Comparative analysis of Solanum stoloniferum responses to probing by the green peach aphid Myzus persicae and the potato aphid Macrosiphum euphorbiae

Abstract Plants protect themselves against aphid attacks by species‐specific defense mechanisms. Previously, we have shown that Solanum stoloniferum Schlechtd has resistance factors to Myzus persicae Sulzer (Homoptera: Aphididae) at the epidermal/mesophyll level that are not effective against Macrosiphum euphorbiae Thomas (Homoptera: Aphididae). Here, we compare the nymphal mortality, the pre‐reproductive development time, and the probing behavior of M. persicae and M. euphorbiae on S. stoloniferum and Solanum tuberosum L. Furthermore, we analyze the changes in gene expression in S. stoloniferum 96 hours post infestation by either aphid species. Although the M. euphorbiae probing behavior shows that aphids encounter more probing constrains on phloem activities–longer probing and salivation time– on S. stoloniferum than on S. tuberosum, the aphids succeeded in reaching a sustained ingestion of phloem sap on both plants. Probing by M. persicae on S. stoloniferum plants resulted in limited feeding only. Survival of M. euphorbiae and M. persicae was affected on young leaves, but not on senescent leaves of S. stoloniferum. Infestation by M. euphorbiae changed the expression of more genes than M. persicae did. At the systemic level both aphids elicited a weak response. Infestation of S. stoloniferum plants with a large number of M. persicae induced morphological changes in the leaves, leading to the development of pustules that were caused by disrupted vascular parenchyma and surrounding tissue. In contrast, an infestation by M. euphorbiae had no morphological effects. Both plant species can be regarded as good host for M. euphorbiae, whereas only S. tuberosum is a good host for M. persicae and S. stoloniferum is not. Infestation of S. stoloniferum by M. persicae or M. euphorbiae changed the expression of a set of plant genes specific for each of the aphids as well as a set of common genes.     

Behavioural responses of the aphid parasitoid Diaeretiella rapae to volatiles from Arabidopsis thaliana induced by Myzus persicae

In response to herbivory by insects, several plant species have been shown to produce volatiles that attract the natural enemies of those herbivores. Using a Y‐tube olfactometer, we investigated responses of the aphid parasitoid Diaeretiella rapae MacIntosh (Hymenoptera: Aphidiidae) to volatiles from Arabidopsis thaliana Columbia (Brassicaceae) plants that were either undamaged, infested by the peach‐potato aphid, Myzus persicae Sulzer (Homoptera: Aphididae), or mechanically damaged, as well as to volatiles from just the aphid or its honeydew. In dual‐choice experiments, female D. rapae given oviposition experience on A. thaliana infested with M. persicae were significantly attracted to volatiles from A. thaliana infested with M. persicae over volatiles from undamaged A. thaliana and similarly were significantly attracted to plants that had been previously infested by M. persicae, but from which the aphids were removed, over undamaged plants. Diaeretiella rapae did not respond to volatiles from M. persicae alone, their honeydew, or plants mechanically damaged with either a pin or scissors. We conclude that an interaction between the plant and the aphid induces A. thaliana to produce volatiles, which D. rapae can learn and respond to. Poor responses of D. rapae to volatiles from an A. thaliana plant that had two leaves infested with M. persicae, with the two infested leaves being removed before testing, suggested the possibility that, at this stage of infestation, the majority of volatile production induced by M. persicae may be localized to the infested tissues of the plant. We conclude that this tritrophic interaction is a suitable model system for future investigations of the biochemical pathways involved in the production of aphid‐induced volatiles attractive to natural enemies.     

Response of Solanum tuberosum to Myzus persicae infestation at different stages of foliage maturity

Young leaves of the potato Solanum tuberosum L. cultivar Kardal contain resistance factors to the green peach aphid Myzuspersicae （Sulzer） （Hemiptera： Aphididae） and normal probing behavior is impeded. However, M. persicae can survive and reproduce on mature and senescent leaves of the cv. Kardal plant without problems. We compared the settling ofM. persicae on young and old leaves and analyzed the impact of aphids settling on the plant in terms of gene expression. Settling, as measured by aphid numbers staying on young or old leaves, showed that after 21 h significantly fewer aphids were found on the young leaves. At earlier time points there were no difference between young and old leaves, suggesting that the young leaf resistance factors are not located at the surface level but deeper in the tissue. Gene expression was measured in plants at 96 h postinfestation, which is at a late stage in the interaction and in compatible interactions this is long enough for host plant acceptance to occur. In old leaves of cv. Kardal （compatible interaction）, M. persicae infestation elicited a higher number of differentially regulated genes than in young leaves. The plant response to aphid infestation included a larger number of genes induced than repressed, and the proportion of induced versus repressed genes was larger in young than in old leaves. Several genes changing expression seem to be involved in changing the metabolic state of the leaf from source to sink.     

