Roles of the volatile terpene, 1,8-cineole,in plant–herbivore interactions: a foraging odor cue as well as a toxin?

Olfaction is an important sense for many animals, yet its role in foraging by herbivores is poorly known. Many plants contain volatile compounds, such as terpenes, that are not only volatile but can be toxic if ingested. Volatile terpenes can be used by herbivores to assess leaf quality, but there is little evidence for whether they are also used as a searching cue. We applied the giving-up density (GUD) framework to examine fine-scale foraging by two free-ranging mammalian herbivores, the brush-tail possum (Trichosurus vulpecula) and the swamp wallaby (Wallabia bicolor), using patches with food and an inedible matrix that varied in content of a volatile terpene, 1,8-cineole. We tested the effect of (1) increasing dietary cineole concentration, and (2) masking the food odor by adding cineole to the inedible matrix, thus overriding the smell released by the diet. In both species GUD was affected by dietary cineole; a high cineole concentration raised GUD, consistent with its role as a toxin. There was a significant effect of masking on GUD for wallabies but not for possums, suggesting that odor was an important foraging cue at the feeding patch only for the former. Differences in ecological niche and diet may explain this pattern. We suggest that herbivores, such as the swamp wallaby, opportunistically eavesdrop on plant volatiles, i.e., take advantage of the signal proffered for a different function. The cost of this eavesdropping for plants, however, is presumably counteracted by other ecological benefits of these volatiles, including a reduction in leaf consumption as a function of toxicity.     

Integrating the costs of plant toxins and predation risk in foraging decisions of a mammalian herbivore

Foraging herbivores must satisfy their nutrient requirements in a world of toxic plants while also avoiding predators. Plant toxins and perceived predation risk at food patches should both reduce patch residency time, but the relative strengths of these factors on feeding decisions has rarely been quantified. Using an arboreal generalist herbivore, the common brushtail possum Trichosurus vulpecula, we tested the effects on food intake of the plant toxin, cineole, and regurgitated pellets from one of its predators, the powerful owl Ninox strenua at the small spatial scale of the food patch. We used the giving-up density (GUD) framework, with animals harvesting food items (sultanas) in an inedible matrix (small pebbles). We ran two consecutive field experiments in a eucalypt woodland in eastern Australia, 1 month apart in the same location. In experiment 1, there was a significant interaction between cineole [at 17% of dry matter (DM)] and owl pellets. The GUD was lowest in the absence of both cineole and owl pellet, intermediate in the presence of owl pellet; and highest with cineole ± owl pellet. The effect of owl pellet diminished over time. In experiment 2, only cineole (at 10% DM) increased the GUD significantly. The difference in effect of owl pellet was probably due to both habituation and freshness of the cue. Our study demonstrates the importance of synthesising predator–prey and plant–herbivore ecology to better understand the complex set of constraints influencing foraging herbivores. The greater effect of toxin than fear on possums is likely to be due to its high, but ecologically relevant concentration. This highlights the need to explore the relative and net impacts of a range of concentrations of plant toxins and predation risks.     

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.     

Plant silicon effects on insect feeding dynamics are influenced by plant nitrogen availability

Although there is growing evidence that silicon (Si)‐based plant defenses effectively reduce both the palatability and digestibility of leaves, and thus impact nutrient assimilation by insect herbivores, much less is known about how this is affected by extrinsic and intrinsic factors. For example, do herbivores exhibit compensatory feeding on poor‐quality diets with Si or are Si defenses less effective in agroecosystems where high N availability increases plant quality? To investigate the interactive effects of N and Si on insect feeding, we conducted insect performance and compensatory feeding bioassays using maize, Zea mays L. (Poaceae), and the true armyworm, Pseudeletia unipuncta Haworth (Lepidoptera: Noctuidae). In the performance assay, the addition of Si alone resulted in increased larval mortality compared with the controls, likely because early instars with poorly developed mandibles could not feed effectively. However, larvae fed on plants treated with both Si and N survived better than on plants treated with Si only, although pupal mass did not differ between treatments. In our compensatory assay, Si addition reduced maize consumption, but increased both armyworm approximate digestibility and N assimilation efficiency, suggesting that enhanced post‐ingestion feeding physiology, rather than compensatory food intake, could have accounted for the lack of Si effects on pupal weight. Overall, our results demonstrate that, similar to other chemical and mechanical defenses, the effectiveness of plant Si defense is influenced by plant nutrient status and consumer compensatory ability.     

