Reproduction and probing behaviour of the bird cherry‐oat aphid Rhopalosiphum padi on detached leaves and leaves on intact seedlings of the winter host,bird cherry Prunus padus

Video recording and electrical penetration graph techniques are used simultaneously to investigate host acceptance (in terms of reproduction) and probing activities of autumn migrants (gynoparae) of the bird cherry‐oat aphid Rhopalosiphum padi (L.) on detached leaves and on leaves on intact plants of the winter host, bird cherry Prunus padus. There are no significant differences between the times taken to first parturition or the number of nymphs produced over a 6‐ or 18‐h period on intact plants or detached leaves. Stylet probing activities (i.e. total probe duration, xylem ingestion, the time to first phloem contact and phloem salivation) are also similar in the two situations.     

Immunocytochemical Localization of Prunasin Hydrolase and Mandelonitrile Lyase in Stems and Leaves of Prunus serotina

In macerates of black cherry (Prunus serotina Ehrh.) leaves and stems, (R)-prunasin is catabolized to HCN, benzaldehyde, and D-glucose by the sequential action of prunasin hydrolase (EC 3.2.1.21) and (R)-(+)-mandelonitrile lyase (EC 4.1.2.10). Immuno-cytochemical techniques have shown that within these organs prunasin hydrolase occurs within the vacuoles of phloem parenchyma cells. In arborescent leaves, mandelonitrile lyase was also located in phloem parenchyma vacuoles, but comparison of serial sections revealed that these two degradative enzymes are usually localized within different cells.     

Host acceptance by aphids: probing and larviposition behaviour of the bird cherry-oat aphid, Rhopalosiphum padi on host and non-host plants

The probing and larviposition behaviour of the bird cherry-oat aphid, Rhopalosiphum padi on summer and winter host plants were investigated using electrical penetration graph (EPG) coupled with simultaneous video recording. In this way the precise probing history prior to parturition can be monitored and the location of possible reproductive stimulants identified. On the host plant, all gynoparae (autumn winged females that give birth to sexual females on bird cherry, Prunus padus, the primary host) and 55% of winged virginoparae (summer females which produce further virginoparae on barley, Hordeum vulgare, a secondary host) initiated larviposition before phloem contact. However, 90% of wingless virginoparae (on barley) contacted the phloem before first larviposition whilst 10% did not. Thus, phloem contact does not appear to be a pre-requisite for these aphid forms to initiate reproduction.     

Are primary woody hosts ‘island refuges’ for host‐alternating aphids and important for colonization of local cereals?

Only few studies are available dealing with the relation between winter host density and spatial distribution and spring colonization of winter cereals by the host‐alternating cereal aphid species Rhopalosiphum padi and Metopolophium dirhodum. Large‐scale studies in climatically different agroecosystems in Germany from 2004 to 2006 revealed for R. padi and M. dirhodum larger spring/summer populations in landscapes with higher densities of winter hosts. A small‐scale study was performed in winter wheat fields adjacent to a large hedge with several typical winter hosts plants, bird cherry (Prunus padus) and wild rose species (Rosa spp.) to indentify distance effects (0–8, 8–24 and 24–60 m). Weekly measurements of aphid density between May to July showed significantly higher densities of R. padi compared with those of other aphids. Statistical analysis (Tukey–Kramer test and regression analyses) revealed significant gradients from the hedge to the field centre for R. padi and M. dirhodum. In comparative studies, winged R. padi from winter and adjacent summer hosts were genotyped using four microsatellite markers. The results showed that individuals from a certain winter host were not genetically similar with individuals from neighbouring summer hosts; it, therefore, seems that winter host clones did not significantly contribute to population built‐up in cereal fields over short distances. It could be concluded that on a regional scale, the density of sources for early migrants of R. padi is important for colonization intensity of surrounding summer hosts, but that the high local movement intensity and the relative small proportion of aphids that could be analysed in such tracking studies are blurring close spatial relations within short time periods.     

