Modelling tritrophic interactions mediated by induced defence volatiles

Many plant species defend themselves against herbivorous insects indirectly by producing and releasing induced volatiles to attract natural enemies of the herbivores. In this paper, we consider the recruitment of natural enemies attracted by plant-induced volatiles and introduce the An–Liu–Johnson–Lovett model into the Lotka–Volterra model in an attempt to add this missing vital link in tritrophic interaction. Increase in attraction strength of plant-induced volatiles to the natural enemy leads to high fluctuation amplitude of plant biomass and herbivore population. When the attack strength of natural enemies reaches a certain level, fluctuation amplitude of plant biomass and herbivore population will decrease and plant biomass will approach to its environmental carrying capacity. The simulation demonstrates that plant volatile compounds induced by insects have led to the introduction of a third tritrophic level, e.g., natural enemies, into the plant–herbivore system, resulting in the coexistence of plants, insects, and natural enemies during the evolution process.     

Herbivore species, infestation time, and herbivore density affect induced volatiles in tea plants

In response to insect herbivory, plants emit volatiles that are used by the herbivores’ natural enemies to locate their host or prey. Herbivore attack also enhances tea aroma. Herbivore-induced plant volatiles (HIPVs) vary both quantitatively and qualitatively with infestation duration and herbivore density. Thus, whether HIPVs can reliably communicate the identities of herbivores is of interest. Here, we studied the tea plant volatiles induced by the tea leafhopper (Empoasca vitis, a piercing–sucking insect), the tea geometrid (Ectropis oblique, a chewing insect), and methyl jasmonate (MeJA, a plant hormone). Geometrid feeding induced more complex volatile blends than did leafhopper infestation. The volatiles induced by both herbivores significantly increased in quality and quantity with time during the first 16 h of infestation, after which the profiles of induced volatile blends and the emission of induced compounds varied diurnally. (E)-β-Ocimene displayed a unique rhythm in which emission peaked at night. The amount of HIPVs significantly increased, while their profiles changed little, with herbivore density. Overall, the leafhoppers and geometrids induced significantly different volatiles from tea plants, while the HIPV profiles varied with a circadian rhythm and were similar at different herbivore densities. Our findings also suggest a new method of enhancing tea flavor using exogenously applied plant hormones, because the volatiles induced by leafhoppers and MeJA were similar in general composition.     

Mass scans from a proton transfer mass spectrometry analysis of air over Mediterranean shrubland browsed by horses

Plants usually emit large amount and varieties of volatiles after being damaged by herbivores. However, analytical methods for measuring herbivore-induced volatiles do not normally monitor the whole range of volatiles and the response to large herbivores such as large mammals is much less studied than the response to other herbivores such as insects. In this paper we present the results of using a highly sensitive proton transfer reaction-mass spectrometry (PTR-MS) technique that allows simultaneous monitoring of leaf volatiles in the pptv range. The resulting mass scans in air over Mediterranean shrubland browsed by horses show 70 to 100% higher concentrations of the masses corresponding to mass fragments 57, 43 and 41 (mostly hexenals, acetone and acetic acid) than scans over control non-browsed shrubland. These compounds are biogeochemically active and they are significant components of the volatile organic carbon found in the atmosphere. They influence the performance of living organisms and, the chemical and physical processes of Earth's atmosphere.     

Why is latex usually white and only sometimes yellow,orange or red? Simultaneous visual and chemical plant defense

Latex is a toxic and sticky defense substance produced by about 20,000 plant species and secreted following wounding. It defends the wounded plants by both chemical and physical components. A striking fact about latex is that the majority of plants produce it white, although yellow, orange, and red latex can sometimes be found. Theoretically, it is possible that there is no function or importance to latex’s color, but it seems unlikely. It is also possible that there are chemical constraints that dictate the production of white latex, but the various colors indicate that this is not a valid explanation. However, since white latex evolved independently many times in various plant taxa, there should have been a strong selection for latex being white, a possibility that has never been addressed, to my knowledge. I propose that latex is generally white because white is the best solution for visual aposematism under the lighted conditions within and under plant canopies, on the background of typical leaf and bark color, even towards color-blind animals. Yellow, orange, and red latex types should naturally also be considered as aposematic because these are typical colors of toxic and aposematic organisms. In addition, latex-exuding plants simultaneously use a chemically based, non-visual olfactory aposematism because wounding perse also causes the emission of non-latex defensive volatiles and because latex contains various but little-studied defensive volatiles. The volatile defensive aspect of latex exudation may also operate against nocturnal or blind herbivores.     

