Searching for synergism: effects of combinations of phenolic compounds and other toxins on oxidative stress in Lymantria dispar caterpillars

Synergistic combinations of plant allelochemicals are commonly believed to increase their effectiveness as defenses against insect herbivores. For example, temperate deciduous trees produce large amounts of phenolic compounds (primarily tannins), but many of these trees also produce smaller amounts of other potentially toxic compounds. This study tested the hypothesis that mixtures of phenolics and other types of toxins produce a greater effect on oxidative stress (a measure of toxicity) than do phenolics alone. Oxidative stress was measured in Lymantria dispar (gypsy moth) larvae as a shift in the redox balance of glutathione (GSH) towards a higher percentage of its oxidized form (glutathione disulfide; GSSG). We began by showing that larvae that ingested ellagitannins and chlorogenic acid (phenolics) contained greatly elevated levels of reactive oxygen species in their midgut contents, but this was not sufficient to shift the redox balance of GSH in their midgut tissues. Therefore, the phenolic compounds were tested in pairwise combinations with juglone (a quinone), rutin, kaempferol, or quercitrin (flavonoids), quinine, berberine, gramine, or glaucine (alkaloids), and soy or Quillaja saponins. When each of these allelochemical combinations was treated on hybrid poplar (Populus tremula × P. alba) leaves and consumed by L. dispar larvae for a 2-day period, none significantly shifted the redox balance of GSH. GSH levels were induced by the combination of phenolics with an alkaloid and a saponin, suggesting that a “cocktail” of allelochemicals might produce a more effective plant defense. GSH levels were also found to increase with larval age, both within and between instars. Increased levels of GSH may help explain the greater resistance of late-instar than early-instar larvae to plant defenses. Overall, the results of this study do not support the hypothesis that the combined effects of phenolic compounds and other toxins in sugar maple (Acer saccharum) leaves produce a synergistic increase in oxidative stress in L. dispar larvae. Further work is needed to test the general hypothesis that the allelochemical diversity in plants functions, in part, to produce synergistic toxicological effects in insect herbivores.     

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.     

Nutritional quality of phloem sap in relation to host plant-alternation in the bird cherry-oat aphid

Summary. Host alternation in aphids has been attributed to complementary growth of host plants, or more specifically to seasonal changes in the nitrogen quality of the phloem sap. In this report, seasonal fluctuation of free amino acids in phloem of the winter and summer host plants (Prunus padus, bird cherry and Hordeum vulgare, barley) of Rhopalosiphum padi (the bird cherry-oat aphid) were investigated in the context of aphid growth and behaviour. Phloem was collected from the cut stylets of aphids taken from plants that were grown outdoors. The total concentration of amino acids in P. padus phloem increased between bud break and late flush (spring), decreased in mature leaves (summer), and increased again in early senescent leaves (autumn). In H. vulgare, however, amino acid concentration fluctuated less from seedlings to flowering. Spring aphids from P. padus grew rapidly on this host from bud break to late flush, but died on mature and early senescent leaves. Summer aphids from H. vulgare grew as fast on this host as spring aphids did on flush leaves of P. padus. Sexual females grew more slowly than other generations and nearly as well on mature as on early senescent P. padus leaves. As judged by aphid growth and phloem nitrogen quality, P. padus during spring equals the summer host H. vulgare. However, the lower growth rates of R. padi on mature and senescent leaves of P. padus appear only loosely correlated with phloem amino acid concentrations. Therefore, factors influencing aphid nutrition, or ecology, other than seasonal changes in phloem sap amino acid concentration may explain the existence of host alternation in R. padi.     