Characterisation of bird cherry-oat aphid (Rhopalosiphum padi L.) behaviour and aphid host preference in relation to partially resistant and susceptible wheat landraces

The bird cherry-oat aphid (Rhopalosiphum padi L.) is a major pest of wheat (Triticum aestivum L.) and can cause up to 30% yield losses. Heritable plant resistance to aphids is both an economically and ecologically sound method for managing aphids. Here we report how the behaviour and performance of R. padi differs on two resistant, one susceptible wheat landrace and a susceptible elite wheat variety. Feeding behaviour differed among the genotypes, with aphids on resistant lines spending longer in the pathway phase and less time phloem feeding. These behaviours suggest that both inter- and intracellular factors encountered during pathway and phloem feeding phases could be linked to the observed aphid resistance. Locomotion and antennal positioning choice tests also revealed a clear preference for susceptible lines. Although feeding studies revealed differences in the first probe indicating that the resistance factors might also be located in the peripheral layers of the plant tissue, scanning electron microscopy revealed no difference in trichrome length and density on the surface of leaves. Aphids are phloem feeders and limiting the nutrient uptake by the aphids may negatively affect their growth and development as shown here in lower weight and survival of nymphs on resistant genotypes and decreased reproductive potential, with lowest mean numbers of nymphs produced by aphids on W064 (54.8) compared to Solstice (71.9). The results indicate that resistant lines markedly alter the behaviour, reproduction and development potential of R. padi and possess both antixenosis and antibiosis type of resistance.     

Tissue and Subcellular Localization of Enzymes Catabolizing (R)-Amygdalin in Mature Prunus serotina Seeds

In black cherry (Prunus serotina Ehrh.) homogenates, (R)-amygdalin is catabolized to HCN, benzaldehyde, and d-glucose by the sequential action of amygdalin hydrolase, prunasin hydrolase, and mandelonitrile lyase. The tissue and subcellular localizations of these enzymes were determined within intact black cherry seeds by direct enzyme analysis, immunoblotting, and colloidal gold immunocytochemical techniques. Taken together, these procedures showed that the two β-glucosidases are restricted to protein bodies of the procambium, which ramifies throughout the cotyledons. Although amygdalin hydrolase occurred within the majority of procambial cells, prunasin hydrolase was confined to the peripheral layers of this meristematic tissue. Highest levels of mandelonitrile lyase were observed in the protein bodies of the cotyledonary parenchyma cells, with lesser amounts in the procambial cell protein bodies. The residual endosperm tissue had insignificant levels of amygdalin hydrolase, prunasin hydrolase, and mandelonitrile lyase.     

Odour stimuli affecting autumn migration of Rhopalosiphum padi (L.) (Hemipter: ornoptera)

R. padi gynoparae settling in the canopy of the winter host, Prunus padus, tended to form aggregations. Differences in the degree of exposure to wind, sun, etc., did not satisfactorily explain the aggregated pattern observed. In laboratory experiments, aphids did not distinguish between leaves taken from nearby aggregations and leaves taken far away from aggregations. The results of laboratory olfactometer tests and field trapping studies indicate that R. padi gynoparae produce an aggregation pheromone. Thus the aggregated settling pattern of gynoparae of R. padi can best be explained as a response to an aggregation pheromone.     

Effect of mineral oils on host plant selection and probing behavior of Rhopalosiphum padi

Aphids that colonize and reproduce on potato are some of the most efficient vectors of Potato virus Y (PVY) (Potyviridae: Potyvirus), and hence these aphids have been the focus of the majority of studies to date. However, other non‐colonizing aphids can also function as vectors. Mineral oil is the only product available to growers that effectively prevents the spread of PVY in potato seed production. Most previous studies focused on the effect of mineral oil on the behavior of aphids on their preferential host plant, and consequently there is a lack of information for non‐colonizing aphids on potato plants. The objective of this study was to determine the effect of spraying potatoes with one of two mineral oils, Superior 70 or Vazyl‐Y, on host selection and probing behavior of the non‐colonizing aphid Rhopalosiphum padi (L.) (Hemiptera: Aphididae). The electrical penetration graph (EPG) technique, combined with ethological observations, determined that there was no difference in R. padi behavior on potato plants treated with Superior 70. However, there were few significant changes in R. padi behavior on plants sprayed with Vazyl‐Y, including a delay in the initiation of stylet penetration and an increase in the duration of xylem sap ingestion. These new data support previous results and confirm that the mode of action of mineral oil in the reduction of the spread of PVY is not solely due to the modification of the behavior of aphids.     

