Physiological mechanisms underlying the costs of chemical defence in Junonia coenia Hu¨bner (Nymphalidae): A gravimetric and quantitative genetic analysis

Several recent studies document that specialist insect phytophages may be less subject to predation than generalists and suggest that hostplant-derived chemical defences may be an important explanation for the predominance of specialized feeding among insect herbivores. The evolution of such chemical defences depends upon both their advantages versus natural enemies and their physiological costs, but data on these costs, particularly genetic data, are few. Here I report the results of an ecological genetic investigation of food use efficiency and allelochemical sequestration in Junonia coenia Hu¨bner (Nymphalidae). I used standard gravimetric techniques to estimate the efficiency of dry matter incorporation and iridoid glycoside sequestration in the larvae of 37 full-sib families fed artificial diets containing trace, low (2%) and high (10%) concentrations of iridoid glycosides. I found a significant reduction in the efficiency of dry matter incorporation on a high iridoid diet that is entirely attributable to reduced digestibility rather than post-digestive toxic effects. Larvae fed high-iridoid diets sequestered them less efficiently, but this difference was due largely to post-digestive effects. Analyses of genetic variation and architecture of dry matter and iridoid budgets reveal substantial genetic variation in both suites of traits, but only chemical defence showed a significant genotype×environment interaction which would be conducive to the evolution of specialization. Neither group of traits showed across-diet trade-offs in the form of negative correlations of family means among diets. Family means correlations of sequestration indices with dry matter indices within diets reveals that chemical defence comes at a cost to growth, but only in the high diet. I also found evidence of specialized physiological machinery for iridoid glycoside processing. These data indicate that even adapted specialists are negatively affected by plant toxins, but in this species, dietary specialization is more likely to result from selection from natural enemies than from hostplant toxins.     

THE GENETICS AND COST OF CHEMICAL DEFENSE IN THE TWO-SPOT LADYBIRD (ADALIA BIPUNCTATA L.)

Ladybirds (Coccinellidae) defend themselves against attack by vertebrate predators by exuding a fluid from the femero-tibial joints. This fluid carries a noxious or toxic alkaloid. The amount of fluid produced during a single attack can be very high (up to 20% of fresh body weight), and the weight of the self-synthesized alkaloid can amount to several percent of the weight of the fluid. A study was carried out on these two defense characters and two other fitness characters (body weight and growth rate) to demonstrate a cost to defense in the form of genetic trade-offs between characters. The two sexes were analyzed separately, and a jackknife procedure was used to attach errors to the estimates of V_a and cov_a. All four characters were associated with high levels of V_a, but the cov_a values were mixed, some being negative and others positive. Principal-component analysis indicated the operation of factors constraining the cov_a values in males, and further possible reasons for the appearance of so many positive values are explored. A matrix analysis showed that the genetic variance/covariance matrices of the two sexes were significantly different from each other. Breeding values derived from sons plotted on breeding values from daughters had correlation coefficients significantly less than +1. This finding indicated that a substantial amount of sex-dependent gene expression was occurring.     

Host plant selection and development in Spodoptera exigua: do mother and offspring know best?

We examined the ovipositional preference and larval development of Spodoptera exigua (Hübner) (Lepidoptera: Noctuidae) on two common hosts in southern California, Chenopodium murale L. (Chenopodiaceae) and Apium graveolens L. (Umbelliferae) to determine if female oviposition preference is correlated with offspring performance. Greenhouse oviposition choice tests indicated that S. exigua oviposit more frequently on C. murale than on A. graveolens. However under laboratory conditions, larvae reared on C. murale had longer development times, lower relative growth rate, and lower survivorship than larvae reared on A. graveolens. larval and pupal masses were significantly greater on A. graveolens than on C. murale. Furthermore, pupal masses were significantly greater for individuals reared on A. graveolens than on C. murale. Because pupal masses and adult fecundity are positively correlated for Spodoptera spp., the fitness of S. exigua on A. graveolens is likely to be substantially higher than its fitness on C. murale. Despite better larval performance on A. graveolens, previous results from choice tests with whole plants and leaf discs indicate that the highly mobile S. exigua larvae strongly prefer C. murale over A. graveolens. Hypotheses attempting to explain this lack of correlation between larval and adult host preference versus development and survival in this system are discussed.     

The cabbage aphid: a walking mustard oil bomb

The cabbage aphid, Brevicoryne brassicae, has developed a chemical defence system that exploits and mimics that of its host plants, involving sequestration of the major plant secondary metabolites (glucosinolates). Like its host plants, the aphid produces a myrosinase (beta-thioglucoside glucohydrolase) to catalyse the hydrolysis of glucosinolates, yielding biologically active products. Here, we demonstrate that aphid myrosinase expression in head/thoracic muscle starts during embryonic development and protein levels continue to accumulate after the nymphs are born. However, aphids are entirely dependent on the host plant for the glucosinolate substrate, which they store in the haemolymph. Uptake of a glucosinolate (sinigrin) was investigated when aphids fed on plants or an in vitro system and followed a different developmental pattern in winged and wingless aphid morphs. In nymphs of the wingless aphid morph, glucosinolate level continued to increase throughout the development to the adult stage, but the quantity in nymphs of the winged form peaked before eclosion (at day 7) and subsequently declined. Winged aphids excreted significantly higher amounts of glucosinolate in the honeydew when compared with wingless aphids, suggesting regulated transport across the gut. The higher level of sinigrin in wingless aphids had a significant negative impact on survival of a ladybird predator. Larvae of Adalia bipunctata were unable to survive when fed adult wingless aphids from a 1% sinigrin diet, but survived successfully when fed aphids from a glucosinolate-free diet (wingless or winged), or winged aphids from 1% sinigrin. The apparent lack of an effective chemical defence system in adult winged aphids possibly reflects their energetic investment in flight as an alternative predator avoidance mechanism.