Immediate alteration of Macrosiphum euphorbiae host plant-selection behaviour after biotic and abiotic damage inflicted to potato plants

The effects of potato [Solanum tuberosum L. (Solanaceae)] plant damage on the host plant-selection behaviour of the potato aphid, Macrosiphum euphorbiae Thomas (Homoptera: Aphididae), were studied. The damage inflicted to the plant was only of short duration and observations on aphid behaviour were made immediately following plant damage. The underlying questions of the study were to know how much time it takes for plant defence mechanisms to be activated and if this activation had noticeable repercussions on aphid behaviour. We considered stresses of various natures: biotic (pre-infestation by conspecifics or by Colorado potato beetles) and abiotic (scissor cuts). Aphid responses to host plant semiochemicals were investigated using a darkened arena bioassay and the probing behaviour was assessed using the electrical penetration graph technique. Aphids were attracted to their host plant (undamaged or damaged). In a preference test (undamaged plant vs. damaged plant), plants previously infested by conspecifics were preferred to undamaged plants, but this preference was not observed for heterospecific and abiotic damage. However, aphid probing behaviour was not modified on plants previously infested by conspecifics, whereas some changes were observed subsequently to heterospecific and abiotic damages. Our data present evidence that plants can respond to biotic and abiotic stresses soon after the damage is inflicted and when the damage is of short duration. The diverse consequences of these various local plant responses on M. euphorbiae behaviour are discussed in the context of plant defence strategies against aphid colonization.     

The effect of co-infestation by conspecific and heterospecific aphids on the feeding behaviour of <Emphasis Type="Italic">Nasonovia ribisnigri</Emphasis> on resistant and susceptible lettuce cultivars

Aphid saliva can suppress the blocking of sieve elements, a reaction that plants employ to inhibit aphid feeding, but aphid saliva can also elicit plant defence responses. Such plant responses might affect interactions between different aphid species and intraspecifically, e.g. among different biotypes. The objectives of our study were to investigate if feeding behaviour and performance of two biotypes of the lettuce aphid Nasonovia ribisnigri are affected by (1) feeding by the other biotype and (2) feeding by the green peach aphid Myzus persicae or the potato aphid Macrosiphum euphorbiae. Additionally the effect of feeding in a group was studied. All experiments were performed on both a resistant and an isogenic susceptible lettuce cultivar. Feeding or probing by conspecific or heterospecific aphids had different effects on Nasonovia ribisnigri biotypes. Aphids were only slightly affected by feeding or probing of the same biotype on both susceptible and resistant lettuce. N. ribisnigri virulent biotype Nr:1 suppressed the resistance against Nr:0 in the resistant cultivar. In contrast, defence was induced by Nr:1 against Nr:0 in susceptible lettuce. Co-infestation by M. euphorbiae and M. persicae had minor effects on Nr:0. Defence against Nr:1 was induced on both susceptible lettuce and resistant lettuce by Nr:0 and M. euphorbiae. Additionally, M. persicae induced defence in resistant lettuce against Nr:1. Effectors in the saliva of Nr:1 aphids are likely responsible for the defence suppression in lettuce. Identification of these effectors could lead to a better understanding of the mechanism of virulence in N. ribisnigri.     

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 induction of top-roll symptoms on potato plants by the aphid Macrosiphum euphorbiae

In the field, caged potato plants of King Edward and Majestic cultivars infested with the potato aphid Macrosiphum euphorbiae developed top-roll symptoms, the proportion of affected plants increasing with the size and persistence of the aphid population. Yield of tubers from plots in which 90% of the plants had top-roll symptoms was 40% less than that from control plots; yield of saleable ware was even less. Foliage produced after the aphids had been killed was symptomless even when it arose from the axil of an affected leaf. Caged field plants treated with phorate granules to prevent aphid attack did not develop top-roll. Prolonged infestation of Pentland Crown, Majestic and King Edward plants by M. euphorbiae in a glasshouse induced rolling of upper leaves similar to top-roll of field plants. Experimental results suggest that rolling was directly attributable to heavy attack by M. euphorbiae, not to an aphid-transmitted pathogen.     