Adaptation of a four-arm olfactometer for behavioural bioassays of large beetles

Summary. We adapted a four-arm olfactometer (55 × 55 × 5.5 cm) for bioassays of large insects and its usefulness was evaluated by testing the responses of three beetles, the palm weevil Rhynchophorus palmarum (L.) (Coleoptera: Curculionidea) and two Dynast scarabs Strategus aloeus (L.) and Oryctes rhinoceros (L.) (Coleoptera: Scarabaeidae) to plant odours and aggregation pheromones. This olfactometer was coupled to a volatile delivery system dispensing highly volatile semiochemicals at constant concentrations. We materialized airflows in the olfactometer by discolouring sensitive papers with acid vapours under various conditions (air straighten by stainless steel grids in inputs and output, 500 ml/min per arm) to visualize air turbulence and to test air-tightness. The volatile delivery system can be used to mix and dispense a broad range of concentrations of semiochemicals in air from diluted aqueous solutions. It was calibrated by measuring the release of the pheromone using solid phase-microextraction (SPME). Dose-response curves to synthetic pheromone were obtained for O. rhinoceros of both sexes. Coupling the volatile delivery system to the fourarm olfactometer provided a system with which the responses of R. palmarum and S. aloeus to their synthetic aggregation pheromones and to natural plant volatiles could be readily observed and studied.     

Associations of plant fitness, leaf chemistry, and damage suggest selection mosaic in plant-herbivore interactions

The geographic mosaic theory of coevolution states that variation in species interactions forms the raw material for coevolutionary processes, which take place over large geographic scales. One key assumption underlying the process of coevolution in plant-herbivore interactions is that herbivores exert selection on their host plants and that this selection varies among plant populations. We examined spatial variation in the existence and strength of phenotypic selection on host plant resistance exerted by specialist herbivores in 17 archipelago populations of the perennial herb Vincetoxicum hirundinaria (Asclepiadaceae). In these highly fragmented populations, V. hirundinaria is consumed by the larvae of two specialist herbivores: the folivorous moth Abrostola asclepiadis and the seed predator Euphranta connexa. Selection imposed on host plants by these herbivores was examined by analyzing the associations between levels of herbivory, plant fitness, and contents of a number of leaf chemicals reflecting plant resistance to and quality for the herbivores. We found extensive spatial variation in the levels of herbivory and in plant fitness. More importantly, the impact of both leaf herbivory and seed predation on plant fitness varied among plant populations, indicating spatial variation in phenotypic selection. In addition, leaf chemistry varied widely among plant populations, reflecting spatial variation in plant quality as food for the herbivores. However, leaf compounds influenced folivory similarly in all the studied plant populations, and interestingly, some of the compounds were associated with the intensity of seed predation. Finally, some of the leaf compounds were associated with plant fitness, and the strength and direction of these associations varied among plant populations. The observed spatial variation in the strength of the interactions between V. hirundinaria and its specialist herbivores suggests a geographic selection mosaic. Because the occurrence and strength of spatial variation varied between the two specialist herbivores, our results highlight the importance of considering multiple enemies when trying to understand evolution of interactions between plants and their herbivores.     

Volatile organic compounds and host-plant specialization in European corn borer E and Z pheromone races

Plant volatile cues are considered the main source of information for ovipositing moths, which use chemical information to locate and recognize the host plant. In Europe, two sympatric populations of European corn borer (ECB; Ostrinia nubilalis, Hübner), the Z and E-pheromone races, feed mainly on maize and hop or mugwort, respectively. We studied the mechanisms of host-plant recognition and fidelity in ECB pheromone races by testing the attractiveness of host plants to gravid females in a flight tunnel and by analyzing the volatiles released from maize, mugwort, and hop during the scotophase, when the ovipositing flight of the ECB females occurs. In the wind tunnel bioassay, the Z-race and E-race females engaged in upwind flight and expressed a strong host fidelity to their respective main host plants; all three of these host plants possess distinctive volatile profiles specific as to blend and ratio. The host plants shared a certain number of ubiquitous volatiles present in various ratios that likely constitute a species-specific cue to host-seeking ECB moths. Our observations therefore suggest that ECB host fidelity is steered by plant volatiles that are present in species-specific ratios of ubiquitous volatile organic compounds.     

