Glandular trichomes as an inflorescence defence mechanism against insect herbivores in Iberian columbines

Glandular trichomes play a defensive role against herbivores in the leaves of many plant species. However, their functional role in inflorescences has not been studied, even though theory suggests that tissues with a higher fitness value, such as inflorescences, should be better defended. Using manipulative experiments, we analysed the defensive role of glandular trichomes against herbivorous insects in the inflorescence of Iberian columbines (genus Aquilegia), and its inter-population and inter-taxa variation in relation to herbivore abundance and potential selective pressure. The experiments were conducted in eight populations belonging to four subspecies of two columbines (Aquilegia vulgaris and Aquilegia pyrenaica). For each population, we estimated the density of glandular trichomes in the inflorescences, the abundance of insects stuck in the inflorescences, the abundance of small herbivorous insects, the incidence of damage on flowers and fruits, and the fruit set. The density of glandular trichomes on the inflorescence of A. vulgaris and A. pyrenaica was higher in regions of higher herbivore abundance. We also found that when the plants lose the protection of glandular trichomes, small insects have better access to flowers and fruits, causing more damage and reducing plant fitness. This study concludes that glandular trichomes are part of an adaptive response against phytophagous insect herbivory. The observed variation in herbivore pressure between taxa, likely caused by habitat differentiation, might have played a role in trait differentiation through divergent selection. This result adds evidence to the differentiation of the Iberian columbines through habitat specialization.     

Selection Mosaic Exerted by Specialist and Generalist Herbivores on Chemical and Physical Defense of Datura stramonium

Selection exerted by herbivores is a major force driving the evolution of plant defensive characters such as leaf trichomes or secondary metabolites. However, plant defense expression is highly variable among populations and identifying the sources of this variation remains a major challenge. Plant populations are often distributed across broad geographic ranges and are exposed to different herbivore communities, ranging from generalists (that feed on diverse plant species) to specialists (that feed on a restricted group of plants). We studied eight populations of the plant Datura stramonium usually eaten by specialist or generalist herbivores, in order to examine whether the pattern of phenotypic selection on secondary compounds (atropine and scopolamine) and a physical defense (trichome density) can explain geographic variation in these traits. Following co-evolutionary theory, we evaluated whether a more derived alkaloid (scopolamine) confers higher fitness benefits than its precursor (atropine), and whether this effect differs between specialist and generalist herbivores. Our results showed consistent directional selection in almost all populations and herbivores to reduce the concentration of atropine. The most derived alkaloid (scopolamine) was favored in only one of the populations, which is dominated by a generalist herbivore. In general, the patterns of selection support the existence of a selection mosaic and accounts for the positive correlation observed between atropine concentration and plant damage by herbivores recorded in previous studies.     

Nonglandular silicified trichomes are essential for rice defense against chewing herbivores

Interspecific New Rice for Africa (NERICA) varieties have been recently developed and used in Sub-Saharan Africa but herbivore resistance properties of these plants remain poorly understood. Here we report that, compared to a local Japanese cultivar Nipponbare, NERICA 1, 4 and 10 are significantly more damaged by insect herbivores in the paddy fields. In contrast to high levels of leaf damage from rice skippers and grasshoppers, constitutive and induced volatile organic compounds for indirect plant defense were higher or similar in NERICAs and Nipponbare. Accumulation of direct defense secondary metabolites, momilactones A and B, and p-coumaroylputrescine (CoP) was reduced in NERICAs, while feruloylputrescine accumulated at similar levels in all varieties. Finally, we found that Nipponbare leaves were covered with sharp nonglandular trichomes impregnated with silicon but comparable defense structures were virtually absent in herbivory-prone NERICA plants. As damage to the larval gut membranes by Nipponbare silicified trichomes that pass intact through the insect digestive system, occurs, and larval performance is enhanced by trichome removal from otherwise chemically defended Nipponbare plants, we propose that silicified trichomes work as an important defense mechanism of rice against chewing insect herbivores.     

