Neem seed oil inhibits aphid transmission of potato virus Y to pepper*

Transmission of potato vims Y to sweet pepper by the green peach aphid, Myzus persicae (Sulzer), was inhibited by foliar applications of 1.0% or 2.0% neem seed oil to infected source plants or to uninfected recipient plants. Neem seed oil interfered with virus acquisition and inoculation in a manner comparable to that of a commercial horticultural oil, while an oil-free neem seed extract did not reduce rates of transmission compared with controls. The finding that neem seed oil inhibits virus transmission, while oil-free neem seed extract does not, suggests that the presence of the oil rather than biologically active limonoids such as azadirachtin interfere with virus transmission. None of the treatments affected rates of infection when potato virus Y was transmitted mechanically, or the resulting virus titre and symptom expression. In addition to direct control of insect pests, formulated neem oils may help reduce or delay the spread of non-persistent plant viruses.     

The role of flying aphid vectors in the transmission of cucumber mosaic virus and potato virus Y to peppers in Israel

More than 44 species of aphids were trapped by suction during the spring seasons of 1981, 1982 and 1983 over a pepper field at Bet Dagan, Israel. Nineteen species transmitted cucumber mosaic virus (CMV), while seven transmitted potato virus Y (PVY) at least once. Over 80% of the CMV and of the PVY infection among test plants (Capsicum annuum cv. Weindale) exposed to trapped aphids was caused by Aphis citricola and two or three other Aphis species, Myzus persicae and Macrosiphum euphorbiae. Landing rate was determined by comparing the proportion of each species found on green tiles or pepper plants with that found in suction traps. A. citricola was the most common but was found in a much lower proportion on plants than either in flight or on green tiles. Aphis spp. and M. persicae were more than 2–5 times more frequent (relative to other species) on green tiles than in flight. M. persicae and M. euphorbiae, which colonise peppers, were found on peppers at a proportion several times higher than either on green tiles or in the air. The relative importance of the different vector species was calculated by multiplying abundance by the proportion of transmitters and the landing rate. A. citricola and Aphis spp. were responsible for more than 50% of the total transmission of either CMV in 1981 and 1982 and of PVY in 1981. Peaks of CMV infection of bait plants coincided with peaks of transmitters of A. citricola and Aphis spp. caught in suction traps. The significance of these findings in primary infection of peppers with CMV and PVY is discussed.     

Morph determination in the bird cherry-oat aphid, Rhopalosiphum padi L

Both crowding and poor nutrition induce the appearance of emigrants in Rhopalosiphum padi L. No emigrants developed when the aphid was reared in isolation for six successive generations on actively growing leaves of bird cherry. However, emigrants developed as soon as the leaves ceased to grow or when the aphids became crowded. Crowding of mothers and postnatal crowding of the nymphs of apterous exules both influenced the induction of alate exules. The highest proportion of alate exules developed when both mother and offspring were crowded. Short day-length induced the appearance of gynoparae and males. For 50% production of gynoparae and males, over the range of temperature 10–18d?C, a 1.75 h reduction in day-length is required for every 4d? increase in temperature. At 18 d?C, even at day-lengths as short as 6 h, exposure for three generations is required before all offspring become gynoparae or males. Low temperature and short day-length was ineffective in inducing the development of gynoparae or males in the first three generations developing from the fundatrix.     

Mapping resistance to the bird cherry-oat aphid and the greenbug in wheat using sequence-based genotyping

Key message

Identification of novel resistance QTL against wheat aphids. First QTL-resistance report for R. padi in wheat and chromosome 2DL for S. graminum . These sources have potential use in wheat breeding.