Predispersal seed herbivores, not pollinators, exert selection on floral traits via female fitness

Herbivores that oviposit in flowers of animal-pollinated plants depend on pollinators for seed production and are therefore expected to choose flowers that attract pollinators. This provides a mechanism by which seed herbivores and pollinators could impose conflicting selection on floral traits. We measured phenotypic selection on floral traits of Lobelia siphilitica (Lobeliaceae) via female fitness to determine the relative strength of selection by pollinators and a specialist predispersal seed herbivore. We were able to attribute selection on flowering phenology to the herbivores. However, no selection could be attributed to pollinators, resulting in no conflicting selection on floral traits. Unlike pollinators, whose preference for certain floral traits does not always translate into higher fitness, any discrimination by seed herbivores is likely to decrease fitness of the preferred floral phenotype. Thus predispersal seed herbivores may be a significant agent of selection on floral traits.     

Ontogenetic and spatio-temporal patterns of induced volatiles in <Emphasis Type="Italic">Glycine max</Emphasis> in the light of the optimal defence hypothesis

Summary. Plants attacked by herbivorous insects emit a blend of volatile compounds that serve as important host location cues for parasitoid wasps. Variability in the released blend may exist on the whole-plant and withinplant level and can affect the foraging efficiency of parasitoids. We comprehensively assessed the kinetics of herbivore-induced volatiles in soybean in the context of growth stage, plant organ, leaf age, and direction of signal transport. The observed patterns were used to test the predictions of the optimal defence hypothesis (OD). We found that plants in the vegetative stage emitted 10-fold more volatiles per biomass than reproductive plants and young leaves emitted >2.6 times more volatiles than old leaves. Systemic induction in single leaves was stronger and faster by one day in acropetal than in basipetal direction while no systemic induction was found in pods. Herbivore-damaged leaves had a 200-fold higher release rate than pods. To some extent these findings support the OD: i) indirect defence levels were increased in response to herbivory and ii) young leaves, which are more valuable, emitted more volatiles. However, the fact that reproductive structures emitted no constitutive or very few inducible volatiles is in seeming contrast to the OD predictions. We argue that in case of volatile emission the OD can only partially explain the patterns of defence allocation due to the peculiarity that volatiles act as signals not as toxins or repellents.     

A review of the chemical ecology of the Cerambycidae (Coleoptera)

Summary. This review summarizes the literature related to the chemical ecology of the Cerambycidae and provides a brief overview of cerambycid biology, ecology, economic significance, and management. Beetles in the family Cerambycidae have assumed increasing prominence as pests of forest and shade trees, shrubs, and raw wood products, and as vectors of tree diseases. Exotic species associated with solid wood packing materials have been notable tree killers in North American urban and peri-urban forests. In forested ecosystems native species respond to disturbances such as fires and windstorms, and initiate the biodeterioration of woody tissue. Eggs are laid by females, on or through the bark surface of stem and branch tissue of moribund, recently killed or decomposing woody plants; larval cerambycids (roundheaded woodborers) typically feed in the phloem and later in the xylem. Females will also, in some cases, select living hosts, e.g. adult conifer and angiosperm trees, for oviposition. Research on the chemical ecology of over 70 species has revealed many examples of attractive kairomones (such as floral volatiles, smoke volatiles, trunk and leaf volatiles, and bark beetle pheromones), repellents and deterrents, oviposition stimulants, short- and long-range sex pheromones, and defensive substances. Emerging generalities are that attractants tend to be monoterpenoids and phenolic esters; oviposition stimulants are monoterpenoids and flavanoids; short-range sex pheromones are female-produced, methyl-branched cuticular hydrocarbons; and long-range sex pheromones are male-produced -hydroxy ketones and (,)-diols ranging in length from 6 to 10 carbons. The latter compounds appear to originate from glands in the male thorax; putative defensive substances originate from metasternal secretory pores or mandibular glands. In one unusual case, a flightless, subterranean female that attacks sugar cane produces a sex pheromone that is derived from the amino acid isoleucine. With significantly more than 35,000 species of Cerambycidae worldwide, these generalities will be subject to change as more species are examined. Addendum The authors would like to point out that the electronic version is more accurate than the printed version.     