Tibor Jermy (1917–2014)

Throughout the course of their evolution, plants have acquired a wide range of chemical and mechanical defenses to protect against herbivores. Ehrlich & Raven's coevolutionary theory suggests that this diversification of defensive traits is driven by the strong impact of novel traits on insect herbivores. However, the impact of plant defenses on insects is difficult to compare between related plant species due to variation in environmental and biotic conditions. We standardized these factors as far as possible by analyzing the effects of chemical and mechanical defensive traits on insects in a local community of 11 Salicaceae species growing in sympatry, and their leaf‐chewing herbivores. Defensive traits (salicylates, flavonoids, tannins, trichomes, and leaf toughness) were generally not inter‐correlated, with the exception of a negative correlation between salicylates and trichomes. The content of salicylates, a novel group of defensive metabolites in the Salicaceae, was correlated with low herbivore diversity and high host specificity. Despite these effects, the phylogeny of the studied species shows loss of salicylates in some Salix species instead of their further diversification. This could be due to salicylates not decreasing the overall abundance of herbivores, despite accounting for up to 22% of the dry leaf mass and therefore being costly. The defense of low‐salicylate willow species is thus probably maintained by other defensive traits, such as trichomes. Our study shows that the balance between costs and benefits of defensive traits is not necessarily in favor of novel compounds and illustrates a process, which may lead to the reduction in a defensive trait.     

Reciprocal feeding facilitation between above- and below-ground herbivores

Interspecific interactions between insect herbivores predominantly involve asymmetric competition. By contrast, facilitation, whereby herbivory by one insect benefits another via induced plant susceptibility, is uncommon. Positive reciprocal interactions between insect herbivores are even rarer. Here, we reveal a novel case of reciprocal feeding facilitation between above-ground aphids (Amphorophora idaei) and root-feeding vine weevil larvae (Otiorhynchus sulcatus), attacking red raspberry (Rubus idaeus). Using two raspberry cultivars with varying resistance to these herbivores, we further demonstrate that feeding facilitation occurred regardless of host plant resistance. This positive reciprocal interaction operates via an, as yet, unreported mechanism. Specifically, the aphid induces compensatory growth, possibly as a prelude to greater resistance/tolerance, whereas the root herbivore causes the plant to abandon this strategy. Both herbivores may ultimately benefit from this facilitative interaction.     

Herbivore handling of a Plants trichome: the case of Heliconius charithonia (L.) (Lepidoptera: Nymphalidae) and Passiflora lobata (Killip) Hutch. (Passifloraceae)

Trichomes reduce herbivore attack on plants by physically and/or chemically inhibiting movement or other activities. Despite evidence that herbivores are negatively affected by trichomes there also reports of insect counter-adaptations that circumvent the plants defense. This paper reports on a study that investigated the likely mechanisms employed by larvae of the nymphalid butterfly, Heliconius charithonia (L.), that allow it to feed on a host that is presumably protected by hooked trichomes (Passiflora lobata (Killip) Hutch). Evidence were gathered using data from direct observations of larval movement and behavior, faeces analysis, scanning electron microscopy of plant surface and experimental analysis of larval movement on plants with and without trichomes (manually removed). The latter involved a comparison with a non specialist congener, Heliconius pachinus Salvin. Observations showed that H. charithonia larvae are capable of freeing themselves from entrapment on trichome tips by physical force. Moreover, wandering larvae lay silk mats on the trichomes and remove their tips by biting. In fact, trichome tips were found in the faeces. Experimental removal of trichomes aided in the movement of the non specialist but had no noticeable effect on the specialist larvae. These results support the suggestion that trichomes are capable of deterring a non specialist herbivore (H. pachinus). The precise mechanisms that allow the success of H. charithonia are not known, but I suggest that a blend of behavioral as well as physical resistance mechanisms is involved. Future studies should ascertain whether larval integument provides physical resistance to trichomes.     

