Cornicle Secretion Research Articles

The Ecological Significance of Aphid Cornicles and Their Secretions.

Aphid cornicles are abdominal appendages that secrete an array of volatile and nonvolatile compounds with diverse ecological functions. The emission of alarm pheromones yields altruistic benefits for clone-mates in the aphid colony, which is essentially a superorganism with a collective fate. Secreted droplets also contain unsaturated triglycerides, fast-drying adhesives that can be lethal when smeared on natural enemies but more often impede their foraging efficiency. The longest cornicles have evolved in aphids that feed in exposed locations and are likely used to scent-mark colony intruders. Reduced cornicles are associated with reliance on alternative defenses, such as the secretion of protective waxes or myrmecophily. Root-feeding and gall-forming lifestyles provide protected feeding sites and are associated with an absence of cornicles. In some eusocial gall-formers, soldier morphs become repositories of cornicle secretion used to defend the gall, either as menopausal apterae that defend dispersing alatae or as sterile first instars that dispatch predators with their stylets and use cornicle secretions as a construction material for gall repair. Collectively, the evidence is consistent with an adaptive radiation of derived cornicle functions molded by the ecological lifestyle of the aphid lineage.

Potential of predatory Neotropical ladybirds and minute pirate bug on strawberry aphid.

Laboratory trials were performed to determine the impact of three Neotropical predatory coccinellids (Cycloneda sanguinea, Eriopis connexa and Coleomegilla quadrifasciata) and a minute pirate bug (Orius insidiosus) on Chaetosiphon fragaefolii, an important strawberry aphid pest. The predation on C. fragaefolii nymphs and adults, as well as the time to the first attack of all predators were compared with predation on Aphis gossypii. Predator preferences for prey and aphid defensive behavior were also evaluated. Moreover, the effect of coccinellids on C. fragaefolii population growth was assessed in experimental greenhouse conditions. The predation rate varied among predators, being significantly lower for O. insidiosus than for the coccinellids. Consumption was higher on A. gossypii than on C. fragaefolii, regardless of the aphids developmental stage. The time to the first attack of all predators was longer in the presence of C. fragaefolii. Walking away and cornicle secretion were the most common antipredator behaviors of aphid against coccinellids and O. insidiosus, respectively. Coccinellids preferred A. gossypii over C. fragaefolii, while O. insidiosus showed indifference. Cycloneda sanguinea and E. connexa exhibited the highest suppression effect on the growth rate of C. fragaefolii. Thus, the four predators evaluated could contribute to reduce strawberry aphid populations, especially C. sanguinea and E. connexa.

Cornicle secretions by Aphis fabae (Hemiptera: Aphididae) result in age-dependent costs and improved host suitability for Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae).

We examined the life history consequences of cornicle secretion by Aphis fabae Scopoli in second and fourth instars, and its effects on host suitability for its parasitoid, Lysiphlebus fabarum (Marshall). Cornicle secretion did not affect aphid fecundity, but secretion in the second instar enhanced life table parameters, whereas secretion in the fourth instar affected them negatively, suggesting a higher cost of secretion in later instars. Secretion in either instar improved host suitability for L. fabarum. Although control and treated aphids were parasitized at similar rates, and with similar success, wasps developed faster and emerged as larger adults in aphids that had secreted, regardless of instar. Transgenerational effects were also evident. Progeny emergence was higher when parental wasps developed in fourth instars than in seconds, whether aphids secreted or not, and progeny were larger when parental hosts secreted in the second instar, but not in the fourth. Secreting fourth instars were preferred to controls by L. fabarum females in choice tests, but not secreting second instars, and fourth-instar secretion improved wasp emergence. When control aphids were attacked, second instars were more likely to secrete than fourth instars, whereas the latter were more likely to kick the parasitoid. Cornicle secretion reduced the probability of subsequent secretion events and the frequency of other aphid defensive behaviors, indicating energetic tradeoffs among defensive tactics. Overall, our results revealed that cornicle secretion by immature A. fabae exacts both physiological and behavioral costs and results in improved host suitability for its parasitoid.

Mechanisms Underlying the Nonconsumptive Effects of Parasitoid Wasps on Aphids.

