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Potential effects of plant protease inhibitors, oryzacystatin I and soybean Bowman-Birk inhibitor, on the aphid parasitoid Aphidius ervi Haliday (Hymenoptera, Braconidae).

Protease inhibitors (PIs) have been shown to cause lethal and sublethal effects on aphids depending on the kind of PI and aphid species. Therefore, these proteins might affect aphid parasitoids directly by inhibiting their digestive proteolysis or indirectly via their development in a less suitable host. In our study, the risk of exposure and the potential effects of soybean Bowman-Birk inhibitor (SbBBI) and oryzacystatin I (OCI) on the aphid endoparasitoid Aphidius ervi were investigated using artificial diet to deliver PIs. Immunoassays showed that both SbBBI and OCI were detected in the honeydew of aphids reared on artificial diet containing these recombinant proteins at 100 microg/mL. However, only SbBBI was detected in parasitoid larvae, while this PI could not be detected in adult parasitoids emerged from PI-intoxicated aphids. Enzymatic inhibition assays showed that digestive proteolytic activity of larvae and adults of A. ervi predominantly relies on serine proteases and especially on chymotrypsin-like activity. Bioassays using SbBBI and OCI on artificial diet were performed. A. ervi that developed on intoxicated aphids had impaired fitness. Thus development and parasitism success of parasitoids exposed to OCI were severely affected. On the contrary, SbBBI only altered significantly female size and sex ratio. Direct exposure to PIs through adult food intake did not affect female's longevity, while SbBBI and OCI (100 microg/mL) induced 69% and 30% inhibition of digestive protease activity, respectively. These studies made it possible to estimate the risk of exposure to plant PIs and the sensitivity of the aphid parasitoid A. ervi to these entomotoxins, by combining immunological, biochemical and biological approaches. First it pointed out that only immature stages are affected by PIs. Secondly, it documented two different modes of effect, according to the nature of the PIs and both host and parasitoid susceptibility. OCI prevented the development of A. ervi mainly due to the host susceptibility, whereas SbBBI only induced sublethal effects on the parasitoid, possibly due to both direct action on the parasitoid susceptible proteases, and host-mediated action through size reduction.

Animals↗

Electron microscopy studies on luteovirid transmission by aphids.

Transmission electron microscopy (TEM) observations have been extensively applied to follow the route of luteovirids in their vectors. Luteovirids are icosahedral plant viruses which are phloem-limited and strictly transmitted in a circulative manner by aphids. Virus particles, acquired by aphids while feeding on an infected plant, circulate in the aphid's body without replication and are internalized during this process in two different cell types (intestinal and accessory salivary gland cells). The endocytosis mechanism at the gut level seems to rely on a clathrin-mediated entry process and virions are observed in the aphid's gut cells in various vesicular structures. After exocytosis from intestinal cells, virions are released in the aphid's body cavity where they are thought to bind to symbionin, an endosymbiotic protein. Transcytosis of the accessory salivary gland cells occurs similarly as at the gut level but in the reverse direction. Using engineered virus mutants, viral proteins required for transmission and involved in virus retention in the hemocoel have been identified. Virus mutants poorly or non aphid-transmitted have also been localized in the aphid's body by TEM. These observations reveal the crucial role of the minor capsid protein in gut internalization. While not strictly required, this protein seems to play an important role in the efficiency of this process by interacting with putative virus receptors localized on the gut apical membrane. More recently, some aphid proteins have also been shown to exhibit in vitro virus binding capacity and could potentially be components of the endocytotic apparatus.

Animals↗

Patterns of resistance to three cereal aphids among wheats in the genus Triticum (Poaceae).

Forty-one accessions of wild and cultivated wheats belonging to 19 Triticum species were tested in the field for resistance to three species of aphids, Rhopalosiphum padi Linnaeus, Sitobion avenae Fabricius and Schizaphis graminum Rondani. Antibiotic resistance was estimated by the increase in biomass of aphids over 21 days on adult plants. Overall resistance was estimated by the plant biomass lost due to aphid infestation. All three species of aphids survived and reproduced on all wheats, and reduced spike biomass compared to uninfested controls. The level of antibiosis varied among wheat species and among accessions, with accessions from three, five and one species showing antibiosis to R. padi, S. avenae and S. graminum, respectively. Overall resistance to the three aphid species was observed in five to seven accessions per aphid species. Resistance was usually specific to one aphid species. The frequency of accessions with antibiosis or overall resistance was associated with the ploidy level of the plant species. Except for overall resistance to R. padi, resistance was highest for diploid species and lowest for hexaploid species. No consistent relationship between resistance and level of domestication was detected. Accessions of the wild wheats, Triticum boeoticum Bois, Triticum tauschii (Coss.) Schmal. and Triticum araraticum Jakubz. exhibited high levels of resistance to aphids, as did Triticum monococcum L. which is derived from T. boeoticum. Nevertheless, individual susceptible or resistant accessions occurred at all levels within the evolutionary tree of wheat.

