defenses, meaning that their combined effects were not greater than additive. There were strong seasonal patterns of the amount of food consumed by ibex, further corroborated by comparison to rainfall levels. Seasonality also interacted with the effectiveness of plant defenses. Thorns were especially ineffective in summer, whereas tannins were most effective in spring. Decreases in seasonal food availability and increased marginal value of energy for ibex may have resulted in thorn ineffectiveness, while seasonal changes in the emergence of young foliage may have resulted in the greater springtime tannin effectiveness. Water was not found to mitigate the detrimental effects of tannins through dilution. The implications for decreased constraints on selective pressures on ibex due to the substitutability of plant defenses are discussed.

Does plant apparency matter? Thirty years of data provide limited support but reveal clear patterns of the effects of plant chemistry on herbivores.

According to the plant-apparency hypothesis, apparent plants allocate resources to quantitative defenses that negatively affect generalist and specialist herbivores, while unapparent plants invest more in qualitative defenses that negatively affect nonadapted generalists. Although this hypothesis has provided a useful framework for understanding the evolution of plant chemical defense, there are many inconsistencies surrounding associated predictions, and it has been heavily criticized and deemed obsolete. We used a hierarchical Bayesian meta-analysis model to test whether defenses from apparent and unapparent plants differ in their effects on herbivores. We collected a total of 225 effect sizes from 158 published papers in which the effects of plant chemistry on herbivore performance were reported. As predicted by the plant-apparency hypothesis, we found a prevalence of quantitative defenses in woody plants and qualitative defenses in herbaceous plants. However, the detrimental impacts of qualitative defenses were more effective against specialists than generalists, and the effects of chemical defenses did not significantly differ between specialists and generalists for woody or herbaceous plants. A striking pattern that emerged from our data was a pervasiveness of beneficial effects of secondary metabolites on herbivore performance, especially generalists. This pattern provides evidence that herbivores are evolving effective counteradaptations to putative plant defenses.

Arbuscular mycorrhizal fungi affect plant tolerance and chemical defences to herbivory through different mechanisms

Summary Arbuscular mycorrhizal fungi (AMF) are ubiquitous plant symbionts that affect plant nutrient status, patterns of resource allocation and rates of plant growth. In addition, AMF influence the expression of plant defence, which may affect subsequent interactions between plants and higher trophic levels. Tolerance to herbivory and chemical defence represent two distinct defence strategies of plants, and there is accumulating evidence that AMF can influence each strategy separately. However, investigations on the simultaneous effects of AMF on tolerance and chemical defence mechanisms are lacking. Using six milkweed (Asclepias) species and three levels of AMF inoculum, we show that plant tolerance to herbivory increases with foliar P concentrations, while both foliar cardenolide concentration and latex exudation (major chemical defences in milkweeds) increase with foliar N concentrations, but decrease with plant growth rate. Additionally, foliar cardenolide concentration increases with foliar P concentration, and latex exudation increases with root biomass. Because AMF have significant effects on all of these traits (foliar N and P concentrations, root biomass and growth rate), AMF can change plant tolerance and chemical defence simultaneously, albeit through different mechanisms. Synthesis. Overall, plant tolerance and chemical defence to herbivores are affected by AMF through changes in basic plant physiology such as nutrient status, allocation patterns and growth rate, and our study suggests that it may be possible to predict effects of AMF on plant defences through these simple traits.

Detection and Adaptation in Parasitic Angiosperm Host Selection

Developmental transitions in some parasitic angiosperms are tied directly to host-derived chemical cues (xenognosins). The obligate hemi-parasite Striga asiatica, initiates the root apical meristem population (germination), development of the host attachment organ (the haustorium), and shoot apical meristem initiation (seed coat shedding) in response to specific xengonosins. These checkpoints synchronize spatial and temporal tissue development. We have now exploited the external control over these developmental transitions to trace functional expression in haustorial organogenesis. Genes associated with phytohormone regulation, metabolism, vascular tissue development, and reactive oxygen species (ROS) production identified in this study suggest an elaborate and global response closely tied to plant defense and redox chemistry that may also be components of a more general quorum sensing-type mechanism in plants.