Natural enemies need not consume herbivores to suppress herbivore populations. Behavioral interactions can adversely impact herbivore fitness from reduced time feeding, investment in defense, or injury from failed attacks. The importance of such "nonconsumptive effects" for herbivore suppression may vary across species based on the specificity and intensity of the herbivore defensive response. In a series of manipulative studies, we quantified the nature and consequences of nonconsumptive interactions between two parasitoid wasps, Aphidius ervi Haliday and Aphidius colemani Viereck, on two aphid species, pea aphids (Acyrthosiphon pisum (Harris)) and green peach aphids (Myzus persicae (Sulzer)). Both wasps successfully parasitize green peach aphids, but only A. ervi parasitizes pea aphids. We observed A. ervi antennating and stinging pea aphids and documented a decrease in pea aphid longevity in response to stinging even when the aphid survived the interaction and no mummy formed. The primary defensive tactic of pea aphids in response to either wasp species was dropping from the host plant. Both wasp species antennated and stung green peach aphids, but they elicited unique defensive behaviors. Green peach aphids kicked or emitted cornicle secretions in response to A. colemani but spent more time off the plant in the presence of A. ervi. Green peach aphid longevity and fecundity were not affected by wasp stings when the aphid survived and no mummy formed. Our study demonstrates the complexity of behavioral interactions between parasitoids and their potential hosts and contributes to a mechanistic understanding of variation in the nonconsumptive suppression of herbivore populations.

Cornicle secretions of Uroleucon nigrotuberculatum (Homoptera: Aphididae) as the last bullet against lady beetle larvae

AbstractAphids have evolved various defense strategies against natural enemies, including secretions from their cornicles. We assessed the defensive function of cornicle secretions by the goldenrod aphid, Uroleucon nigrotuberculatum (Olive), against larvae of the lady beetles Coccinella septempunctata bruckii Mulsant and Propylea japonica (Thunberg) (Coleoptera: Coccinellidae). The aphid secreted red droplets from its cornicles when attacked by the larvae. Two‐thirds of the C. septempunctata bruckii larvae and 46.7% of the P. japonica larvae that preyed on the aphids died before reaching the pre‐pupal stage. The secretions caused molting failure when smeared on the larvae's heads or glued to the larvae's mouthparts, killing 56.7% of C. septempunctata bruckii larvae and 36.7% of P. japonica larvae. Second instar larvae were affected most. About 40% of third and fourth instar larvae of C. septempunctata bruckii vomited soon after ingesting the aphids. In the field, up to 40% of first and second instar larvae were smeared with red secretions. Our results show that these cornicle secretions are an effective and active defense against earlier instars of coccinellid larvae.

Soybean Aphid Response to their Alarm Pheromone E-ß-Farnesene (EBF)

When attacked by natural enemies some insect pests, including many aphid species, alert neighboring conspecifics with alarm pheromones. Cornicle secretions with pheromones benefit the attacked aphid but are costly to produce, while alarm pheromone benefits probably fall largely on alerted conspecifics. Given these variable benefits, the likelihood of a secretion may change depending on aphid density. Thus, we first hypothesized that the common alarm pheromone in aphids, E-s-farnesene (EBF), was present in soybean aphid (Aphis glycines Matsumura) cornicle secretions and would elicit an alarm response in aphids exposed to it. Second, since aphids other than the secretor also benefit from cornicle secretions, we hypothesized that the likelihood of secretion would increase concurrently with the density of neighboring clonal conspecifics. Third, because alarm reaction behavior (e.g. feeding cessation) is probably costly, we hypothesized that alarm reaction behavior would decrease as conspecific density (i.e. alternative prey for an attacking natural enemy) increased. We found that soybean aphids 1) produce cornicle secretions using EBF as an alarm pheromone, 2) are less likely to release cornicle secretions when alone than in a small group (~10 individuals), but that the rate of secretion does not increase further with additional conspecific density, and 3) also exhibit alarm reaction behavior in response to cornicle secretions independent of aphid density. We show that soybean aphids can use their cornicle secretions to warn their neighbors of probable attack by natural enemies, but that both secretion and alarm reaction behavior does not change as density of nearby conspecifics rises above a few individuals.