Analysis of Variance↗

The spatial dynamics of crop and ground active predatory arthropods and their aphid prey in winter wheat.

The distribution of aphid predators within arable fields has been previously examined using pitfall traps. With this technique predominantly larger invertebrate species are captured, especially Carabidae, but the technique provides no estimate of density unless mark-recapture is used. However, many other numerically important aphid predators occur in arable fields and relatively little is known about their distribution patterns nor whether they exhibit a density-dependent response to patches of cereal aphids. Identification of the most effective predators can allow management practices to be developed accordingly. In this study, the distribution of cereal aphids and their predators was examined by suction sampling within a field of winter wheat in Devon, UK, along with visual estimates of weed patchiness. Sampling was conducted on four occasions in 1999 across a grid of 128 sample locations. The distribution of 11 predatory taxa from the Carabidae, Staphylinidae and Linyphiidae was examined. Additionally, the total number of aphid predators and a predation index were used in these analyses. Carabid adults and larvae, along with staphylinid larvae showed the strongest aggregation into patches and the most temporal stability in their distribution. Other taxa had more ephemeral distributions as did the cereal aphids. The distribution of carabid larvae was disassociated from the distribution of cereal aphids for the first two sampling occasions indicating biocontrol was occurring. Other predatory groups showed both association and disassociation. Carabid larvae, Bathyphantes and total numbers of Linyphiidae showed a strong correlation with weed cover for two of the sample dates. Cereal aphids were disassociated from weed cover on three sampling occasions.

Animals↗

The importance of antennae for pea aphid wing induction in the presence of natural enemies.

The pea aphid Acyrthosiphon pisum Harris has been shown to produce an increasing proportion of winged morphs among its offspring when exposed to natural enemies, in particular hoverfly larvae, lacewing larvae, adult and larval ladybirds and aphidiid parasitoids. While these results suggest that wing induction in the presence of predators and parasitoids is a general response of the pea aphid, the cues and mechanisms underlying this response are still unclear. Tactile stimuli and the perception of chemical signals as well as visual signals are candidates for suitable cues in the presence of natural enemies. In this paper the hypothesis that the aphids' antennae are crucial for the wing induction in the presence of natural enemies is tested. Antennae of pea aphids were ablated and morph production was scored when aphids were reared either in the presence or the absence of predatory lacewing larvae over a six-day period. Ablation of antennae resulted in a drastic drop in the proportion of winged morphs among the offspring, both in the presence and the absence of a predator whereas predator presence increased wing induction in aphids with intact antennae, as reported in previous experiments. The results show that antennae are necessary for wing induction in the presence of natural enemies. Critical re-examination of early work on the importance of aphid antennae and tactile stimuli for wing induction suggests that a combination of tactile and chemical cues is likely to be involved not only in predator-induced wing formation but also for wing induction in response to factors such as crowding in the aphid colony.

Analysis of Variance↗

Effects of hydrophilic and hydrophobic kaolin-based particle films on pea aphid (Homoptera: Aphididae) and its entomopathogen Pandora neoaphidis (Entomophthorales: Entomophthoraceae).

Hydrophobic and hydrophilic kaolin-based particle films are effective for control of insect pests in certain agricultural crops. How these products interact with potential biological control agents is not well documented. This study was conducted to evaluate the effects of the hydrophobic (M96-018) and hydrophilic (Surround WP) kaolin-based particle films (Engelhard Corporation, Iselin, NJ) on pea aphid, Acyrthosiphon pisum (Harris), on peas (Pisum spp.), and on the fungal aphid pathogen Pandora neoaphidis (Remaudière and Hennebert) Humber. Over two field seasons (2001 and 2002) in northern Idaho, applications of M96-018 significantly reduced the rate of pea aphid increase on pea, but Surround WP, tested only in 2001, did not reduce aphid population growth rate. Neither particle film treatment was as effective as a standard application of esfenvalerate (DuPont Asana). In the laboratory, particle films suppressed pea aphid populations by up to 30%. M96-018 seemed to have some repellent activity based on aphid distributions after treating plants. When applied along with P. neoaphidis conidia, M96-018 but not Surround WP caused higher percentage of infection mortality of pea aphids by P. neoaphidis than occurred on controls treated only with P. neoaphidis conidia. P. neoaphidis conidia deposited on glass slides coated with M96-018, produced more germ tubes and secondary conidia than those deposited on untreated glass slides or slides treated with Surround WP. This result suggests that greater infection of pea aphids on plants treated with M96-018 is in part a result of a direct enhancement of fungal germination by the particle film.