Early ontogenetic trajectories vary among defence chemicals in seedlings of a fast‐growing eucalypt

Ontogenetic changes in leaf chemistry can affect plant–herbivore interactions profoundly. Various theoretical models predict different ontogenetic trajectories of defence chemicals. Empirical tests do not consistently support one model. In Eucalyptus nitens, a fast‐growing tree, we assessed early developmental changes to seedlings, in foliage concentrations of nitrogen and the full suite of known secondary (defence) chemicals. This included the terpene, α‐pinene, whose impact on marsupial herbivory is unknown. To test for the influence of abiotic conditions on the ontogenetic trajectories we overlaid a nutrient treatment. Ontogenetic trajectories varied among compounds. Sideroxylonals and cineole were barely detected in very young seedlings, but increased substantially over the first 200 days. Total phenolic concentration increased fourfold over this time. In contrast, α‐pinene concentration peaked within the first 60 days and again between 150 and 200 days. Nutrients altered the degree but not the direction of change of most chemicals. A shorter trial run at a different season showed qualitatively similar patterns, although α‐pinene concentration started very high. We investigated the effect of detected levels of α‐pinene and cineole on food intake by two mammalian herbivores, common brushtail possums (Trichosurus vulpecula) and red‐bellied pademelons (Thylogale billardierii). Under no‐choice conditions neither terpene reduced intake; but with a choice, possums preferred α‐pinene to cineole. The ontogenetic trajectories of most compounds were therefore consistent with models that predict an increase as plants develop. Published data from later developmental stages in E. nitens also confirm this pattern. α‐Pinene, however, was the only secondary compound found at significant levels in very young seedlings; but it did not constrain feeding by marsupial herbivores. Models must allow for different roles of defensive secondary chemicals, presumably associated with different selective pressures as plants age, which result in different ontogenetic trajectories.     

Olfactory and visual plant cues as drivers of selective herbivory

Food quality is an important consideration in the foraging strategy of all animals, including herbivores. Those that can detect and assess the nutritional value of plants from afar, using senses such as smell and sight, can forage more efficiently than those that must assess food quality by taste alone. Selective foraging not only affects herbivore fitness but can influence the structure and composition of plant communities, yet little is known about how olfactory and visual cues help herbivores to find preferred plants. We tested the ability of a free‐ranging, generalist mammalian browser, the swamp wallaby Wallabia bicolor, to use olfactory and visual plant cues to find and/or browse differentially on Eucalyptus pilularis seedlings grown under different nutrient conditions. Low‐nutrient seedlings differed from high‐nutrient seedlings, having lighter coloured leaves, red stems and lower biomass and nitrogen content. In the absence of visual cues, wallabies used odour to differentiate vials containing cut seedlings. They visited and investigated patches with high‐nutrient seedling odour most, followed by patches with low‐nutrient seedling odour, and patches with no added odour least. However, when visual and olfactory cues of seedlings were present, wallabies reversed their foraging response and were more likely to browse low‐ than high‐nutrient seedlings. This browsing difference, in turn, disappeared when long‐range visual cues were reduced by pinning seedlings horizontal to the ground. We suggest that visual cues overrode the effects of olfactory cues on browsing patterns of intact seedlings. Our study shows that herbivores can respond to odours of higher nutrient plants but in ecologically realistic scenarios they use a variety of visual and olfactory cues, with a context‐dependent outcome that is not always selection of high nutrient food. Our results demonstrate the importance of testing the sensory abilities of herbivores in realistic multi‐sensory settings to understand their function in selective foraging.     

植物诱导性直接防御

众所周知,植物对植食性昆虫危害的反应表现在3个方面：直接防御,间接防御,和耐害性。直接防御是指植物自身所具有的能影响寄主植物感虫性的所有特性。植物对昆虫危害的直接防御包括：限制食物供给,降低营养价值,减少偏嗜程度,破坏组织结构和抑制害虫代谢途径。目前已知的防御化合物主要包括植物次生代谢物质、昆虫消化酶（蛋白）抑制剂、蛋白酶、凝集素、氨基酸脱氨酶和氧化酶。植物在防御某种昆虫为害时多个因素往往具有累加效应或协同作用,并且对一种昆虫起主导作用的因素在防御另一种昆虫时可能仅仅起次要作用甚至根本不起作用。因此,对寄主植物基因表达、蛋白水平和活性以及代谢物含量在不同时空条件下进行广泛的定量和定性的高通量分析,不仅可以促进对植物直接防御机制的全面理解,而且有助于在农业生产中加快对作物抗性的特定靶标的鉴定。     