Chemical composition of cornicle secretion of the brown citrus aphid Toxoptera citricida

AbstractBrown citrus aphid Toxoptera citricida Kirkadly is considered as an important pest of citrus because it vectors citrus tristeza closterovirus. Aphids secrete a fluid from their cornicles as a defensive mechanism against natural enemies. Earlier studies on cornicle secretions of aphids focus only on triglycerides and fatty acids. In the present study, three different methods are used to investigate the chemical composition of the cornicle fluid of T. citricida. Gas chromatography with flame ionization detection is used to detect and quantify the triglycerides after trimethylsilyl derivatization, and gas chromatography‐mass spectrometry (GC‐MS) is used to determine the fatty acid composition after derivatization with boron trifluoride–methanol. Other compounds are detected using GC‐MS after methoxyamine hydrochloride and N‐methyl‐N‐(trimethylsilyl)trifluoroacetamide derivatization. The major fatty acid in the cornicle secretion of T. citricida is palmitic acid. Oleic, stearic, myristic, myristoleic and sorbic acids are also detected, although in low amounts. Sorboyl, dipalmitoyl (C6‐2, C16, C16) and disorboyl, stearoyl (C6‐2, C6‐2, C18) are the main triglycerides detected in cornicle secretion. Trehalose is the most predominant sugar (558.2 mm), followed by glucose (92.0 mm) and inositol (48.8 mm). Many amino acids, including proline, glycine, alanine and serine, are also detected. In addition, the cornicle secretion is rich in many organic acids, including malic, citric, succinic and lactic acid. Information obtained from the present study improves our understanding of the chemical composition of the cornicle secretion of the brown citrus aphid.

Is there any evidence that aphid alarm pheromones work as prey and host finding kairomones for natural enemies?

1. The aphid alarm pheromone (E)‐β‐farnesene (EBF) is often considered to be used by natural enemies as a prey/host finding kairomone. However, studies show opposing results, some appear to confirm an attraction of aphid natural enemies by EBF whereas others do not provide any evidence for the kairomone function of EBF.2. To clarify if aphid natural enemies are attracted by the amounts of EBF naturally emitted by aphids, the existing literature was reviewed about EBF attractiveness to aphid natural enemies with consideration of the amounts of EBF used in the studies.3. Thirty‐one publications that investigated the ability of EBF, aphid cornicle secretion, and attacked aphids, to attract aphid natural enemies were found. Several studies showed an attraction by EBF, but these used much higher amounts of EBF than usually emitted by aphids during a predator attack. Studies investigating EBF amounts similar to what is emitted by aphids are rare and failed to show attraction. Only two studies document an attraction of natural enemies by attacked aphids.4. As EBF is emitted in very low amounts, not very stable, and only present after an attack, we suggest that aphid‐derived EBF is not a suitable kairomone for most natural enemy species, especially when they are able to use alternative cues. As EBF, amongst other volatiles, is also emitted by herbivore‐induced plants, we propose that natural enemies might use plant‐derived EBF as a synomone to identify aphid‐infested plants via an altered plant volatile bouquet.

The effect of artificial diet on the production of alarm pheromone by Myzus persicae

AbstractThe cornicle secretion of Myzus persicae reared on artificial diet only elicits an alarm response in plant‐reared conspecifics after the young aphids have been transferred to plants for 7 days. Acetate in the form of 0.32% sodium acetate has been added to the diet as an early step in synthesis of the alarm pheromone, (E)‐β‐farnesene (EBF). The cornicle secretion of diet‐reared aphids then elicits an alarm response. However, there is no difference in internal EBF concentration between plant‐ and diet‐reared aphids. Puncturing aphids, either plant‐ or diet‐reared, with a pin shows that both can elicit an alarm response, whereas it is reduced by half with diet‐reared individuals. Although there is no significant difference in the concentration of EBF produced, the total amount in diet‐reared aphids is increased by acetate in the diet to a level similar to that in plant‐reared individuals: the size of aphids reared on an acetate‐supplemented diet is increased and comparable with the size of those that are plant‐reared. Bioassays with a range of EBF concentrations show a high threshold for the alarm response. It is concluded that the different size of aphids reared on plants and standard diet results in them secreting, respectively, above and below the response threshold.