Analysis of Variance↗

Molecular evidence against phylogenetically distinct host races of the pea aphid (Acyrthosiphon pisum).

Pea aphid (Acyrthosiphon pisum) clones have been shown to be adapted to particular host plant species but it is unknown whether there are host races. A 1101 base pair region of the mitochondrial cytochrome oxidase I gene (COI) was sequenced for 21 pea aphid clones that had been collected from different host plants in Canada and the U.S.A. Only five closely related mitochondrial haplotypes were found. A maximum likelihood phylogeny was estimated for these five haplotypes and four related aphid species: Acyrthosiphon macrosiphum, A. kondoi, Fimbriaphis fimbriata, and Macrosiphum creelii. Pea aphids from the same host plant species were no more likely to have the same mitochondrial haplotype than aphids from different host plant species. In addition, aphids from the same geographical regions were no more likely to have the same mitochondrial haplotype than aphids from different geographic regions. I therefore reject the hypothesis that there are monophyletic host races of the pea aphid.

Animals↗

Biological control of cereal aphids in north america and mediating effects of host plant and habitat manipulations.

The predator and parasitoid fauna associated with cereal aphids is described, emphasizing the fauna associated with classical biological control efforts against the greenbug and Russian wheat aphid. We focus on literature from North America and include work from Europe and elsewhere when it is desirable to draw contrasts between approaches that affect cereal aphid biological control. Effects on natural enemies of cereal aphids are described that appear associated with plant traits that are innate, bred, or induced by aphid feeding. Examples of habitat manipulations, within and bordering cereal fields and within the broader landscape in which cereal production resides, affecting predators and parasitoids of cereal aphids are presented. These mediating effects of host plant and habitat manipulations on cereal aphid biological control provide significant and underexplored avenues to optimize cereal aphid management.

Animals↗

Amino acid composition and nutritional quality of potato leaf phloem sap for aphids.

To define plant 'nutritional quality' for aphids, the causal basis of the variation in aphid performance between host plants of different developmental ages was explored using the aphids Myzus persicae and Macrosiphum euphorbiae on potato plants (Solanum tuberosum). Both aphid species performed better on developmentally young ('pre-tuber-filling') plants than on mature ('tuber-filling') plants. Aphid performance did not vary with leaf phloem sucrose:amino acid ratio but could be related to changes in the amino acid composition of the phloem, which included a developmental shift from high glutamine levels in pre-tuber-filling plants to low glutamine levels in tuber-filling plants. Aphid performance on chemically defined 'young' and 'old' diets, with amino acid composition corresponding to that of phloem amino acid composition in pre-tuber-filling and tuber-filling plants, respectively, confirmed that phloem amino acid composition contributed to low aphid performance on tuber-filling plants. The relatively poor performance on 'old' diets could be accounted for, at least in part, by depressed feeding rates. These data suggest that amino acid composition of the phloem is one factor shaping the nutritional quality of plants for aphids.

Amino Acids↗

Evaluation of hairy nightshade as an inoculum source for aphid-mediated transmission of potato leafroll virus.

Potato leafroll virus (PLRV) causes one of the most serious aphid-transmitted diseases affecting yield and quality of potatoes, Solanum tuberosum (L.), grown in the United States. The green peach aphid, Myzus persicae (Sulzer), is considered to be by far the most efficient vector of this virus. Even the most strict aphid control strategy may not prevent the spread of PLRV unless measures also are taken to keep virus source plants within and outside the crop at a minimum. Hairy nightshade, Solanum sarrachoides (Sendtner), is one of the preferred weed hosts for green peach aphid. The potential of this weed as an aphid reservoir and virus source and its spread or perpetuation were investigated. With the use of double antibody sandwich enzyme-linked immunosorbent assay, it was confirmed that green peach aphid can transmit PLRV to hairy nightshade and that aphids can become viruliferous after feeding on infected hairy nightshade plants. Transmission from hairy nightshade to potato is 4 times the rate of potato to potato or potato to hairy nightshade. The green peach aphid preferred hairy nightshade over potato plants and reproduced at a higher rate on hairy nightshade than on potato. Therefore, a low level of PLRV-hairy nightshade infection could enhance the disease spread in the field.