Inducible direct plant defense against insect herbivores: A review

Plants respond to insect herbivory with responses broadly known as direct defenses, indirect defenses, and tolerance. Direct defenses include all plant traits that affect susceptibility of host plants by themselves. Overall categories of direct plant defenses against insect herbivores include limiting food supply, reducing nutrient value, reducing preference, disrupting physical structures, and inhibiting chemical pathways of the attacking insect. Major known defense chemicals include plant secondary metabolites, protein inhibitors of insect digestive enzymes, proteases, lectins, amino acid deaminases and oxidases. Multiple factors with additive or even synergistic impact are usually involved in defense against a specific insect species, and factors of major importance to one insect species may only be of secondary importance or not effective at all against another insect species. Extensive qualitative and quantitative high throughput analyses of temporal and spatial variations in gene expression, protein level and activity, and metabolite concentration will accelerate not only the understanding of the overall mechanisms of direct defense, but also accelerate the identification of specific targets for enhancement of plant resistance for agriculture.     

A quantitative evaluation of major plant defense hypotheses, nature versus nurture, and chemistry versus ants

Variation in plant secondary metabolite content can arise due to environmental and genetic variables. Because these metabolites are important in modifying a plant’s interaction with the environment, many studies have examined patterns of variation in plant secondary metabolites. Investigations of chemical defenses are often linked to questions about the efficacies of plant defenses and hypotheses on their evolution in different plant guilds. We performed a series of meta-analyses to examine the importance of environmental and genetic sources of variation in secondary metabolites as well as the antiherbivore properties of different classes of defense. We found both environmental and genetic variation affect secondary metabolite production, supporting continued study of the carbon-nutrient balance and growth-differentiation balance hypotheses. Defenses in woody plants are more affected by genetic variation, and herbaceous plant defenses are more influenced by environmental variation. Plant defenses in agricultural and natural systems show similar responses to manipulations, as do plants in laboratory, greenhouse, or field studies. What does such variation mean to herbivores? A comparison of biotic, physical, and chemical defenses revealed the most effective defensive strategy for a plant is biotic mutualisms with ants. Fast-growing plants are most often defended with qualitative defenses and slow-growing plants with quantitative defenses, as the plant apparency and resource availability hypotheses predict. However, we found the resource availability hypothesis provides the best explanation for the evolution of plant defenses, but the fact that there is considerable genetic and environmental variation in defenses indicates herbivores can affect plant chemistry in ecological and evolutionary time.     

Interplant variation in creosotebush foliage characteristics and canopy arthropods

Summary We conducted a field study to test the hypothesis that creosotebush (Larrea tridentata) shrubs growing in naturally nutrient-rich sites had better quality foliage and supported greater populations of foliage arthropods than shrubs growing in nutrient-poor sites. Nutrient-rich sites had significantly higher concentrations of soil nitrogen than nutrient-poor sites. Multivariate analysis of variance revealed significant differences between high nutrient and low nutrient shrubs based on a number of structural and chemical characteristics measured. High nutrient shrubs were larger, had denser foliage, greater foliage production, higher concentrations of foliar nitrogen and water, and lower concentrations of foliar resin than low nutrient shurbs. Numbers of foliage arthropods, particularly herbivores and predators, were significantly higher on high nutrient shrubs. Shrub characteristics and foliage arthropod abundances varied considerably from shrub to shrub. Shrub characteristics representing shrub size, foliage density, foliage growth, and foliar nitrogen and water concentrations were positively correlated with arthropod abundances. Foliar resin concentrations were negatively correlated with foliage arthropod abundances. The positive relationship between creosotebush productivity and foliage arthropods is contradictory to the tenet that physiologically stressed plants provide better quality foliage to insect herbivores.     