Effects of three years’ increase in density of the geometrid Epirrita autumnata on the change in metabolome of mountain birch trees (Betula pubescens ssp. czerepanovii)

The principal objective of the study was to characterize effects of increasing density of the autumnal moth, Epirrita autumnata (Lepidoptera, Geometridae), for 3 successive years on the change in metabolome of mountain birch trees (Betula pubescens ssp. czerepanovii). During the three study years (2000–2002), the larval density varied from low to very high (outbreak) density. Samples of leaves were collected from the same 12 trees at the same phenological stage of the trees each year. The leaves were collected from undamaged trees in the first year, from trees slightly damaged by larvae in the second year, and from trees heavily damaged by larvae in the third year. Metabolome analysis showed that the increase in density of E. autumnata larvae and degree of damage of birch trees caused multiple biochemical changes in the leaves, including increased concentrations of phenolic compounds (proanthocyanidins and hydrolysable tannins) and reduced concentrations of nutritive metabolites (monosaccharides, amino acids and some organic acids). These changes reduced the quality of leaves as food for larvae and, probably, were associated with induced chemical resistance of the birches to herbivorous insects. Additionally, the concentration of α-tocopherol was significantly higher in both slightly and heavily damaged trees. Mechanisms of changes in the metabolism of phenolic compounds and carbohydrates, and the role of α-tocopherol in their regulation are discussed.     

Population and leaf-level variation of iridoid glycosides in the invasive weed Verbascum thapsus L. (common mullein): implications for herbivory by generalist insects

Plant–insect interactions, which are strongly mediated by chemical defenses, have the potential to shape invasion dynamics. Despite this, few studies have quantified natural variation in key defensive compounds of invasive plant populations, or how those defenses relate to levels of herbivory. Here, we evaluated variation in the iridoid glycosides aucubin and catalpol in rosette plants of naturally occurring, introduced populations of the North American invader, Verbascum thapsus L. (common mullein; Scrophulariaceae). We examined two scales that are likely to structure interactions with insect herbivores—among populations and within plant tissues (i.e., between young and old leaves). We additionally estimated the severity of damage incurred at these scales due to insect chewing herbivores (predominantly grasshoppers and caterpillars), and evaluated the relationship between iridoid glycoside content and leaf damage. We found significant variation in iridoid glycoside concentrations among populations and between young and old leaves, with levels of herbivory strongly tracking leaf-level investment in defense. Specifically, across populations, young leaves were highly defended by iridoids (averaging 6.5× the concentration present in old leaves, and containing higher proportions of the potentially more toxic iridoid, catalpol) and suffered only minimal damage from generalist herbivores. In contrast, old leaves were significantly less defended and accordingly more substantially utilized. These findings reveal that quantitative variation in iridoid glycosides is a key feature explaining patterns of herbivory in an introduced plant. In particular, these data support the hypothesis that defenses limit the ability of generalists to feed on mullein’s well-defended young leaves, resulting in minimal losses of high-quality tissue, and increasing performance of this introduced species.     

Development of the pH in the intestinal tract of larval turbot

The pH in the gut of turbot larvae and juveniles of turbot was studied from day 11 until the completion of metamorphosis. Dietary effects on the gut pH were estimated when larvae were offered live feed, a microdiet, only microalgae or no feed. The pH in the gut was weakly alkaline until day 24 after hatching with no differences between the foregut, midgut and hindgut. The foregut contents started to turn acidic from day 28 after hatching when the larvae were already weaned successfully, which indicates that an acidic pH is not necessary for the digestion and utilisation of formulated feed. During the following 20 days the pH in the foregut/stomach decreased further to a minimum of pH 3.5, while the pH in the midgut and the hindgut increased slightly to a maximum of pH 9.0. Larvae receiving live feed, microdiet or microalgae had a similar pH in the midgut on day 11, while starved larvae exhibited a lower gut pH. This suggests bicarbonate secretion from the larval pancreas stimulated by ingested microalgae or feed particles.     

Life cycle and population dynamics of Pycnogonum litorale (Pycnogonida) in a natural habitat

A natural population of Pycnogonum litorale Ström was examined every 4 weeks over a period of 15 months and thereafter at yearly intervals for 15 years. Adult pycnogonids – mating couples and males carrying egg batches – and freshly hatched protonymphon larvae within these egg batches were found throughout the year. The second, third, and fourth instar larvae were only found from April to July, during the vegetation period of their hydroid host Clava multicornis. After metamorphosis to the fifth instar (first juvenile instar) the pycnogonids have a significantly larger proboscis than during the larval period, and they feed on the sea anemone Metridium senile. First juvenile instars were found on M. senile from May to August. Older and larger juvenile stages were found over longer time spans throughout the year, and the maximum number of successive instars shifted slowly from June to December. Freshly moulted adults occurred throughout the year. Males, which on the average are smaller, usually reach the adult stage during late autumn of the first year and females, at the end of the following spring. We conclude that in nature the development from egg to adult stage is completed within one year. Continuous reproduction and asynchronous embryonic development provide offspring throughout most of the year. The annual cycle is synchronized by the vegetation period of C. multicornis, the only host of these pycnogonid larvae in the investigated habitat, and by the arrest of growth during low winter temperatures. The low level of locomotory activity of P. litorale probably requires an environment in which both host species coexist. The abundance of C. multicornis, M. senile, and juvenile pycnogonids decreased from 1990 to 1996, maybe due to hydrographic conditions.     