Evaluation of induced responses, insect population growth, and host-plant fitness may change the outcome of tests of the preference-performance hypothesis: a case study

The preference‐performance hypothesis predicts that insect preference should correspond to host suitability for offspring development. We studied the pattern of within‐plant preference in the aphid Sipha flava and its consequences for offspring performance on the host‐plant Sorghum halepense, regarding the role of induced responses of plants to aphid feeding. The consequences of within‐plant preference on aphid population growth and host‐plant traits were also evaluated. Our results showed that winged and wingless aphids preferred to settle on mature rather than young leaves. In contrast, aphid individual growth rate was higher on young leaves when compared with mature leaves, suggesting that the outcome of this test rejected the preference‐performance hypothesis. However, the inclusion of the factor ‘previous aphid infestation’ changed the outcome from a maladaptive choice to a neutral one. Thus, individual growth rates of S. flava increased when aphids developed on leaves that had been previously infested. Interestingly, aphid growth rate on previously infested leaves did not differ between young and mature leaves. On the other hand, aphid population reproductive rate was higher and the percentage of winged aphids lower when infestation occurred on mature rather than young leaves. Aphid infestation reduced plant and shoot biomass, and increased leaf mortality. These negative effects on plant traits related to plant fitness were greater when aphid infestation occurred on young leaves. Likewise, whereas infestation on mature leaves did not cause a significant reduction in the number of flowering plants compared with control plants, aphid infestation on young leaves did reduce the number of plants at the flowering stage. Consequently, if both the reproductive rate of aphids in the mid‐term, and host‐plant fitness are taken into account, the results indicate that aphid preference for mature leaves may be an adaptive choice, thus supporting the preference‐performance hypothesis.     

Quantitative studies on the uptake and retention of potato leafroll virus by aphids in laboratory and field conditions

Enzyme-linked immunosorbent assay (ELISA) was adapted for the efficient detection and assay of potato leafroll virus (PLRV) in aphids. Best results were obtained when aphids were extracted in 0.05 M phosphate buffer, pH 7.0, and the extracts incubated at 37 °C for 1 h before starting the assay. Using batches of 20 green peach aphids (Myzus persicae), about 0.01 ng PLRV/aphid could be detected. The virus could also be detected in single aphids allowed a 1-day acquisition access period on infected potato leaves. The PLRV content of aphids depended on the age of potato source-plants and the position of source leaves on them. It increased with increase in acquisition access period up to 7 days but differed considerably between individual aphids. A maximum of 7 ng PLRV/aphid was recorded but aphids more usually accumulated about 0.2 ng PLRV per day. When aphids were allowed acquisition access periods of 1–3 days, and then caged singly on Physalis floridana seedlings for 3 days, the PLRV content of each aphid, measured subsequently, was not strongly correlated with the infection of P. floridana. The concentration of PLRV in leaf extracts differed only slightly when potato plants were kept at 15, 20, 25 or 30 °C for 1 or 2 wk, but the virus content of aphids kept on leaves at the different temperatures decreased with increase of temperature. PLRV was transmitted readily to P. floridana at all temperatures, but by a slightly smaller proportion of aphids, and after a longer latent period, at 15 °C than at 30 °C. The PLRV content of M. persicae fed on infected potato leaves decreased with increasing time after transfer to turnip (immune to PLRV). The decrease occurred in two phases, the first rapid and the second very slow. In the first phase the decrease was faster, briefer and greater at 25 and 30 °C than at 15 and 20 °C. No evidence was obtained that PLRV multiplies in M. persicae. These results are compatible with a model in which much of the PLRV in aphids during the second phase is in the haemocoele, and transmission is mainly limited by the rate of passage of virus particles from haemolymph to saliva. The potato aphid, Macrosiphum euphorbiae, transmitted PLRV much less efficiently than M. persicae. Its inefficiency as a vector could not be ascribed to failure to acquire or retain PLRV, or to the degradation of virus particles in the aphid. Probably only few PLRV particles pass from the haemolymph to saliva in this species. The virus content of M. euphorbiae collected from PLRV-infected potato plants in the field increased from early June to early July, and then decreased. PLRV was detected both in spring migrants collected from the plants and in summer migrants caught in yellow water-traps. PLRV was also detected in M. persicae collected from infected plants in July and August, and in trapped summer migrants, but their PLRV content was less than that of M. euphorbiae, and in some instances was too small for unequivocal detection.     