Abstract

The aphids Rhopalosiphum padi and Schizaphis graminum are important pests of common wheat (Triticum aestivum L.). Characterization of the genetic bases of resistance sources is crucial to facilitate the development of resistant wheat cultivars to these insects. We examined 140 recombinant inbred lines (RILs) from the cross of Seri M82 wheat (susceptible) with the synthetic hexaploid wheat CWI76364 (resistant). RILs were phenotyped for R. padi antibiosis and tolerance traits. Phenotyping of S. graminum resistance was based on leaf chlorosis in a greenhouse screening and the number of S. graminum/tiller in the field. RILs were also scored for pubescence. Using a sequence-based genotyping method, we located genomic regions associated with these resistance traits. A quantitative trait locus (QTL) for R. padi antibiosis (QRp.slu.4BL) that explained 10.2 % of phenotypic variation was found in chromosome 4BL and located 14.6 cM apart from the pubescence locus. We found no association between plant pubescence and the resistance traits. We found two QTLs for R. padi tolerance (QRp.slu.5AL and QRp.slu.5BL) in chromosomes 5AL and 5BL, with an epistatic interaction between a locus in chromosome 3AL (EnQRp.slu.5AL) and QRp.slu.5AL. These genomic regions explained about 35 % of the phenotypic variation. We re-mapped a previously reported gene for S. graminum resistance (putatively Gba) in 7DL and found a novel QTL associated with the number of aphids/tiller (QGb.slu-2DL) in chromosome 2DL. This is the first report on the genetic mapping of R. padi resistance in wheat and the first report where chromosome 2DL is shown to be associated with S. graminum resistance.     

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.     

Secondary host preferences and reproductive activity of the bird cherry-oat aphid, Rhopalosiphum padi

Of 15 species of grass, including cereals, alate exules and emigrants of Rhopalosiphum padi preferred to colonise Lolium perenne. Emigrants colonised cereals twice as readily as alate exules. Apterous exules also preferred L. perenne to cereals. Although small when reared on L. perenne, R. padi was more fecund per unit weight than on any of the other grasses tested. On the later growth stages of Dactylis glomerata and L. perenne, R. padi developed faster and were more fecund than on Phleum pratense at the same growth stages. At low temperatures R. padi were more fecund on P. pratense than on L. perenne and D. glomerata and vice versa at high temperatures. The secondary host preferences and reproductive activity of R. padi are discussed in relation to possible suitable hosts for viviparous overwintering and the pest status of this aphid in Britain.     

Resistance to potato virus Y and potato virus X in Solanurn brevidens

Although Solanum brevidens could be infected with potato virus X (PVX), potato virus Y⁰ (PVY⁰) and PVY^N, no symptoms of infection were apparent and tests by double antibody sandwich ELISA, electron microscopy and sap transmission to local lesion test plants indicated that the titres of PVX were less than a tenth of those of PVY⁰ and PVY^N were less than a hundredth of those in infected plants of PDH40, a susceptible dihaploid clone of S. tuberosum cv. Pentland Crown. Furthermore, PVY⁰- and PVY^N- infected leaves of S. brevidens were a poor source of inoculum in aphid transmission tests. The possibility of a common mechanism and genetic basis of resistance to PVY, PVX and potato leaf roll virus in S. brevidens is discussed.     

Operating characteristics of full count and binomial sampling plans for green peach aphid (Hemiptera: Aphididae) in potato

Counts of green peach aphid, Myzus persicae (Sulzer) (Hemiptera: Aphididae), in potato, Solanum tuberosum L., fields were used to evaluate the performance of the sampling plan from a pest management company. The counts were further used to develop a binomial sampling method, and both full count and binomial plans were evaluated using operating characteristic curves. Taylor's power law provided a good fit of the data (r2 = 0.95), with the relationship between the variance (s2) and mean (m) as ln(s2) = 1.81(+/- 0.02) + 1.55(+/- 0.01) ln(m). A binomial sampling method was developed using the empirical model ln(m) = c + dln(-ln(1 - P(T))), to which the data fit well for tally numbers (T) of 0, 1, 3, 5, 7, and 10. Although T = 3 was considered the most reasonable given its operating characteristics and presumed ease of classification above or below critical densities (i.e., action thresholds) of one and 10 M. persicae per leaf, the full count method is shown to be superior. The mean number of sample sites per field visit by the pest management company was 42 +/- 19, with more than one-half (54%) of the field visits involving sampling 31-50 sample sites, which was acceptable in the context of operating characteristic curves for a critical density of 10 M. persicae per leaf. Based on operating characteristics, actual sample sizes used by the pest management company can be reduced by at least 50%, on average, for a critical density of 10 M. persicae per leaf. For a critical density of one M. persicae per leaf used to avert the spread of potato leaf roll virus, sample sizes from 50 to 100 were considered more suitable.     

