Isolation of Flower-inducing and Flower-inhibitory Factors from Aphid Honeydew

The aphid Dactynotus ambrosiae Thomas has been allowed to feed on vegetative or flowering plants of the short-day plant Xanthium strumarium L., and the honeydew which they produce is extracted and tested for an effect on flowering using the long-day plant Lemna gibba L., strain G3 for the bioassay. One zone of flower-inducing activity and at least two zones of flower-inhibitory activity are consistently obtained from the honeydew extracts. The levels of flower-inducing and flower-inhibitory activity are not demonstrably different in vegetative and flowering honeydew. The honeydew extracts are inactive on Xanthium but do give some flower induction with the short-day plant Lemna perpusilla Torr., strain 6746. The flower-inducing activity is clearly of plant origin and is present in the phloem since the same active material can be obtained from vegetative or flowering Xanthium by methanol extraction, and honeydew produced by aphids feeding on a chemically defined synthetic diet is completely without flower-inducing activity. This is the first report of successful flower induction in the long-day plant L. gibba G3 by some means other than long-day treatment.     

Host plant recognition in monophagous weevils: Specialization ofCeutorhynchus inaffectatus to glucosinolates from its host plantHesperis matronalis

Several aphid honeydews were incorporated into sucrose solutions and presented to hop aphids, Phorodon humuli (Schrank), as artificial diets in free-choice bioassays. Small additions of honeydew collected from two species of aphid feeding on hop, Humulus lupulus L., arrested the searching behavior of the hop aphid and appeared to stimulate prolonged periods of ingestion. This effect was more dependent on the host plant honeydew source than the species of aphid that produced the honeydew. Aphid honeydews collected from plants other than hop (non-hosts to P. humuli) contained hop aphid phagostimulants that were less effective. Our results indicate that analysis of aphid honeydew could help describe chemical cues involved in the recognition of appropriate host plants by aphid species.     

Plant–rhizobia interactions alter aphid honeydew composition

Both above- and below-ground interspecific interactions contribute to ecosystem functioning in terrestrial systems, and the integration of below- and above-ground interactions is crucial for deepening our knowledge of nutrient cycling and community dynamics in terrestrial ecosystems. The present study explored the effects of plant–microbe interactions on aphid honeydew quality and quantity and important factors mediating ant–aphid mutualisms and below-ground nutrient dynamics. Soybean aphids (Aphis glycines) were inoculated onto two closely related strains of soybean plants: a nodulating strain that associates with rhizobia and a non-nodulating strain that does not harbor any nitrogen-fixing bacteria. As expected, prior to aphid inoculation, nodulating plants were significantly taller and had more leaves than non-nodulating plants. Aphids feeding on nodulating strains were found to reach slightly larger colony sizes and produce honeydew with significantly different sugar profiles than those feeding on non-nodulating plants. The honeydew collected from aphid colonies feeding on nodulating plants contained 160 % more total sugars than honeydew collected from colonies feeding on non-nodulating plants, but there was no difference in total amino acid-N content in honeydew from colonies feeding on the different plant strains. We discuss the implications of honeydew composition for nutrient cycling and community dynamics and suggest areas of future research to elucidate the consequences of altered aphid honeydew composition on ecosystem properties.     

The response of an aphid tending ant to artificial extra-floral nectaries on different host plants

Sap-feeding homopterans, which reduce the fitness of their host plants, are often tended by ants that feed on their honeydew. The composition of the honeydew varies with both the aphid and the host plant. Extra-floral nectaries (EFNs) are believed to have evolved to attract attending ants, protecting the hosts, but it is unknown if EFNs on different plants have the same impact on the relations between an aphid species feeding on those plants and its tending ant. Experimental research was conducted to examine the attraction of Tapinoma erraticum scout ants to honeydew from the aphid Aphis gossypii feeding on two different plants, Prunus amygdalus and Mentha piperita, negligence of tending the aphids, and survival of the aphids in the presence of artificial EFNs. The scout ants were significantly more attracted to artificial nectar dispensed on P. amygdalus leaves than on M. piperita, or aphids on both plants and water. They neglected aphids in the presence of artificial EFNs on M. piperita but not on P. amygdalus. The aphid population on M. piperita did not statistically change in the presence of artificial EFNs during the 8 days of the third experiment. On P. amygdalus, the aphids succeeded in developing fully to winged form. In conclusion, the responses of the ants tending aphids to the presence of artificial EFNs were influenced by the host plant.     

