Density-dependent induction and suppression of soldier differentiation in an aphid social system

We investigated the mechanism underlying the control of soldier production in colonies of a social aphid, Tuberaphis styraci, which has a sterile soldier caste in the second instar. High aphid density was shown to induce soldier production in T. styraci. Analysis of natural colonies revealed that the soldier proportion tended to increase with aphid density but reached a plateau. Artificial diet experiments identified a similar plateau of soldier proportion under high-density conditions. In order to gain insights into the controlling mechanism of soldier production, the effect of soldiers on reproducing adult aphids was examined using the artificial diet system. It was experimentally demonstrated that soldier production was suppressed by coexisting soldiers, whereas coexisting non-soldiers facilitated soldier production. These results suggested that caste ratio in the colony of T. styraci is controlled by positive and negative feedbacks consisting of density-dependent induction and suppression of soldier differentiation.     

Density triggers soldier production in a social aphid

For evolution and maintenance of the social system of insect colonies, investment in the sterile caste should be adequately controlled in response to environmental cues. Recent developments using artificial diet rearing techniques have revealed an underlying mechanism of caste control in a gall-forming aphid, Tuberaphis styraci, which has a soldier caste in the second instar. Statistical analyses of field-collected galls detected a significant positive correlation between aphid density and soldier proportion in the natural colonies of T. styraci. Artificial diet experiments showed that soldiers are produced under crowded conditions. Detailed experiments demonstrated that soldiers are produced in a density-dependent manner rather than in a colony size-dependent manner. From these results, it was concluded that aphid density is the crucial cue that triggers soldier production in T. styraci. This study provides, to our knowledge, the first experimental demonstration of an environmental factor involved in aphid soldier differentiation.     

Social aphids use their antennae to perceive density cue for soldier production

In social insects, local interactions among colony members facilitate information transfer, and allow the whole colony to regulate division of labor and task allocation in an integrated and coordinated manner. In particular, regulation of caste differentiation in response to external cues is important for sustaining social insect colonies. The social aphid Tuberaphis styraci exhibits a caste polyphenism, producing second‐instar soldiers and non‐soldiers. Previous studies using an artificial diet rearing system identified high aphid density as a crucial cue for soldier production, which acts on embryos in the maternal body and newborn first‐instar nymphs to induce soldier differentiation. While direct contact stimuli from live non‐soldiers were suggested to mediate the density effect, how the aphids perceive the stimuli has been unknown. Here we investigated how antennal removal of adult females affects the soldier production in T. styraci. Under a high density condition, intact females produced the highest percentage of soldiers, females deprived of both antennae produced the lowest percentage of soldiers, and females deprived of one antenna exhibited an intermediate percentage of soldiers. Scanning electron microscopic observations of the aphids revealed the existence of sensory organs for chemoreception and tactile sensation on the antennae of the mother aphids. These results indicate that T. styraci females use their antennae to perceive soldier‐inducing density cue, suggesting that maternal perception of density cue is involved in regulation over caste phenotype of their offspring.     

Reproductive schedule and factors affecting soldier production in the eusocial bamboo aphid Pseudoregma bambucicola (Homoptera, Aphididae)

Summary: The reproductive characteristics of the soldier-producing aphid Pseudoregma bambucicola were studied in Kagoshima, Southern Japan, to know the factors affecting soldier production of eusocial aphids. The soldier proportion in aphid colonies was highest from October to November. In some large colonies, soldiers were observed in all seasons except in July when colony size was relatively small. Multiple regression analysis showed that the colony size was a principal factor affecting soldier proportion throughout a year. Other social or environmental factors such as aphid composition, host plant conditions and predator abundance were not always significant. Rearing experiments revealed that large colonies (̿,000 individuals) produced soldiers in almost all seasons while small colonies (<1,000) never produced any soldiers. The caste-production schedule of adult females was examined in the field. When solitary females produced both castes, they usually produced normal nymphs first and then soldiers. Females from large colonies tended to produce more soldiers in the earlier period of their lifetime, whereas females from newly established small colonies produced no or only a few soldiers at later times. The average number of soldiers and normal nymphs produced consecutively by a single female was >10 and >20, respectively. Because they have a small number of ovarioles (<15 on average), females should alter caste production within the same ovarioles according to changes in environmental conditions. Artificial removal or introduction of predators and reduction of colony size did not affect soldier production over two successive generations, revealing maternal effects on soldier production. Females cannot shift caste production quickly in response to changes in predator abundance and colony size. This is probably due to early developmental determination of castes within the mother's body.     

