Pea aphids (Acyrthosiphon pisum Harris) reduce secretion of extrafloral nectar in broad bean (Vicia faba)

1. Herbivores sometimes suppress plant defences. This study tested whether the presence of pea aphids (Acyrthosiphon pisum Harris) on broad bean (Vicia faba) led to decreased secretion of extrafloral nectar (EFN) which functions as an indirect plant defence against herbivores. 2. To determine effects of aphid infestation on EFN secretion, a comparison was done between EFN secretion in uninfested plants and that in plants infested by A. pisum and another aphid species (Aphis craccivora Koch). 3. When broad bean plants were infested by A. pisum, they secreted significantly smaller amounts of EFN than did uninfested plants and A. craccivora‐infested plants. There was no significant difference in EFN secretion between uninfested plants and A. craccivora‐infested plants. The number of extrafloral nectaries did not differ among the three treatments. 4. These results suggest that A. pisum reduced EFN production in broad bean plants.     

Growth and wilting of radish seedlings, Raphanus sativus, infested with the aphid, Myzus persicae

After 10 days of infestation by the aphid Myzus persicae growth of radish seedlings was reduced by 38 %, and the volume of sap bleeding from the stumps of cut stems of infested seedlings was 4% of that exuding from the stumps of uninfested seedlings of the same age. Diminished bleeding could not be accounted for by the 52 % reduction in the weight of the roots of infested seedlings. After 14 days of infestation, necrotic patches developed along the veins of the cotyledons which yellowed noticeably. If freed of aphids the seedlings subsequently resumed growth and their was an increase in the volume of bleeding sap produced by their cut stems. Symptoms associated with aphid infestation cannot be attributed either to virus disease or initially to exhaustion of the seedling's carbohydrate and nitrogen reserves. Possible causes of the symptoms described are discussed.     

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.     

Some factors influencing the occurrence and site of oviposition by aphidophagous Syrphidae (Diptera)

Syrphids were not deterred from ovipositing on suitable plants by the presence of other syrphid eggs, larvae or larval gut contents. Heavy infestations of Aphis fabae Scop, on bean plants (Viciafaba L.) induced Platycheirus manicatus (Meig.) to lay preferentially on neighbouring uninfested plants. Such behaviour was not found for Syrphus species. Nor did flowers, a source of adult food, stimulate oviposition nearby. The site of oviposition in relation to Brevicoryne brassicae L. on brussels sprouts (Brassicae oleracea gemnifera L.) varied markedly according to species: in similar conditions Syrphus luniger Meig. laid over 50% of its eggs touching aphids and less than 1% on uninfested plants, whereas P. manicatus laid less than 5% touching aphids and over 50% on uninfested plants. Most eggs were laid close to the edge of the leaf, especially by Platycheirus species, e.g. P. scutatus Meig. which laid over 90% within 5 mm of the edge of the leaf.     

Changes in growth regulator content of radish seedlings, Raphanus sativus, infested with the aphid Myzus persicae

Following 10 days of infestation by the aphid Myzus persicae there was an increase in the amount of growth inhibiting substances and a decrease in cytokinins, gibberellins and auxins in infested as compared with similar uninfested radish seedlings. Even after previously infested seedlings have been freed of aphids for 10 days, differences in the hormone balance remained. The possible relationships between the changes in hormonal balance and the effect of the aphid infestation on growth, translocation and wilting are discussed.     

Interactions between the stem-mining weevils Ceutorhynchus napi Gyll. and Ceutorhynchus pallidactylus (Marsh.) (Coleoptera: Curculionidae) in oilseed rape

Abstract 1 The rape stem weevil Ceutorhynchus napi Gyll. and the cabbage stem weevil Ceutorhynchus pallidactylus (Marsh.) share the same habitat and food resource within the stems of oilseed rape, Brassica napus L. var. oleifera. Interactions occurring between these two sympatric species on this host were studied under both field and laboratory conditions. 2 The oviposition preference of C. pallidactylus and the within‐plant distribution of eggs and larvae were examined in field plots of oilseed rape. Female C. pallidactylus tended to lay their eggs in plants already infested by eggs and larvae of C. napi rather than in uninfested plants. The within‐plant distribution of the egg batches of C. pallidactylus did not differ significantly between uninfested plants and those preinfested by C. napi. Ovipositing females of C. napi and C. pallidactylus generally showed a significant preference for plants with larger stem diameter. 3 Laboratory choice tests provided further evidence for the oviposition preference of C. pallidactylus. Females laid significantly more eggs in leaves of plants that had been previously infested by C. napi than in leaves of previously uninfested plants. 4 Larvae of C. pallidactylus showed a significant shift of their feeding niche towards the stem base when feeding in individual plants attacked by both species. This possibly indicates ressource partitioning between C. pallidactylus and C. napi. The within‐plant distribution of C. napi larvae was not affected by the simultaneous attack of C. pallidactylus. 5 The size of the head capsule of full‐grown larvae of C. napi and C. pallidactylus was not significantly correlated with the diameter of the stem of their host plant or with the number of conspecific larvae within individual plants.     