The importance of honeydew as food for larvae of Chrysoperla carnea in the presence of aphids

Larvae of the common green lacewing Chrysoperla carnea are predacious and feed on a wide range of small, soft‐bodied arthropods. In addition to their feeding on prey arthropods to cover their nutritional requirements for growth and development, the consumption of non‐prey foods such as honeydew has been reported. It is commonly believed that these food supplements are primarily exploited by the larvae when prey is scarce or of low nutritional quality. Here, we assess whether C. carnea larvae also use honeydew when high‐quality aphid prey are readily available. In a choice experiment, the feeding behaviour of C. carnea larvae was observed in the presence of both aphids and honeydew. The larvae were starved, aphid‐fed, or honeydew‐fed prior to the experiment. The time spent feeding on honeydew compared with feeding on aphids was highest for starved larvae and lowest for honeydew‐fed larvae. Among the three treatments, the aphid‐fed larvae spent the most time resting and the least time searching. In an additional experiment food intake was assessed in terms of weight change when larvae were provided with an ad libitum supply of either aphids or honeydew. Larvae yielded a significant lower relative weight increase on honeydew compared with aphids. The reduced weight increase on honeydew was compensated when larvae were subsequently provided with aphids, but not when honeydew was provided again. This study showed that (i) prior honeydew feeding reduces overall aphid consumption, and (ii) larvae do consume honeydew even after they have been given ad libitum access to aphids. The fact that larvae of C. carnea still use honeydew as a food source in the presence of suitable prey underlines the importance of carbohydrates as foods.     

Feeding location affects demographic performance of cabbage aphids on winter canola

The cabbage aphid, Brevicoryne brassicae L. (Hemiptera: Aphididae), is a perennial pest that specializes on plants of the Brassicaceae family, attacking winter canola (Brassica napus L.) mainly during and after flowering. Under field conditions, cabbage aphid colonizes the upper flowering canopy. Population dynamics of aphids in the flowering canopy could be regulated by differences in either plant quality (bottom‐up) or predatory (top‐down) forces. The goal of our study was to determine the effect of feeding location on cabbage aphid demography. A stage‐structured matrix population model was constructed for aphids restricted to reproductive or vegetative plant tissues of canola. We found that feeding location had a large impact on demography of cabbage aphid; the finite rate of increase (λ ± SEM) was higher when aphids were restricted to reproductive tissues, compared to aphids feeding on vegetative tissues: 1.25 ± 0.01 vs. 1.17 ± 0.01 (leaves). Aphids confined to reproductive tissues with higher λ exhibited shorter generation times (T = 14.2 ± 0.2 days) and 53–75% higher net reproductive rates (R₀ = 23.3 ± 1.7) than aphids feeding on vegetative tissues. Prospective analyses showed that there was a nymph‐skewed stable stage distribution, and elasticity values revealed that λ is most sensitive to changes in stasis of adults staying in the adult stage and to adult survival. Retrospective analyses indicated that variation in adult fecundity (value of 0.05) had the largest effect on population dynamics but collectively, growth of nymphal stage 2–3, 3–4, and 4 to adult accounted for most of the difference in λ between the treatments. Monitoring programs should target adults and penultimate instars colonizing reproductive tissues of canola plants in the field as aphids on these plant structures contribute most to population growth.     

In vitro flowering of Boronia megastigma Nees. and the effect of 6-benzylaminopurine

Successful flower bud initiation and development was achieved on Boronia megastigma in vitro. The effect of cytokinin on flowering was investigated in environmental conditions that promote flowering as well as under conditions that stimulate vegetative growth (nonfloral promotory conditions). Flower initiation and differentiation was enhanced by cytokinin; however, many flower buds reverted when the media contained high levels of cytokinin. Anthesis occurred only on media that had no cytokinin added and under floral promotory conditions.     

The impact of Meu1-mediated resistance in tomato on longevity, fecundity and behavior of the potato aphid, Macrosiphum euphorbiae

The effect of the tomato resistance gene, Meu1, on feeding, longevity, fecundity and developmental rate of the pink biotype of the potato aphid, Macrosiphum euphorbiae (Thomas) (Hemiptera, Aphididae), was determined using nearly isogenic tomato (Lycopersicon esculentum Mill, Solanaceae) lines. Aphid mortality was significantly higher on resistant plants, with 60% of the aphids dying by the 4th day of exposure. By the 10th day, all the aphids on the resistant plants were dead whereas 100% of the aphids on susceptible plants were alive. Meu1-mediated resistance resulted in significantly decreased fecundity with a ten-fold decrease in the net fertility rate (4.5 and 45.7 progeny per aphid on resistant and susceptible tomato, respectively). A qualitative analysis showed that honeydew was produced by aphids on resistant and susceptible plants, suggesting that aphids initiate feeding on both plant types. However, significantly lower quantities of honeydew were present when aphids were caged on resistant plants. There were also significant differences in aphid location on resistant and susceptible leaves. Experiments evaluating behavior in less than 24 h showed that aphids left resistant leaves after relatively short exposure (3–6 h). Aphids transferred from resistant to susceptible tomato at intervals between 3 h and 24 h resumed feeding as evidenced by presence of honeydew. Although the mechanism by which Meu1-mediated resistance operates is not yet known, our data suggest that resistance factors act rapidly after initiation of feeding and that lower fecundity and longevity are related to reduction in aphid feeding.     