Infection of potato plants with potato leafroll virus changes attraction and feeding behaviour of Myzus persicae

Potato leafroll virus (PLRV; genus Polerovirus, family Luteoviridae) is a persistently transmitted circulative virus that depends on aphids for spreading. The primary vector of PLRV is the aphid Myzus persicae (Sulzer) (Homoptera: Aphididae). Solanum tuberosum L. potato cv. Kardal (Solanaceae) has a certain degree of resistance to M. persicae: young leaves seem to be resistant, whereas senescent leaves are susceptible. In this study, we investigated whether PLRV‐infection of potato plants affected aphid behaviour. We found that M. persicae's ability to differentiate headspace volatiles emitted from PLRV‐infected and non‐infected potato plants depends on the age of the leaf. In young apical leaves, no difference in aphid attraction was found between PLRV‐infected and non‐infected leaves. In fact, hardly any aphids were attracted. On the contrary, in mature leaves, headspace volatiles from virus infected leaves attracted the aphids. We also studied the effect of PLRV‐infection on probing and feeding behaviour (plant penetration) of M. persicae using the electrical penetration graph technique (DC system). Several differences were observed between plant penetration in PLRV‐infected and non‐infected plants, but only after infected plants showed visual symptoms of PLRV infection. The effects of PLRV‐infection in plants on the behaviour of M. persicae, the vector of the virus, and the implications of these effects on the transmission of the virus are thoroughly discussed.     

Genetic, biochemical, and behavioral uniformity among populations of Myzus nicotianae and Myzus persicae

Prior to designation as distinct species, an appellation presently in question, the tobacco aphid, Myzus nicotianae Blackman (Homoptera: Aphididae), was classified as a tobacco-feeding form of the green peach aphid, Myzus persicae (Sulzer). In this study, RAPD polymorphisms distinguished members of the Myzus persicae complex (M. persicae and M. nicotianae) from three outgroup Myzus species (M. cerasi (F.), M. hemerocallis Takahashi, and M. varians Davidson). Polymorphisms within the complex did not separate populations on the basis of host association (tobacco versus other host plants) or geographic origin (collections from the United States, Europe, and Japan). Similarly, while GC-MS analysis of cuticular hydrocarbon profiles revealed both developmental and inter-populational differences within the M. persicae complex, it did not separate populations of tobacco feeding aphids from those collected off non-tobacco hosts. Finally, with the exception of their responses to a choice between lettuce and collards, the host preference behavior of a green peach aphid population, a red tobacco aphid population, and a green tobacco aphid population was indistinguishable in host preference experiments. These results add to a growing body of evidence suggesting M. nicotianae and M. persicae are conspecific.     

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.     

Aphid populations showing differential levels of virulence on Capsicum accessions

The green peach aphid,Myzus persicae,is one of the most threatening pests in pepper cultivation and growers would benefit from resistant varietices.Previously,we identified two Capsicum acessions as susceptible and three as resistant to M.persicae using an aphid population originating from the Netherlands(NL).Later on we identified an aphid population originating from a diferent gcographical region(Switserland,SW)that was virulent on all tested Capsicum acessions.The objeetive of the current work is to describe in detail diferent aspects of the interaction between two aphid populations and two sclected Capsicum acessions(one that was susceptible[PB2013046]and one that was resistant[PB2013071]to population NL),including biochemical processes involved.Electrical penetration graph(EPG)recordings showed similar feeding activities for both aphid populations on PB2013046.On acession PB2013071 the aphid population sw was able to devote significantly more time to phloem ingestion than population NL.We also studied plant defense response and found that plants of acession PB2013046 could not induce an accumulation of reactive oxygen species and callose formation after infestation with either aphid population.However,plants of PB2013071 induced a stronger defense response after infestation by population NL than after infestation by population SW.Based on these results,population SW of M.persicae seems to have overcome the resistance of PB2013071 that prevented feeding of aphids from NL population.The potential mechanism by which SW population overcomes the resistance is discussed.     