Body size affects host defensive behavior and progeny fitness in a parasitoid wasp, Lysiphlebus fabarum

AbstractThis study tested effects of maternal body size on foraging behavior and progeny development in a thelytokous population of Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae). Small and large wasps were reared from first and second instar hosts [black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae)], respectively, and each was provided with a patch (bean leaf disk) containing either 15 small (second instar) or 15 large (fourth instar) hosts for a 30‐min foraging period. Neither body size nor host size affected time allocation to various behaviors within a patch, but second instar aphids produced significantly more mummies than fourth instars. The preferred attack orientation was from the side of the aphid, suggesting wasps were sensitive to the risk of smearing with cornicle secretions. Few wasps developed in fourth instar hosts, suggesting later host instars were somewhat resistant to parasitism. Second instar hosts, the most suitable stage for L. fabarum development, relied more on defensive behavior, specifically kicking and secreting cornicle droplets. Large wasps were more likely to elicit a double cornicle secretion, indicating that aphids graded their response to the size of the attacker. Larger wasps were also more likely to be smeared with cornicle secretion, suggesting they were more vulnerable than small wasps. Although small wasps had smaller eggs than large wasps, there was no effect of maternal egg size on the size of progeny. However, daughters of small females emerged with larger egg loads than daughters of large mothers, and their eggs tended to be slightly smaller, although not significantly. Regression analysis revealed a positive correlation between maternal egg size and progeny developmental time for small and large wasps, and between maternal egg size and progeny egg load for small wasps. These results confirm maternal effects of body size in an aphid parasitoid, and reveal that vulnerability to host behavioral defenses is also body size dependent.

Influence of cornicle droplet secretions of the cabbage aphid, Brevicoryne brassicae, on parasitism behavior of naïve and experienced Diaeretiella rapae

Insects have evolved amazing methods of defense to ward off enemies. Many aphids release cornicle secretions when attacked by predators and parasitoids. These secretions contain an alarm pheromone that alerts other colony members of danger, thereby providing indirect fitness benefits to the releaser. In addition, contact with cornicle secretions could also threaten an attacker and could provide direct fitness to the releaser. However, cornicle secretions may also be recruited as a kairomonal cue by aphid natural enemies. In this study, we investigated the effect of the cornicle droplet volatiles of the cabbage aphid, Brevicoryne brassicae (L.), on the host-searching behavior of naïve and experienced female Diaeretiella rapae (M(') Intosh) parasitoids in olfactometer studies. In addition, we evaluated the role of B. brassicae cornicle droplets on the oviposition preference of the parasitoid in a two-choice bioassay. Naïve females did not exhibit any preference between volatiles from aphids secreting cornicle droplets over non-secreting aphids, while experienced parasitoids exploited the secretions in their host location. Experienced females were also able to choose volatiles from both secreting and non-secreting aphids over clean air, while this ability was not observed in naïve females. Although secretion of cornicle droplets did not influence the percentage of first attack in either naïve or experienced females, the success of attack (i.e. resulting in a larva) was significantly different between secreting and non-secreting aphids in the case of experienced parasitoids.

Host selection and parasitism behavior of Lysiphlebus testaceipes: Role of plant, aphid species and instar

The aphid parasitoid Lysiphlebus testaceipes is a potentially valuable biological control agent of Aphis gossypii a major worldwide pest of cotton. One means of increasing the abundance of a biological control agent is to provide an alternative host habitat adjacent to cropping, from which they can provide pest control services in the crop. Host selection and parasitism rate of an alternative host aphid, Aphis craccivora by L. testaceipes were studied in a series of experiments that tested its host suitability relative to A. gossypii on cotton, hibiscus and mungbean. Host acceptance, as measured by number of ovipositions was much greater in A. craccivora compared to A. gossypii, while more host aphids were accepted on mungbean than cotton. When given a choice L. testaceipes attacks more 4th instar and adult stages (63% and 70%, respectively) of both hosts than 2nd instar nymphs (47%). In a switching (host choice) experiment, L. testaceipes preferentially attacked A. craccivora on mungbean over A. gossypii on cotton. Observations of parasitoid contact with A. gossypii cornicle secretion suggest it provides a useful deterrent against parasitoid attack. From these experiments it appears L. testaceipes has a preference for A. craccivora and mungbean compared to A. gossypii and cotton, in this respect using A. craccivora and mungbean as alternative habitat may not work as the parasitoid is unlikely to switch away from its preferred host.

Altruistic defence behaviours in aphids.