Geographic association and temporal variation of chemical and physical defense and leaf damage in Datura stramonium

The evolution of plant defense traits has traditionally been explained trough the “coevolutionary arms race” between plants and herbivores. According to this, specialist herbivores have evolved to cope effectively with the defensive traits of their host plants and may even use them as a cue for host location. We analyzed the geographic association between leaf trichomes, two tropane alkaloids (putative resistance traits), and leaf damage by herbivores in 28 populations of Datura stramonium in central Mexico. Since the specialist leaf beetles Epitrix parvula and Lema trilineata are the main herbivores of D. stramonium in central Mexico, we predicted a positive association between plant defense and leaf damage across populations. Also, if physical environmental conditions (temperature or precipitation) constrain the expression of plant defense, then the geographic variation in leaf damage should be explained partially by the interaction between defensive traits and environmental factors. Furthermore, we studied the temporal and spatial variation in leaf trichome density and leaf damage in five selected populations of D. stramonium sampled in two periods (1997 vs. 2007). We found a positive association between leaf trichomes density and atropine concentration with leaf damage across populations. The interaction between defensive traits and water availability in each locality had a significant effect on the geographic variation in leaf damage. Differences among populations in leaf trichome density are maintained over time. Our results indicate that local plant–herbivore interaction plays an important role in shaping the geographic and temporal variation in plant defense in D. stramonium.     

Costs of glandular trichomes in Datura wrightii: a three-year study

Models accounting for genetic variation for resistance to herbivores within plant populations often postulate a balance between the costs of that resistance and its benefits. The production of glandular trichomes by Datura wrightii was shown to be costly in a previous one-year study because plants producing glandular trichomes (sticky plants), a factor conferring resistance to some insect herbivores, also produced 45% fewer seeds than plants producing nonglandular trichomes (velvety plants) when grown in a common garden. Because sticky plants tended to be larger than velvety plants but produced fewer seed capsules, we postulated an allocation trade-off in which velvety plants are more reproduction-dominated whereas sticky plants are more growth-dominated. If a greater commitment to vegetative growth eventually allows sticky plants to compensate for reduced seed production, we would expect a reduction or elimination of the cost of resistance over time in this perennial plant. We monitored growth, survival, and seed production of plants from defined crosses of local populations for three years in a common garden when exposed to and protected from herbivores, and with and without supplemental water. The majority of plants exposed to herbivores had died by the end of the study. We used standard life-table methods to determine the net reproductive rate (R0) and the finite rate of increase (lambda) of plants of each trichome type. After three years, when plants were protected from herbivores, sticky plants were 187-245% larger than velvety plants, depending upon irrigation treatment, but sticky plants continued to be less efficient in producing seeds per unit of canopy volume. Even though the total seed production of sticky plants eventually equaled that of velvety plants, the advantage of earlier reproduction by velvety plants increased lambda by 55-230% over that of sticky plants, depending upon herbivore and irrigation treatment. Exposure to herbivores reduced lambda by 69-83%, depending upon plant type and irrigation treatment, whereas supplemental irrigation increased lambda by 29-175%, depending upon plant type and exposure to herbivores. Although there was a large allocation trade-off between growth and reproduction, the benefits of such a trade-off did not emerge before most plants were killed by herbivores. The cost of producing glandular trichomes strictly for herbivore resistance continued to exceed its benefits, and in the absence of other, unmeasured benefits from the suite of life-history characters associated with glandular trichome production, natural selection is expected to eliminate this costly resistance trait from D. wrightii populations.     