Host-marking by female pepper weevils, Anthonomus eugenii

Pepper weevils, Anthonomus eugenii Cano (Coleoptera: Curculionidae), feed and oviposit in flower buds and small fruits of plants in the genus Capsicum, as well as several species of Solanum (Solanaceae). Females chew a small hole into the fruit, deposit a single egg within the cavity, and seal the hole with a clear anal secretion that hardens into an ‘oviposition plug’. Female oviposition behavior was studied in a series of small‐arena bioassays to determine whether previous oviposition in Jalapeño pepper fruit deterred subsequent oviposition and to determine what specific cues from an infested fruit influence female behavior. In choice and no‐choice tests, females preferred clean fruit to fruit that had received four eggs 24 h previously (i.e., infested fruit), whether the fruit was infested with conspecific eggs or their own eggs. Further bioassays demonstrated that the presence of female frass, or oviposition plugs alone, in the absence of eggs or any fruit damage, was sufficient to deter oviposition. In addition, females given the choice between an infested fruit with the oviposition plug removed or an unaltered infested fruit preferred the fruit with no plugs, even when eggs, frass, and feeding damage were still present. To determine whether females would avoid infested peppers under more natural conditions, we quantified oviposition on infested and uninfested sentinel pepper fruit within individually caged plants and on clean and infested plants caged together. Females consistently laid more eggs on clean fruit than on infested fruits and moved within and among pepper plants to search for more acceptable oviposition sites. We conclude that oviposition plugs, along with contaminated female, but not male, frass contain a deterrent that, in the absence of any other cue, is enough to alert a female that a patch is occupied.     

The effects of age and weather on egg laying inDanaus plexippus L. (Lepidoptera: Danaidae)

The effects of age and weather conditions on egg laying in D. plexippus were determined for caged females. Age (measured in physiological time), temperature and solar radiation influence egg laying in this species of butterfly. An algorithm taking these factors into account in presented and accounts for 88% of the daily variation in egg laying. Caged D. plexippus begin to lay eggs six—seven days after emergence, peak egg production (about 60 eggs/♀) occurs about 15 days later. Females continue to lay eggs throughout their adult life, which in a flight cage was about 40 days. This egg laying pattern is compared with other published fecundity schedules. The effect and importance of a female being prevented from laying her eggs, on her life-time egg production, is also discussed.     

Changes in photosynthetic and transpiration rates of cotton leaves infested with the cotton aphid, Aphis gossypii: Unrestricted infestation

The influence of aphid, Aphis gossypii, feeding on photosynthesis and transpiration in cotton plants was investigated under greenhouse conditions. Four population densities of 0, 5, 10, and 25 aphids were used to infest individual cotton leaves. Gas exchange rates were determined for single attached cotton leaves after 9, 18, and 27 days of aphid infestation. Aphid feeding changed photosynthetic rates and transpiration rates. These changes were proportional to the number of aphids and the length of infestation period. Photosynthetic rates were significantly reduced in infested leaves with 25 aphids over 18 days, whereas significant reduction in photosynthetic rates was recorded within 27 days in infested leaves with 5, 10, and 25 aphids in comparison to their respective control. Initial population of 10 aphids increased significantly the transpiration rate of infested leaves over 9 and 27 days. Leaves of plants with 25 aphids had significantly greater transpiration rate than the control at all times.     