BackgroundAltruistic anti-predatory behaviours pose an evolutionary problem because they are costly to the actor and beneficial to the recipients. Altruistic behaviours can evolve through indirect fitness benefits when directed toward kin. The altruistic nature of anti-predatory behaviours is often difficult to establish because the actor can obtain direct fitness benefits, or the behaviour could result from selfish coercion by others, especially in eusocial animals. Non-eusocial parthenogenetically reproducing aphids form colonies of clone-mates, which are ideal to test the altruistic nature of anti-predatory defence behaviours. Many aphids release cornicle secretions when attacked by natural enemies such as parasitoids. These secretions contain an alarm pheromone that alerts neighbours (clone-mates) of danger, thereby providing indirect fitness benefits to the actor. However, contact with cornicle secretions also hampers an attacker and could provide direct fitness to the actor.ResultsWe tested the hypothesis that cornicle secretions are altruistic by assessing direct and indirect fitness consequences of smearing cornicle secretions onto an attacker, and by manipulating the number of clone-mates that could benefit from the behaviour. We observed parasitoids, Aphidius rhopalosiphi, foraging singly in patches of the cereal aphid Sitobion avenae of varied patch size (2, 6, and 12 aphids). Aphids that smeared parasitoids did not benefit from a reduced probability of parasitism, or increase the parasitoids' handling time. Smeared parasitoids, however, spent proportionately more time grooming and less time foraging, which resulted in a decreased host-encounter and oviposition rate within the host patch. In addition, individual smearing rate increased with the number of clone-mates in the colony.ConclusionsCornicle secretions of aphids were altruistic against parasitoids, as they provided no direct fitness benefits to secretion-releasing individuals, only indirect fitness benefits through neighbouring clone-mates. Moreover, the use of cornicle secretions was consistent with their altruistic nature, because the occurrence of this behaviour increased with the size of indirect fitness benefits, the number of clone-mates that can benefit. This study provides evidence for a case of kin-directed altruistic defence outside eusocial animals.

Ant mimicry by an aphid parasitoid, Lysiphlebus fabarum

In Iran, Lysiphlebus fabarum (Marshall) (Hymenoptera: Braconidae: Aphidiinae) is a uniparental parasitoid of the black bean aphid, Aphis fabae Scopoli (Hemiptera: Aphididae), that possesses various highly evolved adaptations for foraging within ant-tended aphid colonies. Direct observations and video recordings were used to analyze the behavior of individual females foraging for A. fabae on bean leaf disks in open arenas in the laboratory. Females exploited aphids as hosts and as a source of food, allocating within-patch time as follows: resting - 10.4%, grooming - 8.2%, searching - 11.5%, antennation (host recognition) - 7.5%, antennation (honeydew solicitation mimicking ants) - 31.9%, abdominal bending (attack preparation) 19.7%, probing with the ovipositor (attack) - 10.8%. The mean handling time for each aphid encountered was 2.0 ± 0.5 min. Females encountered an average of 47.4 ± 6.4 aphids per hour, but laid only 1.2 eggs per hour. The ovipositor insertion time for parasitism ranged from 2 sec to longer than a minute, but most insertions did not result in an egg being laid. A. fabae defensive behaviors included kicking, raising and swiveling the body, and attempts to smear the attacker with cornicle secretions, sometimes with lethal results. Food deprivation for 4–6 h prior to testing increased the frequency of ant mimcry by L. fabarum. Females also used ant-like antennation to reduce A. fabae defensive behavior, e.g. the frequency of kicking. L. fabarum attacks primed A. fabae to be more responsive to subsequent honeydew solicitation, such that experienced females improved their feeding success by alternating between the roles of parasitoid and ant mimic. These results reveal the possibility for mutualisms to evolve between L. fabarum and the ant species that tend A. fabae, since L. fabarum receive ant protection for their progeny and may benefit the ants by improving A. fabae responsiveness to honeydew solicitation.