Olfactory responses of Aphidius gifuensis to odors of host plants and aphid‐plant complexes

Abstract The olfactory responses of Aphidius gifuensis to odors from two host plants (Nicotiana tabacum and Brassica napus ssp.) and their complexes with different infestation levels of two host aphids (Myzus persicae and Lipaphis erysimi) were respectively examined in an olfactometer. The results showed that female A. gifuensis did not respond to odors of undamaged or mechanically damaged host plants, but significantly responded to odors of aphid/plant complexes. Moreover, A. gifuensis responded significantly to odors of both M. persicae and L. erysimi/plant complexes when host plants were infested by high levels of aphids, suggesting that quantity of aphid‐induced volatiles could be important for attracting A. gifuensis. When tested between aphid/plant complexes, A. gifuensis did not show its preference for either complex. The efficiency of A. gifuensis against aphids in open fields potentially could be improved by using its olfactory response to aphid/plant complexes.     

Behavioural responses of the seven‐spot ladybird Coccinella septempunctata to plant headspace chemicals collected from four crop Brassicas and Arabidopsis thaliana,infested with Myzus persicae

1 Insects using olfactory stimuli to forage for prey/hosts are proposed to encounter a ‘reliability–detectability problem’, where the usability of a stimulus depends on its reliability as an indicator of herbivore presence and its detectability. 2 We investigated this theory using the responses of female seven‐spot ladybirds Coccinella septempunctata (Coleoptera: Coccinellidae) to plant headspace chemicals collected from the peach‐potato aphid Myzus persicae and four commercially available Brassica cultivars; Brassica rapa L. cultivar ‘turnip purple top’, Brassica juncea L. cultivar ‘red giant mustard’, Brassica napus L. cultivar ‘Apex’, Brassica napus L. cultivar ‘Courage’ and Arabidopsis thaliana. For each cultivar/species, responses to plants that were undamaged, previously infested by M. persicae and infested with M. persicae, were investigated using dual‐choice Petri dish bioassays and circular arenas. 3 There was no evidence that ladybirds responded to headspace chemicals from aphids alone. Ladybirds significantly preferred headspace chemicals from B. napus cv. Apex that were undamaged compared with those from plants infested with aphids. For the other four species/cultivars, there was a consistent trend of the predators being recorded more often in the half of the Petri dish containing plant headspace chemicals from previously damaged and infested plants compared with those from undamaged ones. Furthermore, the mean distance ladybirds walked to reach aphid‐infested A. thaliana was significantly shorter than to reach undamaged plants. These results suggest that aphid‐induced plant chemicals could act as an arrestment or possibly an attractant stimulus to C. septempunctata. However, it is also possible that C. septempunctata could have been responding to aphid products, such as honeydew, transferred to the previously damaged and infested plants. 4 The results provide evidence to support the ‘reliability–detectability’ theory and suggest that the effectiveness of C. septempunctata as a natural enemy of aphids may be strongly affected by which species and cultivar of Brassica are being grown.     

Induced resistance by Myzus persicae in the peach cultivar `Rubira'

The effect of a previous infestation by the green peach aphid Myzus persicae (Sulzer) on the settling behaviour and reproduction of the same aphid species was investigated in the resistant peach cultivar Rubira, and compared with that observed in the susceptible control cultivar GF305. A previous infestation of 48 h triggered induced resistance in Rubira. There were significantly fewer aphids settling on preinfested than on uninfested plants, indicating an increased rejection of Rubira as a host plant. The level of induced resistance in preinfested plants was positively related to the duration of the first infestation. In GF305, previous infestation had no detrimental effect on aphid settlement and even slightly enhanced larviposition by adult females. The aphid probing behaviour after a 48-h preinfestation was also monitored for 8 h with the electrical peneration graph (EPG) technique. On preinfested GF305, most EPG parameters indicated an enhanced host plant acceptance. On preinfested GF305, aphids produced less sieve element salivation and more continuous sap ingestion than on uninfested GF305, indicating that the previous aphids provoked changes in plant properties beneficial to the test aphids. In Rubira, a major induced factor of resistance was thought to be expressed in the sieve element as phloem sap ingestion was 4-fold shorter on preinfested than on uninfested plants. The time taken by the aphid stylets to reach a sieve element was also significantly increased on preinfested Rubira, suggesting the induction of resistance factors outside the phloem. The originality of the Rubira/M. persicae interaction is discussed in the perspective of a better understanding of plant induced responses to aphids.     