The life-cycle and host preferences of the bird cherry-oat aphid, Rhopalosiphum padi L., and their bearing on the theories of host alternation in aphids

Emigrants of the host-alternating bird cherry-oat aphid fly from the primary host to various grasses before the end of June, even in the absence of natural enemies on bird cherry. They fly before the leaves of bird cherry mature and at a time when numbers of insect predators are rapidly increasing. In the autumn gynoparae and males return to bird cherry whose senescing leaves again provide the aphids with a rich source of food. The growth efficiency of R. padi on bird cherry and oats indicates that the quality of the available food on the two hosts is markedly different.
Apterous exules prefer oats to young bird cherry leaves and survive best on oats. Gynoparae prefer bird cherry leaves and only successfully produce offspring on the primary host. This, and other work published on R. padi , support the premise that a change in the aphid's preferences causes the change of host.
The dual-discrimination theory of host alternation in aphids is examined in the light of this and other published work.     

Triticum monococcum lines with distinct metabolic phenotypes and phloem‐based partial resistance to the bird cherry–oat aphid Rhopalosiphum padi

Crop protection is an integral part of establishing food security, by protecting the yield potential of crops. Cereal aphids cause yield losses by direct damage and transmission of viruses. Some wild relatives of wheat show resistance to aphids but the mechanisms remain unresolved. In order to elucidate the location of the partial resistance to the bird cherry–oat aphid, Rhopalosiphum padi, in diploid wheat lines of Triticum monococcum, we conducted aphid performance studies using developmental bioassays and electrical penetration graphs, as well as metabolic profiling of partially resistant and susceptible lines. This demonstrated that the partial resistance is related to a delayed effect on the reproduction and development of R. padi. The observed partial resistance is phloem based and is shown by an increase in number of probes before the first phloem ingestion, a higher number and duration of salivation events without subsequent phloem feeding and a shorter time spent phloem feeding on plants with reduced susceptibility. Clear metabolic phenotypes separate partially resistant and susceptible lines, with the former having lower levels of the majority of primary metabolites, including total carbohydrates. A number of compounds were identified as being at different levels in the susceptible and partially resistant lines, with asparagine, octopamine and glycine betaine elevated in less susceptible lines without aphid infestation. In addition, two of those, asparagine and octopamine, as well as threonine, glutamine, succinate, trehalose, glycerol, guanosine and choline increased in response to infestation, accumulating in plant tissue localised close to aphid feeding after 24 h. There was no clear evidence of systemic plant response to aphid infestation.     

The price of protection:a defensive endosymbiont impairs nymph growth in the bird cherry-oat aphid,Rhopalosiphum padi

Bacterial endosymbionts have enabled aphids to adapt to a range of stressors,but their effects in many aphid species remain to be established.The bird cherry-oat aphid,Rhopalosiphum padi(Linnaeus),is an important pest of cereals worldwide and has been reported to form symbiotic associations with Serratia symbiotica and Sitobion miscanthi L-type symbiont endobacteria,although the resulting aphid phenotype has not been described.This study presents the first report of R.padi infection with the facultative bacterial endosymbiont Hamiltonella defensa.Individuals of R.padi were sampled from populations in Eastern Scotland,UK,and shown to represent seven R.padi genotypes based on the size of polymorphic microsatellite markers;two of these genotypes harbored H.defensa.In parasitism assays,survival of H.defensa-infected nymphs following attack by the parasitoid wasp Aphidius colemani(Viereck)was 5 fold higher than for uninfected nymphs.Aphid genotype was a major determinant of aphid performance on two Hordeum species,a modern cultivar of barley H.vulgare and a wild relative H.spontaneum,although aphids infected with H.defensa showed 16%lower nymph mass gain on the partially resistant wild relative compared with uninfected individuals.These findings suggest that deploying resistance traits in barley will favor the fittest R.padi genotypes,but symbiontinfected individuals will be favored when parasitoids are abundant,although these aphids will not achieve optimal performance on a poor quality host plant.     