The use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L. (Coleoptera: Coccinellidae)

Abstract.  1. To clarify the use of honeydew in foraging for aphids by larvae of the ladybird beetle, Coccinella septempunctata L., searching behaviour of ladybird larvae for Aphis craccivora Koch and Acyrthosiphon pisum Harris and the abundance of honeydew under aphid colonies were examined in laboratory experiments.
2. More larvae responded by climbing the plants with aphids than responded to plants without aphids. When the plants were replaced with sticks, in order to exclude visual and olfactory cues from plants and aphids, more larvae of C. septempunctata climbed sticks above the area that contained honeydew than climbed sticks above the area that did not contain honeydew. Then, ladybird larvae use honeydew as a contact kairomone when foraging for aphids.
3.  Aphis craccivora deposited a larger number of honeydew droplets beneath the plants than did similar numbers of A. pisum. Thus, C. septempunctata larvae licked more frequently the honeydew of A. craccivora than that of A. pisum and spent longer searching on the area containing honeydew of A. craccivora than that of A. pisum . Consequently, a larger number of larvae climbed a stick above honeydew of A. craccivora than that of A. pisum.
4. It may be also considered that C. septempunctata larvae can distinguish honeydew of the two aphid species and respond more strongly to A. craccivora than A. pisum.     

Evidence for gibberellin-like substances in phloem exudate of higher plants

Summary Samples of sieve-tube sap were obtained as honeydew from aphids feeding on three species of higher plants. The honeydew was extracted, chromatographed and tested in several bioassays for the presence of gibberellin-like substances. The bioassay results indicated that gibberellin-like substances were translocated in the phloem of dandelion (Taraxacum officinale), broad bean (Vicia faba) and willow (Salix viminalis). Results obtained with willow showed that the concentration of gibberellin-like substances in the sieve-tube sap is daylength dependent, high levels being present in plants maintained under long days and low levels in short day plants.     

Olfactory response by the aphidophagous gall midge, Aphidoletes aphidimyza to honeydew from green peach aphid, Myzus persicae

Female adults of the aphidopagous gall midge, Aphidoletes aphidimyza (Rondani) (Diptera: Cecidomyiidae), showed an olfactory response to honeydew excreted by the green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae) under laboratory conditions. The response was only elicited by treatments with honeydew, whereas aphids, aphids with pepper plants or leaves, and pepper plants or leaves were not attractive to the midges. Dose‐dependent responses were observed from whole honeydew, honeydew volatiles extracted in pentane, and honeydew volatiles captured on Porapak Q®. When honeydew was eluted with three sequential pentane washes, a positive response was only observed from the midges for the first wash. Female midges laid more eggs on pepper plants infested with higher densities of M. persicae. The olfactory response of midges to honeydew is discussed with respect to prey location.     

Influence of aphid honeydew on the foraging behaviour of Hippodamia convergens larvae

1. Environmental cues associated with prey are known to increase predator foraging efficiency. Ladybird larvae are major predators of aphids. The sugary excretion of aphids (honeydew) has been proposed to serve as a prey‐associated cue for ladybird larvae. 2. Ladybird larvae are regularly found on the ground moving between plants or after falling off plants. The use of prey‐associated cues would be particularly beneficial for ladybird larvae on the ground in that such cues would help them to decide which plants to climb because aphids are patchily distributed within as well as amongst plants and, as a result, many plants are either not infested with aphids or do not host an aphid species of high nutritional value for ladybird larvae. 3. Laboratory experiments with larvae of Hippodamia convergens Guérin‐Méneville (Coleoptera: Coccinellidae) were carried out to explore whether honeydew accumulated on the ground is used as a foraging cue. The study also investigated whether, if honeydew is a foraging cue, larvae show differential responses to honeydew of high‐quality prey Acyrthosiphon pisum Harris compared with that of low‐quality prey Aphis fabae Scopoli (both: Homoptera: Aphididae). 4. Hippodamia convergens larvae stayed longer in areas containing honeydew but did not engage in longer bouts of searching. Furthermore, larvae did not distinguish between honeydew from high‐ and low‐quality aphid prey.     