Maintenance of soldier-producing aphids on an artificial diet

In our efforts to understand the biology of soldier-producing aphids, we attempted to maintain them in the laboratory using a chemically defined artificial diet. The ability of 16 species from the subfamilies Eriosomatinae and Hormaphidinae, most of which are soldier-producing species, to survive on the artificial diet was examined. Some species neither fed nor grew on the diet, whereas other species accepted the diet, grew to some extent, and managed to produce a small number of short-lived offspring. Although they performed poorly on the diet in general, aphid performance was correlated with the stage in the life cycle and the developmental stadium in that aphids of the gall generation tended to accept the diet and survive on it, whereas aphids of the non-gall generation did not. Also, old insects tended to perform better on the diet than young nymphs. Notably, only one species, Tuberaphis styraci, a gall-forming aphid that produces 2nd instar sterile soldier, showed good performance on the diet. Insects collected from galls (generation G0) survived on the diet, grew well, and produced many progeny. Three successive generations (G1, G2 and G3) were produced on the diet. Developmental period, adult body size, and age of first reproduction were almost constant through G0, G1 and G2 whereas fecundity, adult longevity and daily offspring production declined as the generations proceeded. These results are comparable to previous studies in which pest aphids have been maintained on similar artificial diets for several generations. Therefore, it is suggested that the artificial-diet rearing system will provide a useful tool to investigate various biological aspects of the soldier-producing eusocial aphid, T. styraci.     

Comparative analysis of caste differentiation during embryogenesis of social aphids whose soldier castes evolved independently

Summary. We investigated the morphological characters of normal nymphs, soldier nymphs and developing embryos of a social aphid, Colophina arma, which has a sterile soldier caste in the first instar. Morphometric analysis revealed that normal nymphs and soldier nymphs were clearly distinguishable on the basis of several morphological characters. At late embryonic stages, normal embryos and soldier embryos were also distinguishable morphologically. The younger the embryonic stages, the smaller the morphological differences between them. In young embryos of less than 600 m in body length, normal embryos and soldier embryos were no more distinguishable, suggesting that the onset of soldier differentiation occurs at an early embryonic stage. Through the embryonic development, morphological differentiation of soldier caste proceeded gradually: forelegs and midlegs were exaggerated, and growth of mouthpart was suppressed, which resulted in the soldier morphology specialized for attacking behavior. On the basis of these results, developmental aspects in soldier differentiation of C. arma were compared with those of Pseudoregma bambucicola, a social aphid with a first instar soldier caste of independent evolutionary origin. Ecological and evolutionary relevance of the differences between the two social aphids was discussed.Received 30 June 2004; revised 20 October 2004; accepted 9 November 2004.     

Cultivar‐specific plant odour preferences of a generalist aphid parasitoid Aphidius colemani and a possible mechanism for maternal priming of resistance to toxic plant chemistry

Aphidius colemani Viereck, emerging from Myzus persicae (Sulzer) mummies on the Brussels sprout cultivar ‘Bedford Winter Harvest’ (BWH), responds positively in the olfactometer to the odour of that cultivar in comparison with air. Responses to the odours of other sprout cultivars, cabbage and broad bean could be explained by the humidity from plant leaves. In a choice between BWH and other sprout cultivars, the BWH odour is preferred, or that of cv. ‘Red Delicious’ (RD) if the parasitoids are reared on RD. This confirms previous work showing that the secondary chemistry of a cultivar is learnt from the mummy cuticle during emergence. Adults emerging from pupae excised from the mummy show a similar but less pronounced preference. Parasitoids developing in aphids on an artificial diet do not discriminate between the odours of BWH and RD, unless allowed contact with a mummy from the same cultivar that the mother develops on. This suggests a cultivar‐specific maternal cue. This cue is speculated to consist of a small amount of the secondary chemistry (probably glucosinolates in the present study) that are left in or on the egg at oviposition, which subsequently induces enzymes that detoxify plant‐derived toxins in the aphid host. Indeed, when parasitoids emerging from diet‐reared aphids are released on aphid‐infested sprout plants, fewer mummies are produced than by parasitoids emerging from mummies of plant‐reared aphids or from excised pupae. Only parasitoids that emerge from mummies of plant‐reared aphids prefer the cultivar of origin as shown by the number of mummified hosts.     