Chemical aspects of host‐acceptance behaviour in the bird cherry‐oat aphid Rhopalosiphum padi: host‐acceptance signals used by different morphs with the same genotype

Host acceptance by gynoparae and winged virginoparae of the bird cherry‐oat aphid Rhopalosiphum padi (L.) is investigated utilizing leaves and aqueous extracts of the primary and secondary hosts, as well as nonhost plants. Gynoparae are specialized to reproduce on bird cherry Prunus padus L., whereas virginoparae reproduce and feed on various grasses. Host acceptance is assessed using levels of reproduction and survival for adults, as well as survival for nymphs. Little is known of host acceptance by nymphs. The data show that gynoparae and winged virginoparae produce nymphs almost exclusively on their host plants, bird cherry and barley leaves, respectively, over 72 h. When tested with aqueous plant extracts, however, gynoparae produce nymphs almost exclusively on bird cherry extract and progeny numbers are found to be similar to those on plant leaves. Few nymphs are produced on artificial diet. By contrast, winged virginoparae produce nymphs on aqueous extracts of barley, bird cherry and bean, as well as on artificial diet. The numbers of nymphs deposited by gynoparae are similar on aqueous extracts of bird cherry leaves collected at different times during the growing season. When extracts from leaves of various Prunus species are tested, only leaves of P. padus and Prunus virginiana stimulate parturition. Oviparae, the sexual female nymphs of gynoparae, survive well for 96 h on both bird cherry and barley leaves but not on bean seedlings, whereas nymphs of winged virginoparae survive well only on barley leaves. They do not survive for 96 h on any plant‐leaf extracts, although they do survive on artificial diet.     

The role of sinigrin in host plant recognition by aphids during initial plant penetration

Plant penetration behaviour (probing) of the cabbage aphid, Brevicoryne brassicae, and the pea aphid, Acyrthosiphon pisum, was studied on excised leaves of broad beans, Vicia faba, kept in water or in a 1% aqueous solution of sinigrin. Using the DC EPG (Electrical Penetration Graph) technique it was shown that the cabbage aphid on sinigrin-untreated bean leaves showed numerous short probes into epidermis and mesophyll. None of these aphids showed either phloem salivation or ingestion waveforms on untreated leaves. In contrast, on sinigrin-treated bean leaves, 35% of the probing time was spent on phloem sap ingestion (E2) and almost all aphids reached phloem vessels and started feeding. The duration of phloem salivation before phloem ingestion and the mean duration of phloem ingestion periods were similar on a host and a sinigrin-treated non-host plant. However, the total probing time by B. brassicae was 10% longer, the total phloem sap ingestion time was twice as long, and the time to the first phloem phase within a probe was three times shorter on the host plant compared to sinigrin-treated broad beans. Acyrthosiphon pisum also responded to the addition of sinigrin to broad beans, but in this case sinigrin acted as a deterrent. On sinigrin-treated leaves, A. pisum terminated probes before ingestion from phloem vessels, and none of these aphids showed phloem salivation and ingestion on treated leaves. Glucosinolates were detected in the mesophyll cells of the brassicaceous plant, Sinapis alba. Based on this finding and in addition to the foregoing EPG analysis of aphid probing on these plants and broad beans, our hypothesis is that aphids may recognise their host plants as soon as they probe the mesophyll tissue and before they start ingestion from phloem vessels.     