Oviposition site selection of Episyrphus balteatus

The oviposition behaviour of Episyrphus balteatus DeGeer (Diptera: Syrphidae) was investigated in four-choice experiments in screen cages and in experiments with direct behavioural observations. Females laid hardly any eggs in the absence of aphids and preferred larger aphid colonies to small ones. Honeydew attracted the females via olfaction and elicited oviposition. Females were able to perceive the presence of conspecific eggs and oviposited less often in aphid colonies in which eggs were present. The oviposition deterring stimulus was also active when the eggs were removed, hence, the deterrent did not act visually, but probably olfactorily.     

The oviposition behavior ofAphidius ervi andMonoctonus paulensis (Hymenoptera: Aphididae) encountering different host species (Homoptera: Aphididae) in sequential patches

Two-day-old mated females ofAphidius ervi Haliday andMonoctonus paulensis (Ashmead) were each provided with two sequential host patches. Patches were comprised of plastic petri dishes containing either 15 pea aphids,Acyrthosiphum pisum (Harris), or 15 alfalfa aphids,Macrosiphum creelii Davis. Both wasp species parasitized more hosts in patches containing pea aphids than in those containing alfalfa aphids, regardless of sequence. Females ofA. ervi also laid more eggs per aphid in patches containing pea aphids than in patches containing alfalfa aphids. When both patches contained alfalfa aphids,M. paulensis females parsitized more aphids in the second patch than in the first. Fewer alfalfa aphids were parasitized in the second patch when the first patch contained pea aphids, and fewer eggs were laid per alfalfa aphid. Parasitoid females of both species exhibited consistently higher rates of oviposition into their preferred host species and adjusted their reproductive allocation to hosts and host patches as a function of their experience in previous patches.     

Females of Cotesia vestalis,a parasitoid of diamondback moth larvae,learn to recognise cues from aphid‐infested plants to exploit honeydew

1. To maximise their reproductive success, the females of most parasitoids must not only forage for hosts but must also find suitable food sources. These may be nectar and pollen from plants, heamolymph from hosts and/or honeydew from homopterous insects such as aphids. 2. Under laboratory conditions, females of Cotesia vestalis, a larval parasitoid of the diamondback moth (Plutella xylostella) which does not feed on host blood, survived significantly longer when held with cruciferous plants infested with non‐host green peach aphids (Myzus persicae) than when held with only uninfested plants. 3. Naïve parasitoids exhibited no preference between aphid‐infested and uninfested plants in a dual‐choice test, but those that had been previously fed aphid honeydew significantly preferred aphid‐infested plants to uninfested ones. 4. These results suggest that parasitoids that do not use aphids as hosts have the potential ability to learn cues from aphid‐infested plants when foraging for food. This flexible foraging behaviour could allow them to increase their lifetime reproductive success.     

On oviposition in weevils of the genus <Emphasis Type="Italic">Larinus</Emphasis> Dej. (Coleoptera,Curculionidae)

The following actions performed by females of several Larinus Dej. species during egg laying are described: search of an appropriate place on the plant, making the hole for the egg, oviposition proper, and sealing the hole. The hole preparation takes the longest time and the greatest effort. Only one individual usually completes development in one flower head. Females of Larinus vulpes Ol. prefer larger flower heads for oviposition and occasionally lay eggs into stems. The ability of females to distinguish the flower heads with already laid eggs is discussed. Species of Larinus may be divided into two groups with “early” and “late” oviposition. The evolution of egg laying in the genus Larinus is discussed.     

The Foraging Behavior of <Emphasis Type="Italic">Lysiphlebus fabarum</Emphasis> (Marshall), a Thelytokous Parasitoid of the Black Bean Aphid in Iran

A series of laboratory experiments examined the foraging behavior of a thelytokous strain of Lysiphlebus fabarum (Marshall), a strongly proovigenic parasitoid of Aphis fabae Scopoli, in Iran. Females use chemical camoflage to forage undisturbed in ant-tended aphid colonies and solicit honeydew from aphids in the manner of ants. Rates of oviposition are very low (∼ 1.2 eggs / h) despite many aphid encounters and persistent ovipositor probing which appears to prime aphids for subsequent honeydew solicitation. Starved females spent 3.6 times longer in host patches (leaf disks with 15 2nd–3rd instar A. fabae) than did females sated on honey, and 40% of this time was spent soliciting honeydew. Five d-old females spent longer in patches than did one d-old females, and parasitized three times as many aphids. A 24 h pre-trial foraging experience did not reduce mean egg load significantly compared to a one h experience, but was sufficient to reduce patch residence time and number of aphids parasitized. Wasps reared under short day conditions (L:D = 10:14) were more pessimistic foragers (remained longer in patches and parasitized more aphids) than females raised under long days (L:D = 16:8). Wasps that encountered aphids previously parasitized by conspecifics began to superparasitize and remained longer in patches than females that encountered only unparasitized aphids. Encounters with other females had no effect on foraging behavior, possibly because cuticular camoflage interferes with conspecific recognition. The exceptionally low oviposition rate of this wasp may reflect a life history in which individual fitness has evolved to be strongly dependent on continued ant attendance throughout the period of progeny development.     