Explaining differential herbivory in sun and shade: the case of Aristotelia chilensis saplings

Differential herbivory in contrasting environments is commonly explained by differences in plant traits. When several plant traits are considered, separate correlation analyses between herbivory and candidate traits are typically conducted. This makes it difficult to discern which trait best explain the herbivory patterns, or to avoid spurious inferences due to correlated characters. Aristotelia chilensis saplings sustain greater herbivory in shaded environments than in open habitats. We measured alkaloids, phenolics, trichomes, leaf thickness and water content in the same plants sampled for herbivory. We conducted a multiple regression analysis to estimate the relationship between herbivory and each plant trait accounting for the effect of correlated traits, thus identifying which trait(s) better explain(s) the differential herbivory on A. chilensis. We also estimated insect abundance in both light environments. Palatability bioassays tested whether leaf consumption by the main herbivore on A. chilensis was consistent with field herbivory patterns. Overall insect abundance was similar in open and shaded environments. While saplings in open environments had thicker leaves, lower leaf water content, and higher concentration of alkaloids and phenolics, no difference in trichome density was detected. The multiple regression analysis showed that leaf thickness was the only trait significantly associated with herbivory. Thicker leaves received less damage by herbivores. Sawfly larvae consumed more leaf tissue when fed on shade leaves. This result is consistent with field herbivory and, together with results of insect abundance, renders unlikely that the differential herbivory in A. chilensis was due to greater herbivory pressure in open habitats.     

Plant latex and other exudates as plant defense systems: roles of various defense chemicals and proteins contained therein

Plant latex and other exudates are saps that are exuded from the points of plant damage caused either mechanically or by insect herbivory. Although many (ca. 10%) of plant species exude latex or exudates, and although the defensive roles of plant latex against herbivorous insects have long been suggested by several studies, the detailed roles and functions of various latex ingredients, proteins and chemicals, in anti-herbivore plant defenses have not been well documented despite the wide occurrence of latex in the plant kingdom. Recently, however, substantial progress has been made. Several latex proteins, including cysteine proteases and chitin-related proteins, have been shown to play important defensive roles against insect herbivory. In the mulberry (Morus spp.)-silkworm (Bombyx mori) interaction, an old and well-known model system of plant-insect interaction, plant latex and its ingredients - sugar-mimic alkaloids and defense protein MLX56 - are found to play key roles. Complicated molecular interactions between Apocynaceae species and its specialist herbivores, in which cardenolides and defense proteins in latex play key roles, are becoming more and more evident. Emerging observations suggested that plant latex, analogous to animal venom, is a treasury of useful defense proteins and chemicals that has evolved through interspecific interactions. On the other hand, specialist herbivores developed sophisticated adaptations, either molecular, physiological, or behavioral, against latex-borne defenses. The existence of various adaptations in specialist herbivores itself is evidence that latex and its ingredients function as defenses at least against generalists. Here, we review molecular and structural mechanisms, ecological roles, and evolutionary aspects of plant latex as a general defense against insect herbivory and we discuss, from recent studies, the unique characteristics of latex-borne defense systems as transport systems of defense substances are discussed based on recent studies.     

Above‐ and belowground herbivory jointly impact defense and seed dispersal traits in Taraxacum officinale

Plants are able to cope with herbivores by inducing defensive traits or growth responses that allow them to reduce or avoid the impact of herbivores. Since above‐ and belowground herbivores differ substantially in life‐history traits, for example feeding types, and their spatial distribution, it is likely that they induce different responses in plants. Moreover, strong interactive effects on defense and plant growth are expected when above‐ and belowground herbivores are jointly present. The strengths and directions of these responses have been scarcely addressed in the literature. Using Taraxacum officinale, the root‐feeding nematode Meloidogyne hapla and the locust Schistocerca gregaria as a model species, we examined to what degree above‐ and belowground herbivory affect (1) plant growth responses, (2) the induction of plant defensive traits, that is, leaf trichomes, and (3) changes in dispersal‐related seed traits and seed germination. We compared the performance of plants originating from different populations to address whether plant responses are conserved across putative different genotypes. Overall, aboveground herbivory resulted in increased plant biomass. Root herbivory had no effect on plant growth. Plants exposed to the two herbivores showed fewer leaf trichomes than plants challenged only by one herbivore and consequently experienced greater aboveground herbivory. In addition, herbivory had effects that reached beyond the individual plant by modifying seed morphology, producing seeds with longer pappus, and germination success.     