Ecological costs on local adaptation of an insect herbivore imposed by host plants and enemies

Herbivore populations may become adapted to the defenses of their local hosts, but the traits that maximize host exploitation may also carry ecological costs. We investigated the patterns and costs of local adaptation in the pine processionary moth, Thaumetopoea pityocampa, to its host plants, Pinus nigra and P. sylvestris. The two hosts differ in needle toughness, a major feeding impediment for leaf-eating insects. We observed a west-to-east gradient of increasing progeny size in the Italian Alps, matching the pattern in toughness of their respective local host plant. Eastern populations that feed on the native P. nigra with tough needles had larger eggs, and neonate larvae with larger head capsules, than western populations that feed on the native P. sylvestris and the introduced P. nigra with softer foliage. In a reciprocal transfer experiment that involved the eastern-most and the western-most populations of T. pityocampa from this region, and excluded natural enemies, we found evidence for local adaptation to the host plant. Specifically, larvae from the western population only performed well when raised on their local hosts with soft needles, and they suffered near-complete mortality on the tough foliage at the eastern site. In contrast, larvae from the eastern population survived equally well at both sites. Local adaptation involved a trade-off between progeny size and the number of offspring. We hypothesized that an additional cost, imposed by natural enemies, may be associated with increased egg size: we also observed a west-to-east gradient of increased egg parasitism. We tested this hypothesis in a common garden by exposing eggs of both populations to parasitism by two native egg parasitoids, Ooencyrtus pityocampae and Baryscapus servadeii. The eastern population suffered a higher level of parasitoid attack by O. pityocampae than the western population, and performance of hatched adults of both parasitoids was enhanced in large eggs. Thus, increased neonate quality (larger eggs yielding larger larvae) confers an advantage on tough foliage but incurs the ecological cost of increased parasitism, which may constrain further adaptation by this herbivore.     

模拟酸雨对杜仲抗性生理及药用有效成分含量的影响

研究了模拟酸雨对杜仲抗性生理及药用成分绿原酸、桃叶珊瑚甙、总黄酮含量的影响.结果表明,春夏秋季在系列模拟酸雨胁迫下,杜仲叶叶液pH值、K 、Ca2 含量随酸雨pH值降低而降低,呈显著正相关;杜仲叶膜保护酶超氧化物歧化酶(SOD)、过氧化氢酶(CAT)、过氧化物酶(POD)和膜保护物质抗坏血酸(AsA)含量在春夏两季随酸雨pH值降低呈单峰曲线型变化,秋季则均随酸雨pH值降低而降低,且呈显著正相关;抗性物质脯氨酸(Pro)含量春夏两季随酸雨pH值降低而升高,呈显著负相关,秋季则在pH3.0和2.5的强酸雨胁迫下急剧降低;膜脂过氧化产物丙二醛(MDA)含量无论春夏秋季均与酸雨pH值呈显著负相关;春季仅pH2.5的强酸雨对杜仲叶绿原酸、桃叶珊瑚甙和总黄酮等药用成分含量影响较大,夏秋两季各药用有效成分则均随酸雨pH值降低而降低,呈显著正相关.图1表2参27     

Patterns of litter disappearance in a northern hardwood forest invaded by exotic earthworms.