The influence of bite size on foraging at larger spatial and temporal scales by mammalian herbivores

Organisms respond to their heterogeneous environment in complex ways at many temporal and spatial scales. Here, I examine how the smallest scale process in foraging by mammalian herbivores, taking a bite, influences plants and herbivores over larger scales. First, because cropping bites competes with chewing them, bite size influences short-term intake rate of herbivores within plant patches. On the other hand, herbivores can chew bites while searching for new ones, thus influencing the time spent vigilant and intake rate as animals move among food patches. Therefore, bite size affects how much time herbivores must spend foraging each day. Because acquiring energy is necessary for fitness, herbivores recognize the importance of bite size and select bites, patches and diets based on tradeoffs between harvesting rates, digestion, and sheering forces. In turn, induced structural defenses of plants, such as thorns, allow plants to respond immediately to herbivory by reducing bite size and thus tissue loss. Over evolutionary time, herbivores have adapted mouth morphology that allows them to maximize bite size on their primary forage plant, whereas plants faced with large mammalian herbivores have adapted structures such as divarication that minimize bite size and protect themselves from herbivory. Finally, bite size available among plant communities can drive habitat segregation and migration of larger herbivores across landscapes.     

Elevated volatile concentrations in high‐nutrient plants: do insect herbivores pay a high price for good food?

1. A tritrophic perspective is fundamental for understanding the drivers of insect–plant interactions. While host plant traits can directly affect insect herbivore performance by either inhibiting or altering the nutritional benefits of consumption, they can also have an indirect effect on herbivores by influencing rates of predation or parasitism. 2. Enhancing soil nutrients available to trees of the genus Eucalyptus consistently modifies plant traits, typically improving the nutritional quality of the foliage for insect herbivores. We hypothesised that resulting increases in volatile essential oils could have an indirect negative effect on eucalypt‐feeding herbivores by providing their natural enemies with stronger host/prey location cues. 3. Eucalyptus tereticornis Smith seedlings were grown under low‐ and high‐nutrient conditions and the consequences for the release of volatile cues from damaged plants were examined. The influence of 1,8‐cineole (the major volatile terpene in many Eucalyptus species) on rates of predation on model caterpillars in the field was then examined. 4. It was found that the emission of cineole increased significantly after damage (artificial or herbivore), but continued only when damage was sustained by herbivore feeding. Importantly, more cineole was emitted from high‐ than low‐nutrient seedlings given an equivalent amount of damage. In the field, predation was significantly greater on model caterpillars baited with cineole than on unbaited models. 5. These findings are consistent with the hypothesis that any performance benefits insect herbivores derive from feeding on high‐nutrient eucalypt foliage could be at least partially offset by an increased risk of predation or parasitism via increased emission of attractive volatiles.     

Herbivory-induced overcompensation and resource-dependent production of extrafloral nectaries in Luffa cylindrica (Cucurbitaceae)

Extrafloral nectaries (EFNs) are nectar secretory structures involved in the indirect defense of plants. In the sponge gourd (Luffa cylindrica), EFNs commonly occur on the lower surface of leaf blades and stipules and remain functional until leaf senescence. To test the hypothesis that the development of EFNs is influenced by herbivore damage and resource availability, we grew Luffa cylindrica under different concentrations of Hoagland's nutrient solution (nutrient-poor conditions: 10%, 50%; and control condition: 100%) and two herbivory treatments (damaged and undamaged leaves). We collected ten leaves from treated plants to quantify leaf area and EFN density. Overall, leaf area increased and EFN decreased in damaged plants, but this significantly depended on nutritional status. In undamaged plants, EFN density tended to remain constant, whereas foliar area increased with nutrient input. Under herbivory, foliar area increased at 10% but decreased at 50 and 100% of nutrients in relation to undamaged plants, whereas EFN density tended to increase with nutrient availability to exceed undamaged plants under control concentrations. Plants under nutrient-poor conditions subjected to herbivory exhibited an increased foliar area, characterizing a compensatory mechanism. Our results suggest that herbivore-induced indirect defense is a damage- and resource-dependent response in Luffa cylindrica. These findings contribute to understanding the factors that modulate indirect defenses and plant-herbivore-environment interactions.     