Ant tending influences soldier production in a social aphid

The aphid Pseudoregma sundanica (Van der Goot) (Homoptera: Aphididae) has two defence strategies. It is obligatorily tended by various species of ant and also produces sterile soldiers. We investigated how they allocate their investment in these two strategies. We measured the size, number of soldiers, number and species of tending ant, and number and species of predators in P. sundanica populations. We found that the level of ant tending correlated negatively with soldier investment in P. sundanica. The species of tending ant also influenced soldier investment. We excluded ants from aphid populations and recorded changes in population size and structure over four weeks. Ant exclusion led to population decline and extinction. At the same time, surviving populations showed a significant increase in soldier investment. The data demonstrate that social aphids can adjust their investment in soldiers in direct response to environmental change.     

Soldier differentiation during embryogenesis of a social aphid, Pseudoregma bambucicola

To understand the developmental process of aphid soldier differentiation, we investigated the morphological characters of normal nymphs, soldier nymphs and developing embryos of Pseudoregma bambucicola. Results of morphometric analyses showed that normal and soldier nymphs formed discrete clusters on the basis of several morphological characters, although a small number of intermediate individuals, termed ‘intercaste nymphs’, were present. In late embryonic stages, normal and soldier embryos were morphologically distinguishable. The earlier the embryonic stage, the smaller the morphological differences between them. In early embryos less than 1000 µm in length, normal and soldier embryos were not morphologically distinguishable, suggesting that the onset of soldier differentiation occurs at an early embryonic stage. Throughout embryonic development, morphological differentiation of the soldier caste proceeded gradually. Notably, several morphological characters of soldiers grew remarkably upon larviposition. Observation of embryonic leg cuticle revealed a characteristic folding structure, indicating that some morphological traits of the soldier are exaggerated upon larviposition through expansion of the folded cuticle. We suggest that morphological differentiation of the soldier caste in P. bambucicola comprises two phases: gradual growth during embryogenesis and rapid growth upon larviposition.     

The evolutionary biology of sterile soldiers in aphids

The discovery of a sterile soldier caste in aphids by Shigeyuki Aoki offered behavioural ecologists new and interesting subjects for the study of insect sociality. Quantitative studies on the behaviour of soldiers and the population and genetic consequences are, however, scant. This review summarizes the distribution and peculiarities of sterile soldiers in different aphid taxa and also reports on phenomena that may relate to the evolution of aphid soldiers.     

Soldiers with large weapons in predator-abundant midsummer: phenotypic plasticity in a eusocial aphid

Eusocial aphids produce sterile individuals (“soldiers”) that specialize behaviorally and morphologically to protect their colony from predators, while production of soldiers can negatively affect colony growth because of reproductive allocation and opportunity cost. Hence, a cost-saving soldier production strategy is expected to be favored. Here, we hypothesize that, to save the cost, a eusocial aphid Ceratovacuna japonica produces soldiers with smaller weapon in the season when predators are not abundant. The abundance of two specialist lepidopteran predators (i.e., Taraka hamada and Atkinsonia ignipicta) of C. japonica dramatically increased, and aphid colony size significantly decreased, from July to August. In line with these, the soldiers in August had larger weapons (i.e., frontal horns) than those in June, indicating a correlational increase in weapon size with predation pressure. We predict that a reliable prospective signal indicating the coming of midsummer (environmental temperature) induces mother aphids to produce soldiers with larger weapons. Experiments clarified that soldiers produced at 20 °C (typical temperature of July to August) had larger weapons than those produced at 15 °C (typical temperature of May to July). Such phenotypic plasticity appears to be adaptive to maximize the fitness of C. japonica under a temporally variable but predictable predation environment. These results indicate that C. japonica aphids not merely have distinctive reproductive—and soldier castes, but also produce differentially armed soldiers in a habitat with temporally changing predation risks.     