A field study was conducted to evaluate the effects of exotic earthworm invasions on the rates of leaf litter disappearance in a northern hardwood forest in southcentral New York, USA. Specifically, we assessed whether differences in litter quality and the species composition of exotic earthworm communities affected leaf litter disappearance rates. Two forest sites with contrasting communities of exotic earthworms were selected, and disappearance rates of sugar maple and red oak litter were estimated in litter boxes in adjacent earthworm-free, transition, and earthworm-invaded plots within each site. After 540 days in the field, 1.7-3 times more litter remained in the reference plots than in the earthworm-invaded plots. In the earthworm-invaded plots, rates of disappearance of sugar maple litter were higher than for oak litter during the first year, but by the end of the experiment, the amount of sugar maple and oak litter remaining in the earthworm-invaded plots was identical within each site. The composition of the earthworm communities significantly affected the patterns of litter disappearance. In the site dominated by the anecic earthworm Lumbricus terrestris and the endogeic Aporrectodea tuberculata, the percentage of litter remaining after 540 days (approximately 17%) was significantly less than at the site dominated by L. rubellus and Octolasion tyrtaeum (approximately 27%). This difference may be attributed to the differences in feeding behavior of the two litter-feeding species: L. terrestris buries entire leaves in vertical burrows, whereas L. rubellus usually feeds on litter at the soil surface, leaving behind leaf petioles and veins. Our results showed that earthworms not only accelerate litter disappearance rates, but also may reduce the differences in decomposition rates that result from different litter qualities at later stages of decay. Similarly, our results indicate that earthworm effects on decomposition vary with earthworm community composition. Furthermore, because earthworm invasion can involve a predictable shift in community structure along invasion fronts or through time, the community dynamics of invasion are important in predicting the spatial and temporal effects of earthworm invasion on litter decomposition, especially at later stages of decay.     

The effect of seasonality on oxidative metabolism in the sea urchin Loxechinus albus

The objective of the present study was to investigate seasonal variations in the oxidative metabolism of the sea urchin Loxechinus albus gonads. The reported spawning period for this species is from September to November. Lipid radical content showed non-significant changes upon the seasons. Ascorbyl radical content and the content of α-tocopherol were lower in samples collected in spring and summer as compared to the values in winter-collected animals. Ascorbate content decreased in samples collected in fall as compared to those collected in winter. For the lipophilic compartment, the lipid radical content/α-tocopherol content ratio is an indicator of oxidative stress. This index increased significantly in tissues during spawning as compared to the values in samples collected during winter. The ascorbyl content/ascorbate content ratio is an indicator of oxidative stress for the hydrophilic milieu. A significant decrease by 66% was determined in tissues from gonads of animals collected in summer as compared to values in animals collected in winter. The data reported here suggest a different profile of response against oxidative stress at the lipophilic and hydrophilic milieus in L. albus gonads.     

Modelling the spatial population dynamics of the green oak leaf roller (Tortrix viridana) using density dependent competitive interactions: Effects of herbivore mortality and varying host-plant quality

Cyclic population dynamics of forest insects with periods of more than two generations have been discussed in relation to a variety of extrinsic and intrinsic forces. In the present study, we employed the selection pressure of density dependent competitive interactions according to Witting's equations (Witting, 2000) as driver for a discrete spatiotemporal model of the green oak leaf roller (Tortrix viridana). The model was successfully parameterised to rebuild the cyclic population dynamics of an empirical data set of a 30-year leaf roller monitoring in Russia. Our analysis focussed on the role of herbivore mortality and host plant food quality, which have a significant effect on T. viridana population dynamics. An additional egg or larvae mortality lowers population density and can lead to selection pressures that favour individuals with higher growth rate. This increased population growth rate can not only compensate the additional mortality, but also can lead to higher average moth abundances in subsequent generations. Furthermore, we analysed the effect of inter- and intraspecific variation in host plant quality on herbivore population dynamics and the spatial distribution of abundance and defoliation patterns. We found significant effects of the qualitative composition of a trees neighbourhood on the herbivore population of the respective tree. Also, the patchy damage patterns observable in reality have been reproduced by the present model. The applicability of the model approach and the putative genetic processes underlying Witting's model are discussed.     