Herbivory-mediated pollinator limitation: negative impacts of induced volatiles on plant-pollinator interactions

Although induced plant responses to herbivory are well studied as mechanisms of resistance, how induction shapes community interactions and ultimately plant fitness is still relatively unknown. Using a wild tomato, Solanum peruvianum, native to the Peruvian Andes, we evaluated the disruption of pollination as a potential ecological cost of induced responses. More specifically, we tested the hypothesis that metabolic changes in herbivore-attacked plants, such as the herbivore-induced emission of volatile organic compounds (VOCs), alter pollinator behavior and consequentially affect plant fitness. We conducted a series of manipulative field experiments to evaluate the role of herbivore-induced vegetative and floral VOC emissions as mechanisms by which herbivory affects pollinator behavior. In field surveys and bioassays in the plants' native habitat, we found that real and simulated herbivory (methyl jasmonate application) reduced attractiveness of S. peruvianum flowers to their native pollinators. We show that reduced pollinator preference, not resource limitation due to leaf tissue removal, resulted in reduced seed set. Solitary bee pollinators use floral plant volatiles, emitted in response to herbivory or methyl jasmonate treatment, as cues to avoid inflorescences on damaged plants. This herbivory-induced pollinator limitation can be viewed as a general cost of induced plant responses as well as a specific cost of herbivory-induced volatile emission.     

The semiochemically mediated interactions between bacteria and insects

In natural environment, semiochemicals are involved in many interactions between the different trophic levels involving insects, plants and hosts for parasitoids or prey for predators. These volatile compounds act as messengers within or between insect species, inducing particular behaviours, such as the localisation of a source of food, the orientation to an adequate oviposition site, the selection of a suitable breeding site and the localisation of hosts or prey. In this sense, bacteria have been shown to play an important role in the production of volatile compounds which ones act as semiochemicals. This review, focusing on the semiochemically mediated interactions between bacteria and insects, highlights that bacterial semiochemicals act as important messengers for insects. Indeed, in most of the studies reported here, insects respond to specific volatiles emitted by specific bacteria hosted by the insect itself (gut, mouthparts, etc.) or present in the natural environment where the insect evolves. Particularly, bacteria from the families Enterobacteriaceae, Pseudomonaceae and Bacillaceae are involved in many interactions with insects. Because semiochemicals naturally produced by bacteria could be a very interesting option for pest management, advances in this field are discussed in the context of biological control against insect pests.     

Bottom-up multitrophic effects in resprouting plants

Severe damage often provokes compensatory resprouting of plants, which commonly modify plant morphological and phenological traits. Rapid plant growth often results in poorly defended nutrient-rich foliage, which is more susceptible to foliar-chewing herbivores. It is less known how other guilds of arthropods are affected by plant regrowth. We tested the hypotheses that clipping-induced resprouting and nutrient availability, separately and in combination, would (1) influence plant traits, (2) benefit chewing herbivores, sap-suckers, gallers, and pre-dispersal seed predators, and (3) cascade up to the third trophic level by positively affecting herbivores. Resprouted plants were morphologically and phenologically different from undamaged plants; as a result, seed predation, infestation rate, richness, and diversity of seed predators increased, and species composition was altered. Leaf consumption by chewing herbivores was four times higher on resprouted plants. The number of galls decreased, whereas the abundance of sap-sucking and leaf-chewing insects was not affected. The incidence of predators and parasitoids was also higher on resprouted plants and on plants with nutrients added, but the increase was less pronounced compared to the herbivores they feed on. Thus, the effects of resprouting, contingent on nutrient availability, can propagate simultaneously through two independent tri-trophic level pathways.     

Local Species Richness of Leaf-Chewing Insects Feeding on Woody Plants from One Hectare of a Lowland Rainforest

Abstract:  Local species diversity of insect herbivores feeding on rainforest vegetation remains poorly known. This ignorance limits evaluation of species extinction patterns following various deforestation scenarios. We studied leaf-chewing insects feeding on 59 species of woody plants from 39 genera and 18 families in a lowland rainforest in Papua New Guinea and surveyed all plants with a stem diameter at breast height of ≥5 cm in a 1-ha plot within the same area. We used two extrapolation methods, based on randomized species-accumulation curves, to combine these two data sets and estimate the number of species of leaf-chewing herbivores feeding on woody plants from the 1-ha area. We recorded 58,483 feeding individuals from 940 species of leaf-chewing insects. The extrapolation estimated that there were 1567–2559 species of leaf-chewing herbivores feeding on the 152 plant species from 97 genera and 45 families found in 1 ha of the forest. Most of the herbivore diversity was associated with plant diversity on the familial and generic levels. We predicted that, on average, the selection of 45 plant species each representing a different family supported 39% of all herbivore species, the 52 plant species each representing a different additional genus from these families supported another 39% of herbivore species, and the remaining 55 plant species from these genera supported 22% of herbivore species. Lepidoptera was the most speciose taxon in the local fauna, followed by Coleoptera and orthopteroids (Orthoptera and Phasmatodea). The ratio of herbivore to plant species and the estimated relative species richness of the Lepidoptera, Coleoptera, and orthopteroids remained constant on the spatial scale from 0.25 to 1 ha. However, the utility of local taxon-to-taxon species ratios for extrapolations to geographic scales requires further study.     