Evolution of soldier-specific venomous protease in social aphids

In social aphids of the genus Tuberaphis a cysteine protease gene of the family cathepsin B exhibits soldier-specific expression and intestinal protease production. The product is orally excreted and injected by soldier nymphs into natural enemies, thereby exerting an insecticidal activity. In an attempt to gain insights into when and how the novel venomous protease for the altruistic caste has evolved, we investigated the soldier-specific type (S-type) and nonspecific type (N-type) cathepsin B genes from social and nonsocial aphids. All the social aphids examined, representing the genera Tuberaphis, Astegopteryx, and Cerataphis, possessed both the S-type and N-type genes. Phylogenetically distant nonsocial aphids also possessed cathepsin B genes allied to the S-type and the N-type, indicating the evolutionary origin of these genes in the common ancestor of extant aphids. In Tuberaphis species the S-type genes exhibited significant soldier-specific expression and accelerated molecular evolution whereas the N-type genes did not. In Astegopteryx and Cerataphis species, meanwhile, both the S-type and N-type genes exhibited neither remarkable soldier-specific expression nor accelerated molecular evolution. These results suggest that the S-type gene acquired the soldier-specific expression and the venom function after divergence of the genus Tuberaphis. On the structural model of the S-type protease of Tuberaphis styraci the accelerated molecular evolution was associated with the molecular surface rather than the catalytic cleft, suggesting that the venom activity was probably acquired by relatively minor modifications on the molecular surface rather than by generation of a novel active site. In Cerataphis jamuritsu the S-type gene was, although containing a stop codon, structurally almost intact and still transcribed, suggesting recent pseudogenization of the gene copy and possible relevance to relaxed functional constraint in the highly multiplied protease gene family. On the basis of these results we suggest that the massive amplification in aphid cathepsin B genes might have predisposed the evolution of venomous protease in the social aphid lineage and argue that gene duplication, accelerated molecular evolution, and acquisition of novel gene function must have played considerable roles in the evolution of complex biological systems including insect sociality.     

Clonal mixing in the soldier-producing aphid Pemphigus spyrothecae (Hemiptera: Aphididae)

Illuminating the genetic relationships within soldier-producing aphid colonies is an essential element of any attempt to explain the evolution of the altruistic soldier caste. Pemphigus spyrothecae is a soldier-producing aphid that induces galls on the leaf petioles of its host (trees of the genus Populus). At least a quarter of the aphids within the clonally produced gall population are morphologically and behaviourally distinct first-instar soldiers that defend the gall population from predation. Using field trapping and microsatellites, we investigated the degree of clonal mixing within natural gall populations. Field trapping in the UK showed that all the migrants of P. spyrothecae and of two other Pemphigus species were wingless first-instar soldiers. The average degree of mixing estimated from trapping P. spyrothecae migrants was 0.68% (range = 0-15%). Microsatellite genotyping of 277 aphids from 13 galls collected in Italy revealed an average mixing level of 10.4% (range = 0-59%). Six galls contained more than one clone (range = 2-5 clones). Non-kin aphids were not restricted to the soldier caste but were evenly distributed across instars. An additional gall, from which 527 occupants were genotyped, contained 12 non-kin aphids distributed among nine clones, showing that clonal diversity can be high even when mixing is very low. These observations suggest that although soldiers migrate regularly and can moult and reproduce within foreign galls, clonal mixing in this species is generally low and is unlikely to provide a barrier to the evolution of investment by the aphid clones in an altruistic soldier caste.     