Chemical defenses, nutritional quality, and structural components in three sponge species: Ircinia felix, I. campana, and Aplysina fulva

Allocating chemical defenses to regions or tissues most at risk for predatory attack may provide protection while simultaneously minimizing associated metabolic costs. Chemical defense allocation patterns were investigated in the aspiculate sponges Ircinia felix, I. campana, and Aplysina fulva collected between July 2005 and April 2006 from J Reef off the coast of Georgia, U.S.A. It was predicted that chemical defenses would be (1) higher in the outermost 2 mm layer of the sponge; (2) positively correlated with tissue nutritional quality; and (3) correlated with structural components such as spongin fibers. Whereas defensive chemicals were concentrated in the outer 2 mm of A. fulva, the Ircinia species had higher concentrations in deeper tissue layers. Furthermore, no significant positive or negative correlation between chemical defenses and nutritional quality or levels of structural components was observed in these sponges. Overall, these results do not support the prediction that predation pressure by fish and large mobile invertebrates significantly impacts chemical defense allocation in these sponges.     

Fluctuating asymmetry and nutritional condition of Baltic cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) larvae

The level of fluctuating asymmetry (FA), defined as random deviations from perfect bilateral symmetry, is assumed to reflect the developmental instability (DI) of an organism. Because environmental and genetic stress may increase DI, FA has been used to assess the level of stress experienced by, for example, fish. In this study, left–right asymmetry of lapillar otoliths was related to nutritional condition as estimated from RNA/DNA ratios, in order to investigate the utility of FA to detect feeding-related stress in Baltic cod, Gadus morhua L., larvae. Cod larvae in intermediate and good nutritional condition showed similar values of FA, and these were more symmetric than for larvae in poor condition. As increased levels of FA were restricted to larvae in a condition comparable to that of larvae experiencing at least 3 days of starvation in laboratory experiments, it is suggested that FA is an insensitive indicator of short-term feeding success of larval cod. However, FA can be used to reveal severely starved larva populations and probably also populations that have been subjected to prolonged sub-optimal feeding conditions.     

Combining optimal defense theory and the evolutionary dilemma model to refine predictions regarding plant invasion

Optimal defense theory posits that plants with limited resources deploy chemical defenses based on the fitness value of different tissues and their probability of attack. However, what constitutes optimal defense depends on the identity of the herbivores involved in the interaction. Generalists, which are not tightly coevolved with their many host plants, are typically deterred by chemical defenses, while coevolved specialists are often attracted to these same chemicals. This imposes an "evolutionary dilemma" in which generalists and specialists exert opposing selection on plant investment in defense, thereby stabilizing defenses at intermediate levels. We used the natural shift in herbivore community composition that typifies many plant invasions to test a novel, combined prediction of optimal defense theory and the evolutionary dilemma model: that the within-plant distribution of defenses reflects both the value of different tissues (i.e., young vs. old leaves) and the relative importance of specialist and generalist herbivores in the community. Using populations of Verbascum thapsus exposed to ambient herbivory in its native range (where specialist and generalist chewing herbivores are prevalent) and its introduced range (where only generalist chewing herbivores are prevalent), we illustrate significant differences in the way iridoid glycosides are distributed among young and old leaves. Importantly, high-quality young leaves are 6.5x more highly defended than old leaves in the introduced range, but only 2x more highly defended in the native range. Additionally, defense levels are tracked by patterns of chewing damage, with damage restricted mostly to low-quality old leaves in the introduced range, but not the native range. Given that whole-plant investment in defense does not differ between ranges, introduced mullein may achieve increased fitness simply by optimizing its within-plant distribution of defense in the absence of certain specialist herbivores.     

Recruitment,growth, and diel vertical migration of Euphausia pacifica in a temperate fjord