Some host-plant factors affecting oviposition by aphidophagous Syrphidae (Diptera)

Although most aphidophagous syrphid species lay their eggs close to aphids, Platycheirus peltatus (Meig.), Melanostoma scalare (F.) and M. mellinum (L.) laid freely on uninfested brussels sprout plants. The existence of species that will lay in the absence of aphids may be valuable in biological control and also provides a useful tool for the investigation of non-aphid oviposition stimuli. Plant species and plant appearance were both important factors in P. peltatus oviposition. Particular sites on the plant were chosen for oviposition by Syrphus balteatus (Deg.). The nature of the substrate at the oviposition site affected the size of the egg batches laid there by M. scalare, M. mellinum, P. peltatus and P. clypeatus (Meig.). The possibility is discussed that plant-location is the older-established system of host-finding and that an aphid-location system has gradually superseded it during the evolution of the obligatorily aphidophagous Syrphinae.     

The role of larval food resources and adult movement in the population dynamics of the orange-tip butterfly (Anthocharis cardamines)

Numbers of the orange-tip butterfly were recorded on a permanent transect in Monks Wood National Nature Reserve between 1982 and 1993, together with numbers of its only larval food plant in the wood, Cardamine pratensis. Females of the butterfly lay their eggs on the newly opened flower heads of Cardamine and the larvae feed on the developing seed pods. They are extremely selective in their choice of plant for egg laying, choosing mainly large flower heads growing in open sunny locations. Larval survival is greatly reduced if the flower head is more than 8 days old at the time of egg laying, because the seed pods become too tough for feeding. Earlier studies showed that only one larva can survive on a flower head, because larvae are cannibalistic, but females tend to avoid plants carrying more than one egg, in response to an oviposition-deterring pheromone laid down at the time of egg laying. The numbers of flower heads of Cardamine fluctuate enormously between years, and the numbers of eggs follow these closely. There is a strong correlation between number of eggs laid and the availability of suitable flower heads, and this correlation was shown to be real and causal by the experimental provision of extra flower heads, which resulted in more eggs, laid over an extended period of oviposition. The main causes of mortality of the butterfly's young stages were egg infertility, cannibalism, predation, starvation, the grazing of flower heads by deer, and possibly the failure of the newly hatched larvae to feed adequately. Adult numbers were only weakly correlated with the numbers of young stages in the wood, and there appears to be a considerable amount of movement by the butterfly through the wood and surrounding farmland. Received: 8 October 1996 / Accepted: 3 April 1997     

Aphid-host plant interactions: does aphid honeydew exactly reflect the host plant amino acid composition?

Plants provide aphids with unbalanced and low concentrations of amino acids. Likely, intracellular symbionts improve the aphid nutrition by participating to the synthesis of essential amino acids. To compare the aphid amino acid uptakes from the host plant and the aphids amino acid excretion into the honeydew, host plant exudates (phloem + xylem) from infested and uninfested Vicia faba L. plants were compared to the honeydew produced by two aphid species (Acyrthosiphon pisum Harris and Megoura viciae Buckton) feeding on V. faba. Our results show that an aphid infestation modifies the amino acid composition of the infested broad bean plant since the global concentration of amino acids significantly increased in the host plant in response to aphid infestations. Specifically, the concentrations of the two amino acids glutamine and asparagine were strongly enhanced. The amino acid profiles from honeydews were similar for the two aphid species, but the concentrations found in the honeydews were generally lower than those measured in the exudates of infested plants (aphids uptakes). This work also highlights that aphids take large amounts of amino acids from the host plant, especially glutamine and asparagine, which are converted into glutamic and aspartic acids but also into other essential amino acids. The amino acid profiles differed between the host plant exudates and the aphid excretion product. Finally, this study highlights that the pea aphid, a “specialist” for the V. faba host plant, induced more important modifications into the host plant amino acid composition than the “generalist” aphid M. viciae.     