Is flower scent influencing host plant selection of leaf-galling sawflies (Hymenoptera, Tenthredinidae) on willows?

Though it is known that flower scent not only attracts pollinators but also herbivores, little is known about the importance of flower scent on the distribution of leaf herbivores among individuals within a plant species. In this study we determined the distribution of galls induced by the sawfly Pontania proxima (Serville 1823) (Hymenoptera, Tenthredinidae, Nematinae) on flowering and non-flowering representatives of several clones belonging to Salix fragilis and S. × rubens (Salicaceae). Further, amounts and composition of scent of flowering and non-flowering twigs were compared (dynamic headspace-gas chromatography–mass spectrometry, DHS-GC–MS), and a scent sample collected from flowering twigs of S. fragilis was tested by coupled gas chromatography and electroantennographic detection (GC-EAD) on the antennae of P. proxima females. The results show that the presence of flower catkins on plants led to a higher degree of allocation with galls, but the number of galls differed not between flowering and non-flowering plants. The DHS-GC–MS analyses revealed that the total amount of flower scent emitted per flowering twig is about 90 times higher than the scent emitted by a non-flowering twig. Further, several compounds were emitted only by flowering but not by non-flowering twigs. In the GC-EAD analyses, antennae consistently responded not only to green leaf volatiles, but also to compounds emitted only by the flowers (e.g. 1,4-dimethoxybenzene). These flower scent compounds are suggested to affect the host plant choice by attracting more sawflies from the distance to flowering plants compared to non-flowering plants. The EAD-active compounds emitted from vegetative plant parts are assumed to act as long-distance signals especially when flowers are absent on host plants, e.g. during the oviposition period of the second generation of P. proxima.     

Herbivore-induced "rent rise" in the host plant may drive a diet breadth enlargement in the tenant

Inter- and intraspecies variations in host plant traits are presumably involved in many host shifts by insect herbivores, and elucidating the mechanisms involved in such shifts has been a crucial goal in insect-plant research for several decades. Here we propose that herbivore-induced evolutionary increases in host plant resistance may cause oligophagous insect herbivores to shift to other sympatric plants as currently preferred host plants become increasingly unpalatable. We tested this hypothesis in a system based on the perennial herb Filipendula ulmaria (Rosaceae), whose herbivory defense has become gradually stronger due to prolonged selection by Galerucella tenella (Coleoptera: Chrysomelidae) herbivory in a boreal archipelago. We show that Galerucella gradually increases its use of the alternative host plant Rubus arcticus (Rosaceae) in parallel to gradually increased resistance in Filipendula. Our results imply that, by driving the evolutionary increase in Filipendula resistance, Galerucella is also gradually making the original host species more unpalatable and thereby driving its own host-breadth enlargement. We argue that such self-inflicted "rent rises" may be an important mechanism behind host plant shifts, which in turn are believed to have preceded the speciation of many phytophagous insects.     

Variation in composition of predator-attracting allelochemicals emitted by herbivore-infested plants: Relative influence of plant and herbivore

Summary During foraging, natural enemies of herbivores may employ volatile allelochemicals that originate from an interaction of the herbivore and its host plant. The composition of allelochemical blends emitted by herbivore-infested plants is known to be affected by both the herbivore and the plant. Our chemical data add new evidence to the recent notion that the plants are more important than the herbivore in affecting the composition of the volatile blends. Blends emitted by apple leaves infested with spider mites of 2 different species,T. urticae andP. ulmi, differed less in composition (principally quantitative differences for some compounds) than blends emitted by leaves of two apple cultivars infested by the same spider-mite species,T. urticae (many quantitative and a few qualitative differences). Comparison between three plant species — apple, cucumber and Lima bean — reveals even larger differences between volatile blends emitted upon spider-mite damage (many quantitative differences and several qualitative differences).     