The pelagic crustacean Euphausia pacifica Hansen was sampled with a multiple-sample 1.0 m² Tucker trawl and a multiple-sample 1.0 m² vertical net in Dabob Bay, Washington on 17 dates between May 1985 and October 1987. Size (stage) structure and abundance of the population were determined for each date, while vertical distribution and diel migration were determined for 13 dates. Although internannual variability in both timing and magnitude of events occurred, consistent patterns were discernable. The population produced a large pulse of larvae (2 to 5 mm) in late spring of each year, apparently in response to the vernal phytoplankton bloom. Much lower abundances of larvae occurred during summer and autumn of each year, and larvae were completely absent during winter. Recruitment to the juvenile (6 to 9 mm) and adult (10 mm) stages was strongest during the summer, with abundances of these individuals peaking in summer and autumn. Individual growth rates, determined by modal progression analysis, were calculated for E. pacifica. Rates ranged from zero for some adult cohorts during the winter to 0.12 mm d^-1 for larvae during spring. The latter are among the highest ever reported for this species in the field. The vertical distributions and diel vertical migrations (DVM) of E. pacifica varied seasonally and between size (stage) classes. At night, all size classes were distributed in the surface layer (upper 25 m) irrespective of season or year. During the day, the larger/older stages were always distributed at middepths (50 to 125 m). In contrast, the daytime distribution of the larvae was more variable, being concentrated at the surface during spring and early summer of 1985, and at increasing depths later in the summer and autumn of 1985 and again in spring of 1986. This resulted in invariant DVM in the juveniles and adults, but variable DVM in the larvae, the latter of which is hypothesized to be a response to variable abundances of zooplanktivorous fish.     

转sck单价基因和cry1Ac/sck双价基因水稻对二化螟的致死效应及中肠组织病理变化观察

室内采用离体叶片法研究了转sck单价基因抗虫水稻86AS1和转cry1Ac/sck双价基因抗虫水稻MSB不同生育期主茎顶叶对二化螟幼虫的杀虫效果.结果发现,不同生育期的MSB对二化螟初孵幼虫表现出了很强的毒杀效果,二化螟幼虫校正死亡率在扬花期前均达到90%以上;扬花期和灌浆期抗虫性有所下降,校正死亡率高于80%;成熟期校正死亡率在60%以上.在整个生育期,二化螟幼虫取食86AS1主茎顶叶的校正死亡率均未超过50%.MSB对二化螟幼虫的毒杀效果明显高于86AS1.采用透射电镜观察二化螟幼虫取食转cry1Ac/sck双价基因抗虫水稻和转sck单价基因抗虫水稻主茎顶叶后中肠组织的病理变化.结果发现,两种转基因水稻都能够引起二化螟幼虫中肠组织发生病理变化,但病变程度有明显差异.二化螟幼虫取食转cry1Ac/sck双价基因抗虫水稻3d,中肠组织受到严重破坏且病理变化明显;取食转sck单价基因抗虫水稻3d,幼虫中肠组织的病理变化却较缓慢.图2参10     

Food quality, competition, and parasitism influence feeding preference in a neotropical lepidopteran

We surveyed Lepidoptera found on 11 species of Inga (Fabaceae:Mimosoideae) co-existing on Barro Colorado Island, Panama, to evaluate factors influencing diet choice. Of the 47 species of caterpillars (747 individuals) recorded, each fed on a distinct set of Inga. In the field, 96% of the individuals were found on young leaves. Growth rates of caterpillars that were fed leaves in the laboratory were 60% higher on young leaves compared to mature leaves. When caterpillars were fed leaves of nonhost Inga, they grew more slowly. These data provide support for a link between preference and performance. However, among hosts on which larvae normally occurred, faster growth rates were not associated with greater host electivity (the proportion of larvae found on each host species in the field, corrected for host abundance). Growth rates on normal hosts were positively correlated with leaf expansion rates of the host, and fast expansion was associated with leaves with higher nutritional content. Detailed studies on a gelechiid leaf roller, the species with the largest diet breadth, allowed us to assess the importance of factors other than growth that could influence diet electivity. This species showed a 1.7-fold difference in growth rate among Inga hosts and faster growth on species with fast-expanding leaves. However, there was no correlation between caterpillar growth rate and abundance on different host species. Instead, abundance of the gelechiid on each Inga species was significantly correlated with the temporal predictability of food (synchrony of leaf flushing) and was negatively correlated with competition (amount of leaf area removed by species other than the gelechiid). Although rates of parasitism were high (23-43%), there were no differences among hosts. Parasitism was also not related to measures of escape, such as growth rates of caterpillars, leaf expansion rates, and synchrony of leaf production. Together, food availability, parasitism, and competition explained 84% of the variation in host preference by the gelechiid. We suggest that these ecological interactions may be particularly important in determining diet choice initially and that preferences may be reinforced by subsequent divergence in host chemistry and/or the herbivore's ability to tolerate the secondary metabolites.