Snowdrop lectin (Galanthus nivalis agglutinin) in aphid honeydew negatively affects survival of a honeydew- consuming parasitoid

1 Insecticidal proteins can be excreted in the honeydew when sap-sucking insects feed on insect-resistant transgenic plants. Honeydew can be an important source of carbohydrates, thus potentially exposing a broad range of honeydew-feeding insects to transgene products.
2 Snowdrop lectin ( Galanthus nivalis agglutinin; GNA) dissolved in a 2 m sucrose solution had no antifeedant effect on female aphid parasitoids ( Aphidius ervi ) but had a direct negative effect on their longevity.
3 When feeding on honeydew from Rhopalosiphum padi feeding on a GNA-containing artificial diet, Aphidius ervi suffered a longevity reduction that was more pronounced than was to be expected based on the detected GNA concentration in the honeydew.
4 Analysis of carbohydrate and amino acid composition revealed that a change in honeydew composition caused by a GNA-effect on the aphids could be a possible explanation for the additional reduction in parasitoid longevity.
5 When comparing the effect of honeydew from Sitobion avenae and R. padi feeding on GNA-expressing or nontransformed wheat plants on A. ervi longevity, aphid species was found to have a significant effect, whereas the wheat variety had no effect. The latter result was probably due to low GNA expression levels in the plants. Differences in nutritional suitability between honeydew from R. padi and S. avenae could be explained by differences in carbohydrate and amino acid composition.
6 This is the first study to demonstrate that GNA ingested by aphids and transported into the honeydew can negatively affect the parasitoids consuming this honeydew.
7 We recommend that honeydew should be considered as a route of exposure to transgene products in future risk assessment studies.     

A study on common bean cultivars to identify sources of resistance against the black bean aphid,Aphis fabae Scopoli (Hemiptera: Aphididae)

Black bean aphid is an important common bean pest in the world. Aphids damage crops both directly by their feeding and by spreading viruses. Also, they indirectly damages with excretion honeydew and smokes moulds growth through some physiological processes and transmission of viruses. Resistant cultivars application is the main strategy to control Aphis fabae Scopoli. In this experiment, different lines and cultivars were infested with five wingless adult aphids. After 7 and 14?days, plants were evaluated and aphid population was counted and recorded. After 45?days when plants reached to the flowering stage, test was repeated. Result showed that there was a significant difference among cultivars in number of adult aphid in two-leaf stage. Most of the number of adult at the first week in two-leaf stage observed on Goynok cultivar that had significant difference to other lines and cultivars. Number of adults and nymphs among the line and cultivar at flowering stage did not show significant difference. Number of deployed aphids after two weeks was significantly higher than the first week that observed in all cultivars. Totally, the most resistant cultivar was Sayad and the most susceptible cultivar was Goynok.     

The influence of aphids and honeydew on the leaving rate of searching aphid parasitoids from wheat plants

Dispersal cages were used to investigate the effects of aphids and treatment with artificial honeydew on the leaving rate of searching females of the parasitoid Aphidius rhopalosiphi from groups of wheat plants. Parasitoids which flew away from groups of plants placed in the centre of a cage were trapped on the sides and roof of the cage and thus were prevented from returning to the plants. The positions of trapped parasitoids suggested their direction of flight when dispersing from the plants. Parasitoids increased their residence times on groups of plants in the presence of aphids and of artificial honeydew, but the rate of parasitism of the host Sitobion avenae was not raised by the presence of artificial honeydew under the experimental conditions used. The direction of flight taken by the majority of parasitoids suggested that they were leaving the plants in order to locate further plants nearby to continue searching rather than to terminate searching and disperse away from the area. The need to consider plant patch size in studies of parasitoid searching behaviour is stressed.     