Territorial defense against various food competitors in the Tanganyikan benthophagous cichlid <Emphasis Type="Italic">Neolamprologus tetracanthus</Emphasis>

Territorial defense of nonbreeding female Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid, was investigated. Females defended territories (=home ranges, ca. 1m across) against a variety of intruding fishes. Conspecific females were usually attacked outside the territories, heterospecific benthivores (shrimp eaters) and omnivores near the border of the territories, and piscivores, algae and detritus feeders, and herbivores inside the territories. Females used some parts of the sandy substrate in the territories for foraging (foraging areas). Territorial defense prevented most of the conspecific females and benthivores from intruding into the foraging areas. In omnivores, piscivores, and algae and detritus feeders, about half the intruders were repelled from the foraging areas, although herbivores were infrequently repelled in the areas. Soon after removal of the resident females, many food competitors invaded the foraging areas and eagerly devoured prey, suggesting that the territories are maintained for food resource protection from these competitors. Females are likely to discriminate intruding fishes and change their territorial defense primarily on the basis of the degree of dietary overlap, resulting in monofunctional serial territories.     

A novel trade-off of insect diapause affecting a sequestered chemical defense

Diapause allows insects to temporally avoid conditions that are unfavorable for development and reproduction. However, diapause may incur a cost in the form of reduced metabolic energy reserves, reduced potential fecundity, and missed reproductive opportunities. This study investigated a hitherto ignored consequence of diapause: trade-offs involving sequestered chemical defense. We examined the aristolochic acid defenses of diapausing and non-diapausing pipevine swallowtail butterflies, Battus philenor. Pipevine swallowtail larvae acquire these chemical defenses from their host plants. Butterflies that emerge following pupal diapause have significantly less fat, a female fitness correlate, compared to those that do not diapause. However, butterflies emerging from diapaused pupae are more chemically defended compared to those that have not undergone diapause. Furthermore, non-diapausing butterflies are confronted with older, lower quality host plants on which to oviposit. Thus, a trade-off exists where butterflies may have greater energy reserves at the cost of less chemical defense and sub-optimal food resources for their larvae, or have substantially less energetic reserves with the benefit of greater chemical defense and plentiful larval food resources.     

A Specialist Herbivore Uses Chemical Camouflage to Overcome the Defenses of an Ant-Plant Mutualism

Many plants and ants engage in mutualisms where plants provide food and shelter to the ants in exchange for protection against herbivores and competitors. Although several species of herbivores thwart ant defenses and extract resources from the plants, the mechanisms that allow these herbivores to avoid attack are poorly understood. The specialist insect herbivore, Piezogaster reclusus (Hemiptera: Coreidae), feeds on Neotropical bull-horn acacias (Vachellia collinsii) despite the presence of Pseudomyrmex spinicola ants that nest in and aggressively defend the trees. We tested three hypotheses for how P. reclusus feeds on V. collinsii while avoiding ant attack: (1) chemical camouflage via cuticular surface compounds, (2) chemical deterrence via metathoracic defense glands, and (3) behavioral traits that reduce ant detection or attack. Our results showed that compounds from both P. reclusus cuticles and metathoracic glands reduce the number of ant attacks, but only cuticular compounds appear to be essential in allowing P. reclusus to feed on bull-horn acacia trees undisturbed. In addition, we found that ant attack rates to P. reclusus increased significantly when individuals were transferred between P. spinicola ant colonies. These results are consistent with the hypothesis that chemical mimicry of colony-specific ant or host plant odors plays a key role in allowing P. reclusus to circumvent ant defenses and gain access to important resources, including food and possibly enemy-free space. This interaction between ants, acacias, and their herbivores provides an excellent example of the ability of herbivores to adapt to ant defenses of plants and suggests that herbivores may play an important role in the evolution and maintenance of mutualisms.     

ECOLOGICAL COSTS OF PLANT RESISTANCE TO HERBIVORES IN THE CURRENCY OF POLLINATION

In this paper, we examine how ecological costs of resistance might be manifested through plant relationships with pollinators. If defensive compounds are incorporated into floral structures or if they are sufficiently costly that fewer rewards are offered to pollinators, pollinators may discriminate against more defended plants. Here we consider whether directional selection for increased resistance to herbivores could be constrained by opposing selection through pollinator discrimination against more defended plants. We used artificial selection to create two populations of Brassica rapa plants that had high and low myrosinase concentrations and, consequently, high and low resistance to flea beetle herbivores. We measured changes in floral characters of plants in both damaged and undamaged states from these populations with different resistances to flea beetle attack. We also measured pollinator visitation to plants, including numbers of pollinators and measures of visit quality (numbers of flowers visited and time spent per flower). Damage from herbivores resulted in reduced petal size, as did selection for high resistance to herbivores later in the plant lifetime. In addition, floral display (number of open flowers) was also altered by an interaction between these two effects. Changes in floral traits translated into overall greater use of low-resistance, undamaged plants based on total amount of time pollinators spent foraging on plants. Total numbers of pollinators attracted to plants did not differ among treatments; however, pollinators spent significantly more time per flower on plants from the low-resistance population and tended to visit more flowers on these plants as well. Previous work by other investigators on the same pollinator taxa has shown that longer visit times are associated with greater male and female plant fitness. Because initial numbers of pollinators did not differ between selection regimes, palatability and/or amount of rewards offered by high- and low-resistance populations are likely to be responsible for these patterns. During periods of pollinator limitation, less defended plants may have a selective advantage and pollinator preferences may mediate directional selection imposed by herbivores. In addition, if pollinator preferences limit seed set in highly defended plants, then lower seed set previously attributed to allocation costs of defense may also reflect greater pollinator limitation in these plants relative to less defended plants.     