Effects of systemic potato response to wounding and jasmonate on the aphid Macrosiphum euphorbiae (Sternorryncha: Aphididae)

Plant induced responses are activated by multiple biotic and abiotic stresses, and may affect the interactions between a plant and phytophagous insects. The objective of this work was to evaluate the effects of different stresses inflicted to potato plants (Solanum tuberosum) on the potato aphid (Macrosiphum euphorbiae). Abiotic wounding, biotic wounding by Leptinotarsa decemlineata and treatment with volatile methyl jasmonate (MeJA) were evaluated with regard to the orientation behaviour, the feeding behaviour and the development of the potato aphids. Dual‐choice olfactometry showed that plants treated with MeJA lost their attractiveness for the potato aphids, while both abiotic and biotic wounding did not alter the orientation of aphids. Electropenetrography revealed that the feeding behaviour of aphids was only slightly disturbed by a previous L. decemlineata wounding, while it was highly disturbed by mechanical wounding and MeJA treatment. Aphid nymph survival was reduced on mechanically wounded plants, the pre‐reproductive period was lengthened and the fecundity reduced on plants treated with MeJA. Our results bring new information about the effects of various stresses inflicted to S. tuberosum on M. euphorbiae. We showed that wounding and MeJA treatment induced an antixenosis resistance in potato plants against M. euphorbiae, which may influence aphid colonization processes.     

Patterns in aphid honeydew production parallel diurnal shifts in phloem sap composition

Variation in phloem sap composition is important in determining aphid performance and is known to occur at both diurnal timescales and in response to plant age. For field grown potato plants, Solanum tuberosum L. (Solanaceae), we determined diurnal variation in components of phloem sap, measured by ethylene diamine tetra‐acetate exudation, and tested for impacts of plant age. The effects of plant age and diurnal cycles on honeydew production by Macrosiphum euphorbiae (Thomas) and Myzus persicae (Sulzer) (both Hemiptera: Aphididae) were also quantified. Both the ratio of sucrose to amino acids and the composition of amino acids in phloem sap varied significantly with time of day. Dietary essential amino acids contributed a smaller proportion of amino acids in the phloem sap of older plants and during early phases of the diurnal cycle. The only significant effect on aphid honeydew production was of the diurnal cycle for Ma. euphorbiae, although increased honeydew production during the day when compared with the production at night, was consistent across the two species. In contrast with studies carried out at seasonal scales, we found limited evidence for variation in phloem sap composition in response to plant age, consistent with our results for honeydew production. These data highlight the need for improved understanding of how seasonal and diurnal physiology of plants influence performance in phloem sap feeding insects.     

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.     

Probing behavior of aposymbiotic green peach aphid (Myzus persicae) on susceptible Solanum tuberosum and resistant Solanum stoloniferum plants

The green peach aphid, Myzus persicae Sulzer (Hemiptera: Aphididae) is one of the potato important pests; it is the most efficient vector of potato viruses. Myzus persicae harbors the endosymbiotic bacteria Buchnera aphidicola which supplements their diet. There is increasing evidence that B. aphidicola is involved in plant–aphid interactions and we previously demonstrated that B. aphidicola disruption (aposymbiosis) affected the probing behavior of M. persicae on radish plants, delaying host plant acceptance. In this work, we evaluated the effect of aposymbiosis on the probing behavior of M. persicae on 2 Solanum species with different compatibility with M. persicae, Solanum tuberosum (susceptible) and Solanum stoloniferum (resistant) with the electrical penetration graph technique (EPG). To disrupt B. aphidicola, rifampicin was administered to aphids through artificial diets. Aposymbiotic aphids, on both plant species, showed increased pathway activities, mechanical problems with the stylets, and delayed salivation in the phloem. The extended time in derailed stylet mechanics affected the occurrence of most other probing activities; it delayed the time to the first phloem phase and prevented ingestion from the phloem. The effect of aposymbiosis was more evident in the compatible interaction of M. persicae–S. tuberosum, than in the incompatible interaction with S. stoloniferum, which generated the M. persicae–S. tuberosum interaction to become incompatible. These results confirm that B. aphidicola is involved in the plant–aphid interaction in relation to plant acceptance, presumably through a role in stylets penetration in the plant.