Hoverfly oviposition response to aphids in broccoli fields

The oviposition response of predacious hoverflies (Diptera: Syrphidae) to Brevicoryne brassicae L. and Myzus persicae (Sulzer) (Homoptera: Aphididae) in commercial broccoli, Brassica oleracea var. italica L., Plenck (Brassicaceae), fields was investigated at two sites over the course of a growing season. The hoverfly oviposition responses to these aphid species on different parts of the broccoli plant canopy were also examined. There were no hoverfly eggs on broccoli plants without aphids, egg numbers were very low on plants with fewer than 50 aphids, and no peak in oviposition relative to aphid numbers was observed. Within individual plants that were colonized by aphids, there was some oviposition on individual leaves without aphids, and no hoverfly eggs were seen on leaves that had more than 400 aphids. Leaves in the broccoli plant canopy, and the datasets associated with them, were divided into three sections vertically, ‘upper’, ‘middle’, and ‘lower’. Brevicoryne brassicae was more abundant in the upper and middle canopy sections, while M. persicae was found mostly in the lower section. The rate of hoverfly oviposition per aphid was higher in the upper section than in the two other sections. Modeling of the oviposition response using logistic regression showed that the presence of hoverfly eggs was positively correlated with numbers of each aphid species and sampling date.     

Aphid infestation promotes survival of a seed predator: observations and experiments on a tritrophic community module

We investigated whether aphid presence and abundance influence the survival of an endophagous pre-dispersal seed predator of the same host plant. We studied a terrestrial community module consisting of one plant (Laburnum anagyroides) and four insect species/groups (an aphid, Aphis cytisorum, a pre-dispersal seed predator bruchid, Bruchidius villosus, aphid-attending ant species, and parasitoids of the bruchid). Two complementary investigations were carried out in parallel: (a) a plant-aphid-ant complex was experimentally manipulated by excluding aphids, ants, or both for 5 years to assess their impacts on the seed predator’s survival and parasitism rate; and (b) different aphid infestation levels on randomly selected infructescences were correlated with plant traits, nutrient allocation pattern, and variables of seed predator’s survival, such as the number of eggs laid and adults emerged influenced by parasitoid activity, for 7 years. We found that ants did not affect bruchid oviposition negatively, but egg-parasitism was significantly decreased by their presence. Plant traits, such as the number of seeds and seed mass, as well as seed predator performance were negatively affected by heavy aphid infestation. Seed predator -infested seeds had no effect on the mass of remaining seeds in the pods. This study suggests that aphids were nevertheless promoting bruchid abundance and survival, depending on their infestation rate.     

Role of terpenes from aphid-infested potato on searching and oviposition behavior of Episyrphus balteatus

To cope with pathogen and insect attacks, plants develop different mechanisms of defence, in both direct （physical and chemical） and indirect ways （attractive volatiles to entomophagous beneficials）. Plants are then able to express traits that facilitate ＂top-down＂ control of pests by attracting herbivore predators. Here we investigate the indirect defence mechanism of potato plants by analyzing the volatile patterns of both healthy and aphid- infested plants. Important changes in the emitted terpene pattern by the Myzus persicae infested host plant were observed. Using Solid Phase MicroExtraction （SPME） and GC-MS, the （E）-fl-farnesene （EBF） appeared to be emitted by aphid-infested potato and not by healthy plants. To assess the infochemical role of these volatile releases after aphid damage on the aphidophagous predators Episyrphus balteams, the hoverfly foraging behavior was assessed using the Observer 5.0 software （Noldus, Wageningen, The Netherlands）. Aphidfree potato plants were also used as a control volatile source in the predator behavioral study. While aphid-infested plants induced efficient searching and acceptation behaviors leading to egg-laying, no kairomonal effect of healthy potato plants was observed, leading to longer immobility durations and shorter searching periods in the net cage. High oviposition rate of E. balteatus was observed when aphid-infested potato was used （mean of 48.9 eggs per laying and per female）. On the other hand, no egg was produced by the hoverfly on healthy aphid-free plants. The E. balteatus foraging and reproductive behaviors according to the volatile emission from aphid-infested plants are discussed in relation to the potential use of active infochemical molecules in integrated aphid pest management.