Non-protein amino acids in plant defense against insect herbivores: representative cases and opportunities for further functional analysis

Chemical defense against herbivores is of utmost importance for plants. Primary and secondary metabolites, including non-protein amino acids, have been implicated in plant defense against insect pests. High levels of non-protein amino acids have been identified in certain plant families, including legumes and grasses, where they have been associated with resistance to insect herbivory. Non-protein amino acids can have direct toxic effects via several mechanisms, including misincorporation into proteins, obstruction of primary metabolism, and mimicking and interfering with insect neurological processes. Additionally, certain non-protein amino acids allow nitrogen to be stored in a form that is metabolically inaccessible to herbivores and, in some cases, may act as signals for further plant defense responses. Specialized insect herbivores often possess specific mechanisms to avoid or detoxify non-protein amino acids from their host plants. Although hundreds of non-protein amino acids have been found in nature, biosynthetic pathways and defensive functions have been elucidated in only a few cases. Next-generation sequencing technologies and the development of additional plant and insect model species will facilitate further research on the production of non-protein amino acids, a widespread but relatively uninvestigated plant defense mechanism.     

The Arabidopsis trichome is an active mechanosensory switch

Trichomes (‘hair cells’) on Arabidopsis thaliana stem and leaf surfaces provide a range of benefits arising from their shape and disposition. These include tempting herbivores to sample constitutive toxins before they reach the bulk of the tissue. We asked whether, in addition, small mechanical disturbances such as an insect can make elicit signals that might help the plant respond to herbivory. We imaged, pressed and brushed trichomes in several ways, most notably with confocal microscopy of trichomes transgenically provided with apoplastic pH reporter apo‐pHusion and cytosolic Ca²⁺ reporter cameleon. In parallel, we modelled trichome wall mechanics with finite element analysis. The stimulated trichome focuses force on a pliant zone and the adjoining podium of the stalk. A buckling instability can further focus force on a skirt of cells surrounding the podium, eliciting oscillations of cytosolic Ca²⁺ and shifts in apoplastic pH. These observations represent active physiological response. Modelling establishes that the effectiveness of force focusing and buckling is due to the peculiar tapering wall structure of the trichome. Hypothetically, these active mechanosensing functions enhance toxin synthesis above constitutive levels, probably via a priming process, thus minimizing the costly accumulation of toxins in the absence of herbivore attack but assuring rapid build‐up when needed.     

Incidence of the introduced parasitoids Cotesia kazak and Microplitis croceipes (Hymenoptera: Braconidae) from Helicoverpa armigera (Lepidoptera: Noctuidae) in tomatoes, sweet corn, and lucerne in New Zealand

Morphological defense traits of plants such as trichomes potentially compromise biological control in agroecosystems because they may hinder predation by natural enemies. To investigate whether plant trichomes hinder red imported fire ants, Solenopsis invicta Buren (Hymenoptera: Formicidae), as biological control agents in soybean, field and greenhouse experiments were conducted in which we manipulated fire ant density in plots of three soybean isolines varying in trichome density. Resulting treatment effects on the abundance of herbivores, other natural enemies, plant herbivory, and yield were assessed. Trichomes did not inhibit fire ants from foraging on plants in the field or in the greenhouse, and fire ant predation of herbivores in the field was actually greater on pubescent plants relative to glabrous plants. Consequently, fire ants more strongly reduced plant damage by herbivores on pubescent plants. This effect, however, did not translate into greater yield from pubescent plants at high fire ant densities. Intraguild predation by fire ants, in contrast, was weak, inconsistent, and did not vary with trichome density. Rather than hindering fire ant predation, therefore, soybean trichomes instead increased fire ant predation of herbivores resulting in enhanced tritrophic effects of fire ants on pubescent plants. This effect was likely the result of a functional response by fire ants to the greater abundance of caterpillar prey on pubescent plants. Given the ubiquity of lepidopteran herbivores and the functional response to prey shown by many generalist arthropod predators, a positive indirect effect of trichomes on predation by natural enemies might be more far more common than is currently appreciated.     