Antibiosis and antixenosis of six commonly produced potato cultivars to the green peach aphid, Myzus persicae Sulzer (Hemiptera: Aphididae)

The antibiotic and antixenotic resistance of six commonly produced potato cultivars in Iran including Aozonia, Agria, Cosima, Cosmos, Kondor and Savalan to the green peach aphid, Myzus persicae Sulzer, were investigated under laboratory conditions at 20 ± 2oC, 65 ± 5% RH and 16:8h (L:D) in 2009. Antibiosis experiments showed significant differences in the developmental time, nymphal survivorship, fecundity, adult longevity of the green peach aphid among the potato cultivars. Intrinsic rate of natural increase (r m) for apterous aphids varied significantly with the potato cultivars on which the aphids were reared. This value ranged from 0.225 to 0.293 females/female/day, which was lowest on Cosmos and highest on Aozonia. Additionally, the estimated net reproductive rate (R0) and finite rate of increase (λ) for apterous aphids were the lowest on Cosmos. For the antixenosis experiment, no significant difference was found in aphid's preference to the potato cultivars. However, Aozonia was preferred more than the other five cultivars by the apterous aphids. Therefore, our results demonstrated that among the investigated cultivars the Cosmos cultivar is moderately resistant to the green peach aphid.     

Ovariole number and fecundity for the two emigrating generations of the bird cherry-oat aphid (Rhopalosiphum padi) in Sweden

Abstract. 1. The number of ovarioles and the potential fecundity in first (fundatrigenia) and second generation emigrants of Rhopalosiphum padi (L.) was investigated.
2. The relationships between weight and host quality on ovariole number were investigated.
3. The first generation of emigrants had a greater number of ovarioles and a higher fecundity than the second.
4. There was a positive relationship between ovariole number and weight.
5. The number of ovarioles and weight decreased with decreasing host quality.
6. The results do not agree with a recently proposed reproductive strategy of aphids where the number of ovarioles are supposed to be independent of weight and host quality.
7. In years with a high initial population on bird cherry, the migration to grasses, e.g. cereals, will be earlier, probably larger and, to a greater extent, consists of aphids which have a higher reproductive rate than in years with a small initial number on bird cherry.     

Varietal differences in susceptibility to potato virus Y

In addition to giving different kinds of symptoms when infected with potato virus Y , individual potato varieties also differ in their susceptibility to infection, in the concentration of virus attained in their sap, and in their efficiency as sources of virus for aphides. Their relative susceptibility in the open when exposed to equal chances of infection is correlated with the ease with which they become infected when colonized with infective aphides, and can be assessed from tests made under glass. Methods for making such tests are described; these need few tubers and give reproducible results. It is considered that they could be applied in studying the inheritance of this type of resistance and to test the behaviour of new seedlings. The American variety Katahdin was the most resistant of those tested, but there were significant differences between commercial British varieties.
In the open, all varieties were equally colonized by aphides and resistance to infection with virus Y was not correlated with resistance to leaf roll.     

The transmission of potato virus Y by aphids of different vectoring abilities

Adult apterae of Myzus persicae and Macrosiphum euphorbiae that did not transmit potato virus Y^N (PVY^N) in a first test were as likely to transmit the virus in a subsequent test as those that did transmit on the first occasion. Only 16% of M. persicae that were allowed a single acquisition probe into a leaf infected with both PVY^O and PVY^N transmitted both strains, 37% transmitted either PVY^O or PVY^N and 47% did not transmit. There was no difference in the duration of probes that did or did not result in virus transmission. Statistical models were fitted to data on the frequency of transmission of PVY^O, PVY^N or both PVY^O and PVY^N by M. persicae and by aphids of poorer vector species, M. euphorbiae and Rhopalosiphum padi. Transmission of the two viruses ocurred independently of each other and consequently transmission of both was rare with M. euphorbiae and R. padi. Mineral oil applied to leaves infected with both strains diminished the frequency of transmission by M. persicae. Fitted models suggested that the aphids that probed through the oil droplets on leaves treated 30 min previously did not transmit virus, and that 24 h later, when the droplets had spread, aphids probing through them could transmit but with a decreased ability.     