Evidence for the transfer of a soil‐borne contaminant from plants to ants via an aphid mediator

1. Uptake of environmental contaminants by lower trophic groups can have negative effects on higher trophic groups. This study tested the ability of selenium, an environmental contaminant found in high concentrations throughout the tissues of certain accumulating plants, to be transferred to ants via aphid tissue and honeydew. 2. Plants of the selenium accumulator, Raphanus sativus (wild radish), were watered with three different selenium treatments (0, 0.25 or 0.5 µg Se ml^?1). Aphids, Myzus persicae, and Argentine ant colonies, Linepithema humile, were added to each caged plant and allowed to interact freely. Ant colonies were supplemented with one of three different food options to encourage the consumption of aphids, aphid honeydew, or aphids and honeydew. 3. The accumulation of selenium by each trophic group and a trophic transfer factor (TTF) was calculated. The TTF for plants to aphids was > 1, indicating biomagnification, whereas the TTF for aphids to worker ants was < 1, indicating only biotransfer. Accumulated levels by worker ants did not statistically differ as a result of diet. 4. The amounts of selenium acquired by ants as a factor of diet and caste were compared. Plants, aphids and worker ants accumulated selenium in a dose‐dependent manner. Ant queens did not contain detectable amounts of selenium. Honeydew contained comparable amounts of selenium to plant selenium levels. 5. Access to toxic compounds via honeydew and insect protein may have negative effects on the range expansion of invasive species, such as the Argentine ant.     

Changes in Gene Expression from Diploid to Autotetraploid Status of Lycopersicon esculentum

The cytokinin activity of the root exudate, the leaves, and the apices of vegetative and flowering white lupin plants (Lupinus albus L.) was investigated. The level of cytokinin activity in the root exudate decreased over the 11-week experimental period. Four peaks of cytokinin activity were recorded in the root exudate of 8-week-old plants after fractionation on Sephadex LH-20. Two of these peaks co-eluted with zeatin and zeatin riboside. It is suggested that the remaining peaks represent nucleotide and glucoside cytokinins. The cytokinin levels in extracts of the mature leaves fluctuated slightly over the experimental period. Three peaks of activity co-eluting with zeatin, zeatin riboside and the glucoside cytokinins were recorded in extracts of mature leaves of 8-week-old plants. In the apices cytokinin activity could only be detected in the inflorescences of flowering plants. It would appear that cytokinins produced by the roots accumulate in the fully expanded mature leaves, but are utilized in the rapidly growing apical region and in young expanding leaves.     

The effects of barley yellow dwarf virus, aphids and honeydew on Cladosporium infection of winter wheat and barley

Prior infection of both wheat and barley plants by BYDV predisposed their ears to infection by Cladosporium spp. and Verticillium spp. Aphids and honeydew increased the incidence of Cladosporium on wheat ears but not on barley. This difference between crops was attributed to the larger number ot aphids on the wheat. In the glasshouse, aphids and honeydew, but not honeydew alone, increased Cladosporium populations.     

Induction of early flowering in Cymbidium niveo-marginatum Mak in vitro

 Many orchids take several years to flower. We have been able to induce early flowering in the temperate orchid Cymbidium niveo-marginatum Mak in vitro. The combined treatment of cytokinin (6-benzylaminopurine), restricted nitrogen supply with phosphorus enrichment, and root excision (pruning) induced transition of the Cymbidium shoot from a vegetative to a reproductive stage. Nearly 100% of the plants flowered within 90 days only when the combined treatment was applied. When root excision and/or 6-benzylaminopurine were omitted from the combined treatments, flower induction was significantly reduced. The auxin transport inhibitor, 2,3,5-triiodobenzoic acid prevented flowering of Cymbidium in vitro, although auxin (α-naphthaleneacetic acid) itself did not induce flowering. Gibberellic acid markedly delayed flowering in C. niveo-marginatum even when the flower-promoting treatment was applied. Paclobutrazol, an anti-gibberellin agent, totally blocked the inductive effects of either cytokinin or pruning. These observations suggest that concerted actions of auxin, cytokinin, and gibberellin, as well as nutrient concentration and putative promoting/suppressing agents, determine the timing of Cymbidium orchid transition from the vegetative to reproductive stage. Received: 22 July 1998 / Revision revised: 23 November 1998 / Accepted: 2 December 1998     

Spruce stands provide natural food sources to adult hymenopteran parasitoids of bark beetles

Adults of synovigenic hymenopteran parasitoids of bark beetles need supplemental feeding to maximise their lifetime realised fecundity, and yet little is known about the natural sources that provide this food. Here we show that, in spite of the reduced diversity of flowering plants expected in even-aged spruce plantations, several plant species are present that might provide nectar and pollen to the hymenopteran parasitoid complex of Ips typographusL. (Coleoptera: Scolytidae), throughout the growing season. Conifer aphids are also abundant in these stands and produce large amounts of honeydew, which might also be consumed by the parasitoids. Laboratory experiments demonstrate that longevity is significantly increased in Coeloides bostrychorum Giraud (Hymenoptera: Braconidae) when the wasps are exposed to flowers of various species commonly found in spruce stands.