Salicylic acid-mediated reductions in yield in Nicotiana attenuata challenged by aphid herbivory

Aphid herbivory decreases primary production in natural ecosystems and reduces crop yields. The mechanism for how aphids reduce yield is poorly understood as some studies suggest aphid feeding directly impedes photosynthesis, whereas other studies suggest a change in allocation of resources from growth to defense compounds reduces yield. To determine the mechanisms underlying reduced plant growth by aphids, Nicotiana attenuata plants, native tobacco, were infested with Myzus persicae ssp. nicotianae, tobacco-adapted green peach aphids, at low and high densities, and plant performance including fitness was assessed. To test the direct defense capacity of salicylic acid (SA) on aphid performance, we fed aphids an artificial diet with varying levels of SA and measured their survivorship and fecundity. There was no detectable effect of aphid herbivory on net photosynthesis, yet herbivory reduced plant growth, final biomass (43 % at high aphid density), and seed set (18 % at high aphid density) at both low and high aphid infestation levels. High-density aphid attack during the rosette and flowering stage caused an increase in SA levels, but caused only a transient decrease in jasmonic acid concentration at low aphid density. SA concentrations similar to those found in infested flowering plants decreased aphid fecundity, suggesting that SA was an effective chemical defense response against aphids. These results suggest that as aphid densities increased the proximal cause of reduced growth and yield was not reduced photosynthesis, but instead resources may have been mobilized for defense via the SA pathway, decreasing the availability of resources for building plant biomass.     

Direct and indirect sublethal effects of Galanthus nivalis agglutinin (GNA) on the development of a potato-aphid parasitoid, Aphelinus abdominalis (Hymenoptera: Aphelinidae)

Snowdrop lectin (Galanthus nivalis agglutinin, GNA), has been shown to confer partial resistance to two potato aphids Myzus persicae and Aulacorthum solani, when incorporated in artificial diet and/or expressed in transgenic potato. First-tier laboratory-scale experiments were conducted to assess the potential effect of GNA on the aphid parasitoid Aphelinus abdominalis. GNA (0.1% w/v) was successfully delivered to Macrosiphum euphorbiae via artificial diet and induced a reduced growth rate and increased mortality compared to aphids fed a control diet. As aphid parasitoid larvae are endophagous, they may be exposed to GNA during their larval development and potential "chronic toxicity" on A. abdominalis was investigated. The amounts of GNA present in aphid and parasitoid tissues were estimated by western blotting. Results suggest that parasitoids excrete most of the GNA ingested. Sublethal effects of GNA on several parasitoid fitness parameters (parasitism success, parasitoid development and size, emergence success, progeny survival and sex ratio) were studied. No direct detrimental effect of GNA on A. abdominalis was observed. However, GNA had an indirect host-size-mediated effect on the sex ratio and the size of parasitoids developing in GNA-fed aphids. This work highlights the need to determine the exact "causes and effects" when assessing the ecological impact of transgenic plants on non-target beneficial insects. Such bioassays form the basis of a tiered risk assessment moving from laboratory studies assessing individuals towards field-scale experiments assessing populations.     

Evaluation of artificial diets for rearing Aphis glycines (Hemiptera: Aphididae)

Artificial aphid diets have been previously developed for the pea aphid, Acyrthosiphon pisum (Harris), and the green peach aphid, Myzus persicae (Sulzer). The ability to rear aphids on an artificial diet allows for selectively adding or subtracting compounds from an aphid's food source to determine the effect on fecundity and longevity. Five diets previously developed for the green peach aphid and the pea aphid were tested for their suitability for rearing soybean aphid, Aphis glycines Matsumura. The best diet, originally developed for the green peach aphid and based on the amino acid profile of young potato plants, allowed 12 generations of soybean aphids to develop. For all diets tested, aphid fecundity, and longevity were greatly reduced in comparison with aphids reared on soybean, Glycine max (L.) Merr., plants or on detached soybean leaves. In addition, mean developmental time was significantly longer for aphids reared on artificial diets.     