Entomotoxic action of jackbean lectin (Con A) in bird cherry-oat aphid through the effect on insect enzymes

The mechanism of the toxicity of plant lectins is not clearly understood. The insecticidal activity of plant lectins results from effects on insect metabolism by interfering with gut enzymatic function. Thus, a greater understanding of the mechanisms of plant lectin toxicity in insects is required. This study reports the effects of dietary ingestion of the glucose-mannose binding lectin Concanavaline A (Con A) on bird cherry-oat aphid (Rhopalosiphum padi L.) enzymes involved in protein digestion [aminopeptidase N and cathepsin L (CatL)], sugar (α- and β-glucosidases), and phosphorus (alkaline and acid phosphatase) metabolism. An aphid bioassay test using artificial diets containing Con A is also presented. An increase in the concentration of Con A generally suppressed the activity of glucosidases and phosphatases, and increased the activity of CatL in apterae morphs. Bird cherry-oat aphid performance was affected by the presence of Con A in artificial diets. The lectin added to the liquid diet increased the pre-reproductive period, mortality, and the average time of generation development (T) and decreased fecundity and the intrinsic rate of natural increase (r_m). Aphicidal activity of Con A might be linked to its interference in the activity of digestive enzymes.     

Identifying bird cherry-oat aphid Rhopalosiphum padi emigrants, alate exules and gynoparae: application of multivariate methods to morphometric and anatomical features

The host microhabitat location behavior of females of the generalist parasitoid Campoletis sonorensis (Cameron) (Hymenoptera: Ichneumonidae) was studied in a wind tunnel. Visual cues associated with the host plant cotton, Gossypium hirsutum L., were important and significantly more parasitoids completed flights to a damaged 4-leaf cotton plant bearing a Heliothis virescens (F.) (Lepidoptera: Noctuidae) larva and frass than to a similarly damaged single leaf with frass and a larva. This difference in completed flights was not due to differences in amounts of volatiles released by the two stimuli. Both naive and experienced parasitoids responded differently to an undamaged cotton leaf, a mechanially damaged leaf, a naturally damaged leaf with the host removed and a naturally damaged leaf with a host larva. Parasitoids completed significantly fewer flights to the undamaged sources of volatiles than to damaged sources of volatiles. Experienced females responded strongly to all types of damage. The number of flights completed by naive females to the three types of damage differed but not significantly and was less than the number completed by experienced females. Components of the preflight experience were varied to determine which factors were responsible for the higher response of experienced females to the host/plant complex. Oviposition was the most important component of this experience. Contact with host frass or plant damage followed by oviposition did not increase the response over that exhibited by females allowed oviposition only. When frass or damaged plant material were contacted without subsequent oviposition, females completed fewer flights than naive females.     

Costs and benefits of hydroxamic acids-related resistance in winter wheat against the bird cherry-oat aphid, Rhopalosiphum padi L.

Estimations of infestation by the bird cherry-oat aphid (Rhopalosiphum padi) as well as measurements of grain yield in 26 Hungarian winter wheat cultivars under field conditions were correlated with the concentration of hydroxamic acids (Hx) in seedlings of those cultivars. The significant inverse relationship between infestation ratings and Hx levels in wheat showed that Hx, despite their decreased accumulation at later plant phenological stages, may be able to confer resistance against aphid infestation in the field. Since no significant relationship was found between grain yield and Hx levels in plants it is suggested that Hx accumulation does not impose a cost to the plant in terms of yield. These findings support earlier claims stressing the potential of Hx as breeding targets for aphid resistance in wheat.     

Fixed precision sequential sampling plans for the greenbug and bird cherry-oat aphid (Homoptera: Aphididae) in winter wheat