Focus on Metabolism: Alteration of Plant Primary Metabolism in Response to Insect Herbivory

Plants in nature, which are continuously challenged by diverse insect herbivores, produce constitutive and inducible defenses to reduce insect damage and preserve their own fitness. In addition to inducing pathways that are directly responsible for the production of toxic and deterrent compounds, insect herbivory causes numerous changes in plant primary metabolism. Whereas the functions of defensive metabolites such as alkaloids, terpenes, and glucosinolates have been studied extensively, the fitness benefits of changes in photosynthesis, carbon transport, and nitrogen allocation remain less well understood. Adding to the complexity of the observed responses, the feeding habits of different insect herbivores can significantly influence the induced changes in plant primary metabolism. In this review, we summarize experimental data addressing the significance of insect feeding habits, as related to herbivore-induced changes in plant primary metabolism. Where possible, we link these physiological changes with current understanding of their underlying molecular mechanisms. Finally, we discuss the potential fitness benefits that host plants receive from altering their primary metabolism in response to insect herbivory.Plants in nature are subject to attack by a wide variety of phytophagous insects. Nevertheless, the world is green, and most plants are resistant to most individual species of insect herbivores. To a large extent, this resistance is due to an array of toxic and deterrent small molecules and proteins that can prevent nonadapted insects from feeding. Although many plant defenses are produced constitutively, others are inducible (i.e. defense-related metabolites and proteins that are normally present at low levels become more abundant in response to insect feeding). Inducible defense systems, which allow more energy to be directed toward growth and reproduction in the absence of insect herbivory, represent a form of resource conservation. Well-studied examples of inducible plant defenses include the production of nicotine in tobacco (Nicotiana tabacum; Baldwin et al., 1998), protease inhibitors in tomato (Solanum lycopersicum; Ryan, 2000), benzoxazinoids in maize (Zea mays; Oikawa et al., 2004), and glucosinolates in Arabidopsis (Arabidopsis thaliana; Mewis et al., 2005). Additionally, herbivore-induced plant responses can include the production of physical defenses such as trichomes or thickened cell walls that can make insect feeding more difficult. Some plant defensive metabolites are highly abundant, suggesting that their biosynthesis can have a significant effect on overall plant metabolism. For instance, benzoxazinoids can constitute 1% to 2% of the total dry matter of some Poaceae (Zúñiga et al., 1983), and up to 6% of the nitrogen in herbivore-induced Nicotiana attenuata can be devoted to nicotine production (Baldwin et al., 1998).In addition to the herbivore-induced production of physical and chemical defenses, numerous changes in plant primary metabolism occur in response to insect herbivory. Among other observed effects, these can include either elevated or suppressed photosynthetic efficiency, remobilization of carbon and nitrogen resources, and altered plant growth rate. However, although the defensive value of induced toxins such as nicotine, terpenes, benzoxazinoids, and glucosinolates is clear, it is sometimes more difficult to elucidate the function of herbivore-induced changes in plant primary metabolism. Insects may also manipulate plant primary metabolism for their own benefit, making it challenging to determine whether the observed changes are actually a plant defensive response.Here, we describe commonly observed changes in plant primary metabolism, focusing on carbohydrates and nitrogen, and discuss their possible functions in plant defense against insect herbivory. There are large differences among published studies involving different plant-herbivore combinations, and no universal patterns in the herbivory-induced changes in plant primary metabolism. Therefore, we also discuss how the potential benefits can depend on the tissue that is being attacked, the extent of the tissue damage, and the type of insect herbivore that is involved in the interaction.     