Soldier caste with morphological and reproductive division in the aphid tribe Nipponaphidini

Summary. We investigated demographic, morphological and histological aspects of Distylaphis foliorum, a gall-forming nipponaphidine aphid from Java, Indonesia, whose first instar nymphs had been reported to have enlarged forelegs and attack other insects. The gall inhabitants of D. foliorum consisted of two discrete populations; one developing normally and the other remaining at the first instar. Morphometric analysis identified two types of first instar nymphs; nymphs with relatively long forelegs and slender abdomen, and nymphs with relatively short forelegs and fat abdomen. The former nymphs were found from both young galls and mature galls, while the latter nymphs were from young galls only. Histological analysis strongly suggested a reproductive division in the first instar. In the former nymphs from mature galls, ovaries and mycetomes were degenerate and replaced by well-developed fat body cells. These results indicated that D. foliorum has a morphologically differentiated sterile soldier caste in the first instar. This study is the first report of a soldier caste with morphological and reproductive division in the aphid tribe Nipponaphidini, and suggests that highly specialized soldier castes have evolved at least four times in aphids.Received 25 June 2004; revised 24 September 2004; accepted 11 October 2004.     

Effect of nymphal diet on adult predation behavior in Orius majusculus (Heteroptera: Anthocoridae)

The predatory bug Orius majusculus (Reuter) was reared on 2 different diets during the nymphal stages. The 1st group was exclusively offered eggs of Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), a standard diet for O. majusculus production. The 2nd group was exclusively offered 4th instars of the pea aphid, Acyrtosiphon pisum (Harris). Subsequently, adult predatory behavior in experimental arenas containing A. pisium was recorded using 2 video cameras. One camera permitted observation of the predator's contact with the prey, where the 2nd camera viewed the arena from above to record the path taken by O. majusculus adults before and after contact with prey. When O. majusculus were reared on aphids, adult bugs successfully located and consumed 55% of experimental prey and continued prey search behavior after each aphid meal. O. majusculus adults that had no experience of aphid predation as nymphs, did not prey on aphids in the experimental arena. The mean walking speed of this group of predators increased from 5.9 +/- 1.2 mm/s to 9.8 +/- 0.7 mm/s after contact or detection of prey, indicating that predators rapidly moved away from unfamiliar prey. Moreover, for egg-reared O. majusculus, all contacts between aphid and predator were lateral, along the side of the prey and were effectively repelled by an aphid kicking response. In contrast, 83% of attacks by aphid-reared O. majusculus were directed at the head or posterior abdomen for which the prey could not defend themselves adequately. When egg-reared O. majusculus were exposed to novel aphid prey for 1-8 d, the frequency of aphid attack increased significantly. We conclude that the standard diet used for rearing O. majusculus may adversely affect the efficiency of this predator as an agent of biological control.     

Direct and Indirect Impacts of Infestation of Tomato Plant by Myzus persicae (Hemiptera: Aphididae) on Bemisia tabaci (Hemiptera: Aleyrodidae)

The impacts of infestation by the green peach aphid (Myzus persicae) on sweetpotato whitefly (Bemisia tabaci) settling on tomato were determined in seven separate experiments with whole plants and with detached leaves through manipulation of four factors: durations of aphid infestation, density of aphids, intervals between aphid removal after different durations of infestation and the time of whitefly release, and leaf positions on the plants. The results demonstrated that B. tabaci preferred to settle on the plant leaves that had not been infested by aphids when they had a choice. The plant leaves on which aphids were still present (direct effect) had fewer whiteflies than those previously infested by aphids (indirect effect). The whiteflies were able to settle on the plant which aphids had previously infested, and also could settle on leaves with aphids if no uninfested plants were available. Tests of direct factors revealed that duration of aphid infestation had a stronger effect on whitefly landing preference than aphid density; whitefly preference was the least when 20 aphids fed on the leaves for 72 h. Tests of indirect effects revealed that the major factor that affected whitefly preference for a host plant was the interval between the time of aphid removal after infestation and the time of whitefly release. The importance of the four factors that affected the induced plant defense against whiteflies can be arranged in the following order: time intervals between aphid removal and whitefly release > durations of aphid infestation > density of aphids > leaf positions on the plants. In conclusion, the density of aphid infestation and time for which they were feeding influenced the production of induced compounds by tomatoes, the whitefly responses to the plants, and reduced interspecific competition.     

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.