The numbers of greenbugs, Schizaphis graminum (Rondani), and bird cherry-oat aphids, Rhopalosiphum padi L., per wheat tiller (stem) were estimated in 189 production winter wheat (Triticum aestivum L.) fields located throughout Oklahoma. Taylor's power law regressions were calculated from these data and used to construct fixed precision sequential sampling schemes for each species. An evaluation data set was constructed from 240 samples taken during three growing seasons from winter wheat fields at four locations in Oklahoma. Wheat cultivar and growth stage were recorded for each field on the day of sampling. Taylor's power law parameters for evaluation fields differed significantly for both species among growing seasons, locations, and plant growth stages. Median precision achieved using the fixed precision sequential sampling schemes for each species departed <20% from expected precision over the range population intensity in the evaluation data. For the 10% of samples with greatest deviation between observed and expected precision, observed precision was 13.8-81.8% greater than that expected precision depending on aphid species and population intensity. For the greenbug, the distribution of the percentage deviation between observed and expected precision was positively skewed, so that the sampling scheme tended to over-predict precision. For the bird cherry-oat aphid, the distribution was more symmetric. Even though precision observed using the sampling schemes frequently varied from expected precision, because of the inevitable consequence of sampling error and environmental variation, the sampling schemes yielded median observed precision levels close to expected precision levels over a broad range of population intensity.     

Discrimination of Arabidopsis PAD4 activities in defense against green peach aphid and pathogens

The Arabidopsis (Arabidopsis thaliana) lipase-like protein PHYTOALEXIN DEFICIENT4 (PAD4) is essential for defense against green peach aphid (GPA; Myzus persicae) and the pathogens Pseudomonas syringae and Hyaloperonospora arabidopsidis. In basal resistance to virulent strains of P. syringae and H. arabidopsidis, PAD4 functions together with its interacting partner ENHANCED DISEASE SUSCEPTIBILITY1 (EDS1) to promote salicylic acid (SA)-dependent and SA-independent defenses. By contrast, dissociated forms of PAD4 and EDS1 signal effector-triggered immunity to avirulent strains of these pathogens. PAD4-controlled defense against GPA requires neither EDS1 nor SA. Here, we show that resistance to GPA is unaltered in an eds1 salicylic acid induction deficient2 (sid2) double mutant, indicating that redundancy between EDS1 and SID2-dependent SA, previously reported for effector-triggered immunity conditioned by certain nucleotide-binding-leucine-rich repeat receptors, does not explain the dispensability of EDS1 and SID2 in defense against GPA. Mutation of a conserved serine (S118) in the predicted lipase catalytic triad of PAD4 abolished PAD4-conditioned antibiosis and deterrence against GPA feeding, but S118 was dispensable for deterring GPA settling and promoting senescence in GPA-infested plants as well as for pathogen resistance. These results highlight distinct molecular activities of PAD4 determining particular aspects of defense against aphids and pathogens.     

Characterization of a new densovirus infecting the green peach aphid Myzus persicae

A new icosahedral DNA virus was isolated from aphids (Myzus persicae) that showed abnormal growth and development. The purified virus particles have a diameter of 20 nm and contain a single-stranded DNA molecule of approximately 5.7 kb. The viral particles are composed of five structural proteins (92, 85, 68, 64, and 57 kDa). As the main biophysical properties of this virus are similar to those of the members of the genus Densovirus it was tentatively named Myzus persicae densovirus (MpDNV). A PCR-based detection method and a polyclonal antiserum raised against MpDNV allowed the detection of the virus in a single-infected aphid. MpDNV is immunologically related to Junonia coenia densovirus, but not to other members of the subfamily Densovirinae. Biological assays showed that MpDNV could be both transmitted transovarially and horizontally via honeydew and saliva. MpDNV was able to infect whiteflies but not other aphid species tested.     

Expression of genes for resistance to potato virus Y in potato plants and protoplasts

Plants of several potato clones with major gene resistance to potato virus Y (PVY) developed necrotic local lesions and systemic necrosis after manual inoculation with common (PVY_o) or veinal necrosis (PVY_N) strains of the virus. The clones reacted similarly, although their resistance genes are thought to be derived from four different wild species of Solarium. Mesophyll protoplasts from each clone became infected when inoculated with RNA of PVY_o by the polyethylene glycol method. The proportion of protoplasts infected, assessed by staining with fluorescent antibody to virus particles, was similar to that of protoplasts of susceptible potato cultivars. In contrast, plants of potato cultivars Corine and Pirola, which possess gene Ry from S. stoloniferum, developed few or no symptoms when manually inoculated or grafted with PVY_o. Moreover, only very few protoplasts of these cultivars produced virus particle antigen after inoculation with PVY_o RNA. The extreme resistance to PVY of cvs Corine and Pirola was therefore expressed by inoculated protoplasts whereas the resistance of the necrotic-reacting potato clones was not.