Functional bases of host‐acceptance behaviour in the aphid parasitoid Aphidius ervi

Abstract The host acceptance behaviour in Aphidius ervi is investigated, assessing the role of both external and internal host‐associated cues, offered to the experimental parasitoids with parafilm‐made aphid dummies. The reaction to internal cues present in the host haemolymph is clearly evident, and its intensity is enhanced by external cues. Parasitoid females lay few eggs in aphid dummies filled with host haemolymph. A significant increase in the number of both oviposition reactions (host stinging) and egg laying is observed only when these dummies are coated with cornicle secretion. However, this enhancement is not observed when the aphid dummies contain distilled water. Thus, the host acceptance behaviour of A. ervi females appears to be controlled by the integration of both external and internal chemical cues. The physiological basis of this behavioural response is investigated with a detailed study on the anatomy and ultrastructure of A. ervi ovipositor. The detection of chemical cues present in the host haemolymph that act as kairomones is made possible by the presence of gustatory sensilla on the tip of the ovipositor. These sensilla consist of porous areas, reached by unbranched dendrites running inside both the lower valves (i.e. first valvulae) and the unpaired upper valve (i.e. second valvulae). The mechanosensory function during oviposition appears to be provided mostly by the basiconic sensilla found on the tip of external valves (i.e. third valvulae). A tentative functional model accounting for the observed oviposition behaviour of A. ervi is proposed.

Playing the hare or the tortoise in parasitoids: could different oviposition strategies have an influence in host partitioning in two Aphidius species?

In this paper, we compare the host selection behaviours of two parasitoids, Aphidius rhopalosiphi and Aphidius picipes, in order to analyse whether behavioural adaptations to the defensive behaviour of their host (the grain aphid Sitobion avenae) could, in part, be responsible for the simultaneous presence of both species in cereal fields. The oviposition behaviour of A. picipes differed from that of A. rhopalosiphi by including a number of ‘fluttering wings’ sequences followed by immobility. It resulted in a 44 times longer host-handling time for A. picipes than for A. rhopalosiphi. Hosts attacked by A. picipes exhibited fewer defensive behaviours than hosts attacked by A. rhopalosiphi. A. picipes and A. rhopalosiphi rejected respectively 0% and 53% of unparasitized hosts presenting cornicle secretions, one of the defensive means of aphids. Furthermore, A. picipes females rejected 100% of the hosts that were already parasitized, whereas A. rhopalosiphi was previously described to reject only 20 to 40% of such hosts. Such differences could be explained by the way the two species deal with the aphid defensive behaviour. Field analyses, showed that A. rhopalosiphi was already present in wheat fields in early April whereas A. picipes appeared later and only achieved a low level of parasitism. However, when both species were present simultaneously, they shared the same resource.

Acceptance and suitability of Acyrthosiphon pisum and Sitobion avenae as hosts of the aphid parasitoid Aphidius ervi (Hymenoptera: Braconidae)

Aphidius ervi Haliday is a parasitoid of natural populations of both Acyrthosiphon pisum (Harris) on alfalfa and Sitobion avenae (Fabricius) on wheat in Chile. In this study the performance of the aphid parasitoid on both host species was evaluated. Regardless of origin, both females and males of A. ervi were significantly smaller when reared on S. avenae than on A. pisum. Males and females of A. ervi reared on S. avenae on wheat also took significantly longer to develop than when reared on A. pisum. There was no significant difference in the sex ratio of the parasitoid when reared on these two host aphids. Survival of parasitoids from the A. pisum - alfalfa system was significantly lower when reared on S. avenae relative to those reared on A. pisum, but no significant difference in survival on both hosts was detected in parasitoids from the S. avenae - wheat system. Behaviour of parasitoid females during oviposition, such as frequency of encounters, number of attacks and stabbings of host aphids, were significantly higher in the A. pisum - A. ervi interaction than in the S. avenae - A. ervi interaction, regardless of the origin of the parasitoid. Aphid defensive reactions such as kicking or production of cornicle secretion, were significantly higher in the A. pisum - A. ervi than the S. avenae - A. ervi interaction. The results are discussed in relation to the effect of host quality and host aphid body size on parasitoid development.

Superparasitism limitation in an aphid parasitoid: cornicle secretion avoidance and host discrimination ability

Superparasitism avoidance by the endoparasitoid Aphidius rhopalosiphi De Stefani Perez on the grain aphid, Sitobion avenae was studied. Experiments were carried out in which aphids were exposed to two consecutive attacks by parasitoids. Results showed that superparasitism avoidance in A. rhopalosiphi was mediated by two successive stimuli whose effectiveness depended on the time interval between attacks. For short time intervals (<16 h), host rejections were mainly associated with the presence of dried cornicle secretion on the host's body which was exuded during the first attack. The repellency of this secretion declined with the time interval between attacks, becoming ineffective 2 days after the first parasitization, and allowed females to reject up to 30% of parasitized hosts. For longer time intervals (≥16 h), host rejection behavior was a response of parasitoid females to internal changes in host quality associated with parasite development. This response gradually increased with an increase in time interval, reaching no more than 60%, 96 h after initial parasitization. This host discrimination ability did not allow females to distinguish between hosts parasitized by themselves or by conspecifics. Consequently, these findings suggest that superparasitism is a common event in A. rhopalosiphi and especially on recently parasitized hosts.