Diurnal feeding as a potential mechanism of osmoregulation in aphids

Diurnal variation in phloem sap composition has a strong infuence on aphid performance.The sugar-rich phloem sap serves as the sole diet for aphids and a suite of physiological mechanisms and behaviors allowv them to tolerate the high osmotic stress.Here,we tested the hypothesis that night-time feeding by aphids is a behavior that takes advantage of the low sugar diet in the night to compensate for osmotic stress incurred while feeding on high sugar diet during the day.Using the electrical penetration graph(EPG)technique.we examined the eiects of diurmal rhythm on feeding behaviors of bird cherry-oat aphid(Rhopalosiphurm padi L.)on wheat.A strong diurmal rhythm in aphids as indicated by the presence of a cyclical pattern of expression in a core clock gene did not impact aphid feeding and similar feeding behaviors were observed during day and night.The major difference observed between day and night feeding was that aphids spent significantly longer time in phloem salivation during the night compared to the day.In contrast,aphid hydration was reduced at the end of the day-time feeding compared to end of the night-time fepding.Gene expression analysis of R.padi osmoregulatory genes indicated that sugar break down and water transport into the aphid gut was reduced at night.These data suggest that while diumal variation occurs in phloem sap composition,aphids use night time feeding to overcome the high osmotic stress incurred while feeding on sugar-rich phloem sap during the day.     

Feeding responses of two grain aphids to barley yellow dwarf virus-infected oats

The probing behavior of greenbugs, Schizaphis graminum (Rondani), and oat-bird cherry aphids, Rhopalosiphum padi (L.), was electronically monitored on healthy oats, Avena sativa (L.), and on oats infected with the RPV-NY isolate of barley yellow dwarf virus (BYDV). S. graminum fed better on infected oats than on healthy oats. This was manifested by a shorter time before initiation of committed phloem ingestion, and by increased duration of ingestion from phloem of infected compared to healthy plants. In addition, on infected oats S. graminum made fewer interruptions in their probing once their stylets were inserted into tissues, including phloem. R. padi fed similarly on infected and healthy oats, except that these aphids made fewer short probes to the phloem (lasting <15 min) on infected compared to healthy oats. BYDV infection of oats increased the rate of population growth of S. graminum relative to that on healthy oats but had no effect on the population growth of R. padi. The proportion of aphids of R. padi which developed into alatae on BYDV-infected oats was significantly greater than on healthy oats, but S. graminum showed no such response.

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Résumé Le comportement de sondage de S. graminum (Rond.) et de R. padi (L.) a été suivi électroniquement sur avoine (Avena sativa L.) saine ou contaminée par l'extrait RPV-NY du virus jaune du nanisme de l'orge (BYDV). S. graminum s'est mieux alimenté sur avoine contaminée que saine; ceci se traduisait par un temps de latence inférieur avant l'ingestion de phloème et par une prolongation de la période d'ingestion. De plus, sur avoine contaminée, S. graminum a moins souvent interrompu le sondage, une fois que les stylets avaient étè insérés dans les tissus, y compris le phloème. Les résultats avec R. padi présentaient les mêmes tendances, mais les améliorations des performances alimentaires de cette espèce sur avoine contaminée ont été moins nettes que pour S. graminum. La contamination de l'avoine par BYDV a accru le taux de croissance de la population de S. graminum par rapport à ce qui a été observé sur avoine saine; l'effet était moins prononcé pour R. padi. La proportion de pucerons devenant aliés était plus élevée sur avoine contaminée que saine, rien de tel n'a été observé avec S. graminum.

     

Effects of invasive European bird cherry (Prunus padus) on leaf litter processing by aquatic invertebrate shredder communities in urban Alaskan streams

European bird cherry (Prunus padus) (EBC) is an invasive ornamental tree that is spreading rapidly in riparian forests of urban Alaska. To determine how the spread of EBC affects leaf litter processing by aquatic invertebrate shredders, we conducted complementary leaf pack experiments in two streams located in Anchorage, Alaska. The first experiment contrasted invasive EBC with three native tree species—thin-leaf alder (Alnus tenuifolia), paper birch (Betula neoalaskana), and black cottonwood (Populus trichocarpa)—in one reach of Chester Creek; finding that EBC leaf litter broke down significantly faster than birch and cottonwood, but at a similar rate to alder. The second experiment contrasted EBC with alder in four reaches of Campbell and Chester creeks; finding that while EBC leaf litter broke down significantly faster than alder in Chester Creek, EBC broke down at a similar rate to alder in Campbell Creek. Although EBC sometimes supported fewer shredders by both count and mass, shredder communities did not differ significantly between EBC and native plants. Collectively, these data suggest that invasive EBC is not currently exhibiting strong negative impacts on leaf litter processing in these streams, but could if it continues to spread and further displaces native species over time.     