Animals↗

Biotypic variation among north American Russian wheat aphid (Homoptera: Aphididae) populations.

The Russian wheat aphid, Diuraphis noxia (Mordvilko) (Homoptera: Aphididae), has been a major economic pest of small grains in the western United States since its introduction in 1986. Recently, a new Russian wheat aphid biotype was discovered in southeastern Colorado that damaged previously resistant wheat, Triticum aestivum L. Biotype development jeopardizes the durability of plant resistance, which has been a cornerstone for Russian wheat aphid management. Our objective was to assess the relative amount of biotypic diversity among Russian wheat aphid populations collected from cultivated wheat and barley, Hordeum vulgare L. We conducted field surveys from May through June 2002 and August 2003 from seven counties within Texas, Kansas, Nebraska, and Wyoming. Based upon a foliar chlorosis damage rating, three new Russian wheat aphid biotypes were identified, one of which was virulent to all characterized sources of Russian wheat aphid resistance. The future success of Russian wheat aphid resistance breeding programs will depend upon the continual monitoring of extant biotypic diversity and determination of the ecological and genetic factors underlying the development of Russian wheat aphid biotypes.

Animals↗

[Symbiotic relationships between ants and aphids].

The classical and contemporary literature devoted to symbiotic relationships between ants and aphids was analyzed. The most studied issues are: the totality of morphological and anatomical adaptations of partners to symbiosis, different degrees of aphids myrmecophilia, benefits and costs of insects during interaction. A lot of issues are still discussible. For instance, it is unknown whether the trophobiotical relationships between ants and aphids are mutualistic or intermediate between mutualism and exploitation. To clarify this vague point the study of symbionts behaviour is of great importance, however, it remains practically non-investigated either for aphids or ants. It is known only that nonmyrmecophilous aphids living within galls have rather complicated behaviour--their colonies are guarded with specialized castes of aphids-soldiers. As for the ants, till nowadays trophobionts (individuals looking after the aphids), which make up special functional group within ants family, have been described as passive foragers being occupied only with the honeydew collection and transportation. The recent investigations revealed that ants use behavioural schemes with different degree of complication interacting with aphids: from lonely foraging to professional specialization in working groups.

Adaptation, Physiological↗

Effect of aphid-infested plants on the host location and learning behaviour of the parasitoid Aphelinus abdominalis.

Aphelinus abdominalis Dalman (Hymenoptera: Aphelinidae) is a generalist parasitoid of cereal (e.g. Sitobion avenae) and greenhouse aphids (e.g. Macrosiphum euphorbiae). It differs from Aphidius spp. (Hymenoptera: Braconidae) by its robust constitution, longevity and its habit to use aphids not only as hosts for its progeny but also as prey (host feeding). In addition, A. abdominalis does not evoke Aphidius-characteristic defence responses by attacked aphids as this wasp approaches its hosts very carefully and then inserts its ovipositor nearly unnoticed by the aphids. In order to obtain more information on the host location behaviour of A. abdominalis, the effect of aphid-infested plants on its orientation and learning behaviour after mummy emergence was first studied in a series of Y-tube olfactometer experiments. When females reared on S. avenae were offered a 2 h foraging experience on M. euphorbiae-infested sweet pepper (Capsicum annuum) and then had the choice between the odour of uninfested sweet pepper plants and plants infested by aphid hosts (M. euphorbiae), they were significantly attracted to the odour of infested plants. Naive A. abdominalis, i.e. females that prior to the experiments were not offered any contact with a plant-host complex (PHC) after their emergence from S. avenae mummies and hence had no chance to learn the odour of the PHC, were not attracted by aphid-infested plants. The effects of adult experience on long-range host location by A. abdominalis (again reared on S. avenae) were then investigated in wind tunnel experiments by using M. euphorbiae-infested and uninfested sweet pepper and aubergine (Solanum melongena) plants. Females experienced with one of the offered PHCs significantly preferred the odour of the learnt PHC to that of the different PHC. A. abdominalis females obviously learn specific volatile signals emitted by aphid-infested plants for long-range host location.

Animals↗

Role of host plant alternation in the precocity and intensity of parasitoid activity on of cereal aphids.