Are lower-latitude plants better defended? Palatability of freshwater macrophytes

Increased herbivory at lower latitudes is hypothesized to select for more effective plant defenses. Feeding assays with seaweeds and salt marsh plants support this hypothesis, with low-latitude plants experiencing greater damage in the field and being less palatable than higher-latitude plants. We tested this hypothesis for freshwater macrophytes because they offered an independent plant lineage and habitat type for testing this general hypothesis and because the patchiness of consumer occupancy across isolated water bodies might produce local variance in herbivory that would override geographic variance and produce different results for this habitat type. When we fed eight congeneric pairs of live plants from four sites in Indiana vs. four sites in South Florida (-215 and 0 frost days/yr respectively) to three species of crayfishes and one species of snail, three of the four herbivores significantly preferred high-latitude to low-latitude plants. For two crayfishes that differed in feeding on live plants (one favoring high-latitude plants and one not), we retested feeding using foods composed of freeze-dried and finely ground plants, thus removing structural characteristics while retaining most chemical/nutritional traits. In this assay, both herbivores strongly preferred high-latitude plants, suggesting that lower-latitude plants had been selected for more deterrent chemical traits. When we collected 22 pairs of congeneric plants from 9 sites throughout Indiana vs. 13 sites in Central Florida (-215 and -95 frost days/yr respectively) and tested these in feeding assays with three crayfishes using dried, ground, and reconstituted plant material, we found a significant effect of latitude for only one of three species of herbivore. Overall, our results suggest a preference for high-latitude plants, but the strength of this relationship varied considerably across small scales of latitude that differed considerably in numbers of frost-free days. The difference in results suggests that large changes in frost frequency over small spatial scales may affect selection for plant defenses, that local variance in herbivory overrode differential selection at geographic scales, or that these possibilities interact when durations of cold weather periodically exclude herbivores from shallower habitats, producing heterogeneous selection for defenses at small spatial scales.     

Maize field odorscape during the oviposition flight of the European corn borer

Most crop pests find a suitable host through chemical cues released from plants, but little is known about the odorscape encountered by host-seeking gravid females under natural, outdoor conditions. In this field study, the volatile organic compound (VOC) composition of maize (Zea mays, L.), a host for the European corn borer (ECB) (Ostrinia nubilalis Hüb.) was characterized during the oviposition flight and compared with a forest odorscape. VOCs from maize fields and the forest atmosphere were collected by solid phase microextraction and characterized by gas chromatography-mass spectrometry. The electroantennographic (EAG) response of female ECB antennae to candidate VOCs was tested. Analyses revealed clear differences between the maize field and the forest odorscapes, mainly composed of ubiquitous VOCs but in specific ratios. The maize field odorscape is more complex than the forest odorscape for maize found 18 VOCs but only eight in the forest. Both biotopes shared seven VOCs—green leaf volatiles (GLV), monoterpènes (MT) and homoterpenes. In addition, we found in the forest a distinctive sesquiterpene (SQT) identified as isoledene. The highest EAG responses were elicited by two GLVs and a MT shared by the two biotopes. SQT elicited weak EAG responses, except β-farnesene, only found in the maize field odorscape. Our results suggest that the two biotopes produce specific chemical signatures that insects may use as host cues. To the best of our knowledge this paper is the first report on the maize odorscapes under field conditions. The putative role of the VOCs in host plant detection and selection is discussed.     

Making scents of defense: do fecal shields and herbivore-caused volatiles match host plant chemical profiles?

Many plant families have aromatic species that produce volatile compounds which they release when damaged, particularly after suffering herbivory. Monarda fistulosa (Lamiaceae) makes and stores volatile essential oils in peltate glandular trichomes on leaf and floral surfaces. This study examined the larvae of a specialist tortoise beetle, Physonota unipunctata, which feed on two M. fistulosa chemotypes and incorporate host compounds into fecal shields, structures related to defense. Comparisons of shield and host leaf chemistry showed differences between chemotypes and structures (leaves vs. shields). Thymol chemotype leaves and shields contained more of all compounds that differed than did carvacrol chemotypes, except for carvacrol. Shields had lower levels of most of the more volatile chemicals than leaves, but more than twice the amounts of the phenolic monoterpenes thymol and carvacrol and greater totals. Additional experiments measured the volatiles emitted from M. fistulosa in the absence and presence of P. unipunctata larvae and compared the flower and foliage chemistry of plants from these experiments. Flowers contained lower or equal amounts of most compounds and half the total amount, compared to leaves. Plants subjected to herbivory emitted higher levels of most volatiles and 12 times the total amount, versus controls with no larvae, including proportionally more of the low boiling point chemicals. Thus, chemical profiles of shields and volatile emissions are influenced by the amounts and volatilities of compounds present in the host plant. The implications of these results are explored for the chemical ecology of both the plant and the insect.