Host plant trichomes and the advantage of being big: progeny size variation of the pipevine swallowtail

1. Plants possess numerous traits that confer resistance against insect herbivores, and herbivores, in turn, can evolve traits to ameliorate the effectiveness of these traits. The pipevine swallowtail, Battus philenor, is an extreme specialist on plants in the genus Aristolochia. The only host plant available to the California population of B. philenor is A. californica. Aristolochia californica is distinct from most other B. philenor host plants in that it is pubescent. 2. The progeny of B. philenor are larger in California compared with populations examined in Texas. Size differences persist throughout larval development. 3. Regardless of maternal host plant, population differences in progeny size persist, and crosses between California (large progeny) and Texas (small progeny) B. philenor populations resulted in offspring producing intermediate sized progeny, indicating a heritable component to progeny size variation. 4. California neonate caterpillars more easily overcame the trichomes of A. californica compared with Texas neonates. When trichomes were removed from A. californica, time to feeding establishment was reduced for caterpillars from both populations. Texas caterpillars established feeding sites on A. californica with trichomes removed, in the same time required to establish feeding on their non‐pubescent host plant, A. erecta. 5. This study shows that plant trichomes might impose selection pressure on progeny size.     

Effects of decomposers and herbivores on plant performance and aboveground plant-insect interactions

Most ecologists acknowledge that plants are subject to complex interactions between both below- and aboveground dwelling animals. However, these complex interactions are seldomly investigated simultaneously. In a factorial common garden experiment we tested single and combined effects of decomposers, root herbivores and leaf herbivores on the growth, flower visitation, and abundance of naturally colonizing aphids and parasitoids on wild mustard ( Sinapis arvensis ). We found that the individual presence of either root herbivores or decomposers resulted in increased aphid abundance, demonstrating that the same aboveground plant–insect interaction can be released by different belowground processes. Enhanced aphid densities caused higher numbers of parasitoids. Furthermore, decomposers increased plant growth and plant fitness (measured as the number of seeds produced), indicating that mustard may benefit from nutrients provided by decomposers, regardless whether plants are attacked by root herbivores or leaf herbivores, or both simultaneously. More flower visits were observed in plants attacked by root herbivores but without leaf herbivores than in plants with both herbivores, suggesting that root herbivory can modify flower attractivity to pollinators. Our results suggest that patterns in plant–insect interactions above the ground are not only affected by aboveground factors but also by a wealth of different belowground processes mediated by the plant.     

Chemical and Mechanical Defenses Vary among Maternal Lines and Leaf Ages in Verbascum thapsus L. (Scrophulariaceae) and Reduce Palatability to a Generalist Insect

Intra-specific variation in host-plant quality affects herbivore foraging decisions and, in turn, herbivore foraging decisions mediate plant fitness. In particular, variation in defenses against herbivores, both among and within plants, shapes herbivore behavior. If variation in defenses is genetically based, it can respond to natural selection by herbivores. We quantified intra-specific variation in iridoid glycosides, trichome length, and leaf strength in common mullein (Verbascum thapsus L, Scrophulariaceae) among maternal lines within a population and among leaves within plants, and related this variation to feeding preferences of a generalist herbivore, Trichopulsia ni Hübner. We found significant variation in all three defenses among maternal lines, with T. ni preferring plants with lower investment in chemical, but not mechanical, defense. Within plants, old leaves had lower levels of all defenses than young leaves, and were strongly preferred by T. ni. Caterpillars also preferred leaves with trichomes removed to leaves with trichomes intact. Differences among maternal lines indicate that phenotypic variation in defenses likely has a genetic basis. Furthermore, these results reveal that the feeding behaviors of T. ni map onto variation in plant defense in a predictable way. This work highlights the importance of variation in host-plant quality in driving interactions between plants and their herbivores.     

Peltate glandular trichomes of Colquhounia seguinii harbor new defensive clerodane diterpenoids

Glandular trichomes produce a wide variety of secondary metabolites that are considered as major defen-sive chemicals against herbivore attack. The morphology and secondary metabolites of the peltate g...