Endophage-ectophage ratios and plant defense

Summary Endophagous folivores, which are concealed inside leaf tissue for much of their life cycle, or which live externally but feed internally, should be more successful on heavily defended plants than ectophagous species. This is because endophages are more facile at feeding selectively and can manipulate tissue development to avoid physical and chemical defenses and to enhance nutrition. As a result, endophage-ectophage ratios should increase on more heavily defended hosts. This pattern will likely be strengthened by negative asymmetrical interactions with ectophages and pathogens, which may displace endophages from lightly defended hosts. The hypothesis predicts that endophages should be particularly abundant in resource-poor habitats which seem to support a preponderance of heavily defended hosts. Although data do not yet exist for a rigorous test, several observed cases where endophage distributions seem biased toward heavily defended hosts are at least consistent with the hypothesis. Plant defense levels may have little influence on the total number of herbivores associated with a host, but I suggest that guild structure can be profoundly altered.     

Family matters: effect of host plant variation in chemical and mechanical defenses on a sequestering specialist herbivore

Insect herbivores contend with various plant traits that are presumed to function as feeding deterrents. Paradoxically, some specialist insect herbivores might benefit from some of these plant traits, for example by sequestering plant chemical defenses that herbivores then use as their own defense against natural enemies. Larvae of the butterfly species Battus philenor (L.) (Papilionidae) sequester toxic alkaloids (aristolochic acids) from their Aristolochia host plants, rendering larvae and adults unpalatable to a broad range of predators. We studied the importance of two putative defensive traits in Aristolochia erecta: leaf toughness and aristolochic acid content, and we examined the effect of intra- and interplant chemical variation on the chemical phenotype of B. philenor larvae. It has been proposed that genetic variation for sequestration ability is ??invisible to natural selection?? because intra- and interindividual variation in host-plant chemistry will largely eliminate a role for herbivore genetic variation in determining an herbivore??s chemical phenotype. We found substantial intra- and interplant variation in leaf toughness and in the aristolochic acid chemistry in A. erecta. Based on field observations and laboratory experiments, we showed that first-instar larvae preferentially fed on less tough, younger leaves and avoided tougher, older leaves, and we found no evidence that aristolochic acid content influenced first-instar larval foraging. We found that the majority of variation in the amount of aristolochic acid sequestered by larvae was explained by larval family, not by host-plant aristolochic acid content. Heritable variation for sequestration is the predominant determinant of larval, and likely adult, chemical phenotype. This study shows that for these highly specialized herbivores that sequester chemical defenses, traits that offer mechanical resistance, such as leaf toughness, might be more important determinants of early-instar larval foraging behavior and development compared to plant chemical defenses.     

Ecological and Evolutionary Interactions between Wild Crucifers and Their Herbivorous Insects

Abstract Insects feeding on ten species of wild crucifer were investigated. Differences in host plant range and insect community structure were examined with regard to anti-herbivore defense mechanisms. Most of the crucifer species deterred insect herbivory by disappearing in the summer or by lowering their intrinsic quality as food for insects. Species with these defense mechanisms were exploited by only a few specialized herbivorous insects that seemed to have counter defenses. The plants without these defense mechanisms were used by many herbivorous insect species. Rorippa indica lacked direct defenses, but supported a low total density of herbivore individuals. This crucifer has an indirect defense mechanism: ants attracted to floral nectar defended the plant from deleterious herbivores. Crucifers that disappeared seasonally lacked other anti-herbivore defense mechanisms. This suggests that the phonological response is an alternative other responses to herbivore attack.