Integrated pest management programs tend to reduce the chemical input by enhancing the development of biological control. Cereal aphids cause important damages to winter wheat in Europe but are currently under the pressure of several parasitoid species (Braconidae: Aphidiinae). Previous ecological studies have reported the existence of an asynchrony between aphid and parasitoid populations in early spring in cereal cultures. Here, we tested the presence of rose bushes (Rosa rugosa) as a host plant for alternative aphid-host. Aphid and parasitoid densities were recorded for two years using the plant cutting sampling method. The main results were: (i) rose bushes constitute a potential reservoir of alternative aphid hosts species for a number of parasitoid and predator species, (ii) rose aphids appear earlier in spring and were more abundant than the aphids on wheat, (iii) parasitism activity in rose bushes is synchronized with cereal fields, however (iv) rose bushes management did not induce a decrease of cereal aphid population.

Animals↗

An aphid-borne bacterium allied to the secondary symbionts of whitefly.

Bacterial 16S rDNA amplified by PCR from the pea aphid Acyrthosiphon pisum included a sequence with >98% similarity to secondary symbionts in the whitefly Bemisia tabaci. The 'pea aphid Bemisia-like bacterium' (PABS) and B. tabaci secondary symbionts are estimated to have diverged 17-34 million years ago, a time considerably more recent than the common ancestor of aphids and whitefly and suggestive of horizontal transmission of this bacterial lineage. PABS was scored in both the gut and ovaries of aphids by PCR and identified as a small rod by in situ hybridisation. PABS was not universal in pea aphids: 2/3 laboratory strains and 13/35 of field aphids were PABS-positive. It is suggested that the incidence of PABS in pea aphids is determined by the balance between loss (processes may include occasional failure of vertical transmission and selection against PABS-positive aphids) and horizontal transfer between insects.

Journal Article↗

Growth Kinetics of the Endosymbiont Buchnera aphidicola in the Aphid Schizaphis graminum.

The aphid Schizaphis graminum is dependent on its prokaryotic endosymbiont, Buchnera aphidicola. As a means of determining B. aphidicola numbers during the growth cycle of the aphid we have used the quantitative PCR to measure the number of copies of rrs (the gene coding for 16S rRNA, which is present as one copy in the B. aphidicola genome). In addition we have measured the aphid wet weight and the DNA and protein content. The results indicate an approximately parallel (23- to 31-fold) increase of these properties during the period of aphid growth. A 1-day-old aphid (24 mug [wet weight]) has 0.2 x 10 copies of rrs, while a 9-day-old aphid (497 mug [wet weight]) has 5.6 x 10 copies. The coupling of endosymbiont and aphid growth is consistent with the requirement of the endosymbiont for growth and reproduction of the aphid.

Journal Article↗

Partitioning of Symbiotic Bacteria between Generations of an Insect: a Quantitative Study of a Buchnera sp. in the Pea Aphid (Acyrthosiphon pisum) Reared at Different Temperatures.

The population of symbiotic Buchnera bacteria in parthenogenetic females of the pea aphid Acyrthosiphon pisum was determined by quantitative hybridization of a DNA probe (groESL) to aphid homogenates. The aphids bore 1 x 10(sup7) to 2 x 10(sup7) bacterial cells per mg (fresh weight). In teneral aphids (i.e., aphids that had moulted to adulthood but that had not initiated reproduction), >75% of the bacteria were in the embryos, and the density of bacteria in the embryos was consistently greater than that in the maternal tissues. The bacterial density in teneral aphids increased from 1.3 x 10(sup7) to 2.0 x 10(sup7) cells mg (fresh weight) of aphids(sup-1) with temperature between 15 and 25(deg)C. This variation could be attributed to a temperature-dependent increase in both the density of bacteria in the embryos and embryo content of the aphids.

Journal Article↗

Nutritional interactions in insect-microbial symbioses: aphids and their symbiotic bacteria Buchnera.

Most aphids possess intracellular bacteria of the genus Buchnera. The bacteria are transmitted vertically via the aphid ovary, and the association is obligate for both partners: Bacteria-free aphids grow poorly and produce few or no offspring, and Buchnera are both unknown apart from aphids and apparently unculturable. The symbiosis has a nutritional basis. Specifically, bacterial provisioning of essential amino acids has been demonstrated. Nitrogen recycling, however, is not quantitatively important to the nutrition of aphid species studied, and there is strong evidence against bacterial involvement in the lipid and sterol nutrition of aphids. Buchnera have been implicated in various non-nutritional functions. Of these, just one has strong experimental support: promotion of aphid transmission of circulative viruses. It is argued that strong parallels may exist between the nutritional interactions (including the underlying mechanisms) in the aphid-Buchnera association and other insect symbioses with intracellular microorganisms.

Journal Article↗