Change in response of Rhopalosiphum padi spring migrants to the repellent winter host component methyl salicylate

Olfactometry showed that the response of spring migrants of the bird cherry-oat aphid, Rhopalosiphum padi (L.) (Homoptera: Aphididae), to the repellent winter host volatile methyl salicylate changes with age of the adult aphid. Between three and four days after becoming adult, and having left the winter host Prunus padus L., aphids lost their negative response to the chemical. The change in response was not associated with contact with a summer host, oats. In a settling choice bioassay, migrants avoided oats which had been exposed to volatile methyl salicylate. Aphids with removed antennal tips did not avoid the exposed plant, indicating that plant choice was influenced by cues from the plant surface. The results are discussed in relation to the use of methyl salicylate in integrated control.     

Development of the Potential for Cyanogenesis in Maturing Black Cherry (Prunus serotina Ehrh.) Fruits

Biochemical changes related to cyanogenesis (hydrogen cyanide production) were monitored during maturation of black cherry (Prunus serotina Ehrh.) fruits. At weekly intervals from flowering until maturity, fruits (or selected parts thereof) were analyzed for (a) fresh and dry weights, (b) prunasin and amygdalin levels, and (c) levels of the catabolic enzymes amygdalin hydrolase, prunasin hydrolase, and mandelonitrile lyase. During phase I (0-28 days after flowering [DAF]), immature fruits accumulated prunasin (mean: 3 micromoles/fruit) but were acyanogenic because they lacked the above enzymes. Concomitant with cotyledon development during mid-phase II, the seeds began accumulating both amygdalin (mean: 3 micromoles/seed) and the catabolic enzymes and were highly cyanogenic upon tissue disruption. Meanwhile, prunasin levels rapidly declined and were negligible by maturity. During phases II (29-65 DAF) and III (66-81 DAF), the pericarp also accumulated amygdalin, whereas its prunasin content declined toward maturity. Lacking the catabolic enzymes, the pericarp remained acyanogenic throughout all developmental stages.     

Induced proteolysis within the bird cherry leaves evoked by Rhopalosiphum padi L. (Hemiptera, Aphidoidea)

The objectives of this study were to elucidate the impact of bird cherry-oat aphid (Rhopalosiphum padi L.) feeding on functioning of the proteolytic machinery in bird cherry leaves. Biochemical analyses proved that R. padi feeding in tissues of primary host stimulated activity of the two major fractions of proteinases (extracted at the optimal pH values: 5.0 and 7.0). Additionally, it has been demonstrated that aphids' feeding on bird cherry led to a decline in levels of albumins and globulins (main protein fractions in P. padus leaves). The opposite tendency, regarding the amounts of these protein fractions was ascertained at the phase of disappearance of R. padi population on tested shoots. Furthermore, it is reported that an increase in activity of the analysed enzymes and a decline in the content of tested protein fractions, were proportional to density of aphid individuals developing on P. padus side shoots. It is hypothesized that long-term R. padi feeding may lead to intensifying the catabolic processing of proteins by the activated proteolytic machinery in bird cherry leaves. The multi-level biological functions of endogenous plant proteinases and their significance in triggering the defense reactions in aphid-infested plant tissues are discussed.     

Winter Host Plant Specialization in a Host-Alternating Aphid

The host alternating aphid, Rhopalosiphum padi (L.), feeds in summer on several different species of grasses but is monophagous on its winter host, Prunus padus L. The monophagy on P. padus could be a result of the restricted host range of the several different generations colonizing, or feeding, on this host during autumn–winter–spring. This study shows that the winter host plant specificity of R. padi is controlled mainly by the preference of the females remigrating (gynoparae) to the winter host, P. padus, in autumn. The other generations living on the winter host, i.e., sexual females, males, and spring generations, all accept a broader range of winter hosts. One alternative host plant, Prunus spinosa L., could be utilized by all generations associated with the winter host, except for the females remigrating at autumn.