Physical and chemical cues influencing the oviposition behaviour of Aphidius ervi

AbstractThe oviposition behaviour of the aphid parasitoid Aphidius ervi Haliday is influenced by both chemical and physical cues. Oviposition attack responses were elicited by paint pigments sealed into the tip of a glass capillary tube. Parasitoids reacted to yellow pigments with repeated oviposition attack responses, but they did not react to green pigments. The spectrum of reflected light from the yellow pigments was very similar to that from the ‘green’ natural host Acyrthosiphon pisum (Harris), with a high proportion of the total radiation energy being emitted in the yellow‐orange wavebands (580–660 nm). Pea aphid cornicle secretion also elicited oviposition attack responses, which were not exclusively induced by its pale yellow‐green colour. In fact, the oviposition attack response to capillary tips coated with cornicle secretion remained evident under red light conditions, which, in contrast, nearly completely suppressed the response to yellow pigments. Chemical compounds from cornicle secretion do not appear to be involved in parasitoid orientation, even though they stimulate intense oviposition attack responses. Olfactometer experiments showed that the putative kairomone involved acts only at very short range or on contact. Host exuviae, which also elicited strong and persistent oviposition reactions from A. ervi females, appear to be a good alternative source of ovipositional kairomone(s). This work confirms the existence of an aphid cuticular kairomone.

Epicuticular wax ester and triacylglycerol composition in relation to aphid infestation and resistance in red raspberry ( Rubus idaeus L.)

Epicuticular waxes from two cultivars of red raspberry ( Rubus idaeus) were collected from the newly emerging crown leaves, and also from the group of four more mature leaves immediately below the crown. One cultivar, Autumn Bliss, was identified as aphid-resistant, and the other, Malling Jewel, as aphid-susceptible following bioassay with the large raspberry aphid, Amphorophora idaei, just prior to collection of the wax. Biological activity was primarily associated with the more mature leaves. Epicuticular wax esters consisted predominantly of long-chain aliphatic compounds in which even-carbon-number acids were esterified to even-carbon-number alcohols. Lesser amounts of odd-carbon-number esters were also present. The acid : alcohol combinations of the major esters were C 38: 14 : 24, 16 : 22, 20 : 18; C 40: 14 : 26, 16 : 24, 18 : 22, 20 : 20; C 42: 16 : 26, 20 : 22; 22 : 20, C 44: 20 : 24, 22 : 22, 24 : 20; C 46: 20 : 26, 22 : 24, 24 : 22; C 48: 20 : 28, 22 : 26, 24 : 24, 26 : 22; C 50: 20 : 30, 22 : 28, 24 : 26, 26 : 24, 28 : 22 and C 52: 22 : 30, 24 : 28, 26 : 26. Terpenyl esters were also present and these consisted of α- and β-amyrin and cycloartenol esterified to C 16, C 18 and C 20 acids. Compositional differences between the more mature leaves which may relate to resistance to A. idaei were higher levels of cycloartenyl esters and α-amyryl esters in wax from the resistant cultivar Bliss. There were also differences between the cultivars in the distribution of individual alkyl esters and their component acids and alcohols. Esters with longer acid : shorter alcohol combinations were more abundant in Jewel than Bliss. There were compositional differences between leaves at the different developmental stages. Alkyl esters were more abundant and cycloartenyl esters were not detected in wax from the immature leaves. Small amounts of an unusual class of triacylglycerol were found only on leaves of the aphid-susceptible cultivar, Jewel, which had been subject to bioassay with raspberry aphid. These compounds, which have a C 6 acid at C-2 of the glycerol backbone, were derived from the aphid, and are the major component in the insect’s cornicle secretions. The triacylglycerols probably arise from the presence on the leaf surface of shed aphid skins, or by incorporation of cornicle fluid into the leaf wax. The abundance of aphid triacylglycerols on the leaf surface may provide a measure of aphid-susceptibility.