Aphids exhibit seasonally alternating asexual and sexual reproductive modes. Different morphs are produced throughout the life cycle. To evaluate morph-specific fitness during reproductive switching, holocyclic Sitobion avenae were induced continuously under short light conditions, and development and reproduction were compared in each morph. Seven morphs, including apterous and alate virginoparae, apterous and alate sexuparae, oviparae, males, and fundatrices, were produced during the life cycle. The greatest proportions of sexuparae, oviparae, males, and virginoparae were in the G1, G2, G3, and G4 generations, respectively. Regardless of asexual or sexual morphs, alate morphs exhibited a marked delay in age at maturity compared with that of apterous morphs. Among the alate morphs, males had the longest age at maturity, followed by sexuparae and virginoparae. Among the apterous morphs, sexuparae were older at maturity than the fundatrices, virginoparae, and oviparae. The nymphs of each morph had equal survival potentials. For the same wing morphs, apterous sexuparae and oviparae exhibited substantial delays in the pre-reproductive period and considerable reductions in fecundity, compared with those of apterous virginoparae and fundatrices, whereas alate sexuparae and alate virginoparae had similar fecundity. The seven morphs exhibited Deevey I survivorship throughout the life cycle. These results suggest that sexual production, particularly in males, has short-term development and reproduction costs. The coexistence of sexual and asexual morphs in sexuparae offspring may be regarded as an adaptive strategy for limiting the risk of low fitness in winter.

Several studies have shown the negative effects of mature plant resistance (MPR) on aphids in sugar beet, which is correlated to the formation of black deposits in their stomach. However, the underlying mechanism of MPR still needs to be elucidated, by understanding the toxicity effects of MPR on aphids and the role of the plant phenological stage and the environment. Here, we report that MPR in sugar beet does not only affect Myzus persicae mortality rate and the formation of a black deposit in the aphid stomach, but also aphid fecundity and behaviour. In addition, experiments in climate-controlled and field settings showed quantitative variation in MPR to M. persicae between six genotypes of sugar beet. Our results indicate that environmental effects, such as temperature, play a major role in MPR and underscore the importance of proper climate-controlled experiments for investigating MPR. In climate-controlled experiments, 83.3% of aphids on old leaves developed a black deposit, in contrast to only 16.8% of aphids on young leaves. This shows that not only plant age, but also leaf age plays a major role in the intensity of MPR. Further research will be needed to identify the underlying mechanism, before MPR can be used as a viable and sustainable solution to aphid pests in sugar beet.

Plants have a variety of defense mechanisms that are often induced following attacks by herbivores; this benefits those plants by decreasing performance or preference of herbivores that attack the plants later. We investigated the effects of previous exposure of plants to the safflower aphid, Uroleucon carthami, cotton bollworm, Helicoverpa armigera, and mechanical wounding on subsequent safflower aphid infestations using commercial safflower (Carthamus tinctorius) cultivars and wild safflower species (C. oxyacantha). The experiments were conducted in a greenhouse with two treatments: previously induced plants via direct herbivory or mechanical wounding, and control plants that had never experienced herbivory. To test the performance of safflower aphid on different plant treatments, five unwinged aphids were placed on each plant and allowed to reproduce for 14 days. Finally, the total numbers of aphids on each plant were counted and the percentage of produced winged individuals was calculated. The number of aphids on plants that were previously infested or injured was significantly lower than in control plants. Percentage of winged aphids was significantly higher on induced plants, which is an indicator for unsuitable conditions. Also, significant increase in total phenolic content and hydrogen peroxide was observed in induced plants, showing that the levels of these compounds were either treatment, cultivar and/or genotype × treatment dependent, highlighting the specificity of these interactions. Overall, among the safflower cultivars the lowest number of aphids and the highest percentage of winged aphid individuals were observed on Mahali-Isfahan cultivar and wild safflower, showing that this cultivar is more sensitive to herbivory and/or responds to it more than other cultivars. These findings could contribute to a better utilization of induced defense in the integrated pest management of safflower fields.

Secondary metabolites are central to understanding the evolution of plant–animal interactions. Direct effects on phytophagous animals are well-known, but how secondary consumers adjust their behavioural and physiological responses to the herbivore's diet remains more scarcely explored for some metabolites. Caffeine is a neuroactive compound that affects both the behaviour and physiology of several animal species, from humans to insects. It is an alkaloid present in nectar, leaves and even sap of numerous species of plants where it plays a role in chemical defences against herbivores and pathogens. Caffeine effects have been overlooked in generalist herbivores that are not specialized in coffee or tea plants. Using a host–parasitoid system, we show that caffeine intake at a relatively low dose affects longevity and fecundity of the primary consumer, but also indirectly of the secondary one, suggesting that this alkaloid and/or its effects can be transmitted through trophic levels and persist in the food chain. Parasitism success was lowered by ≈16% on hosts fed with caffeine, and parasitoids of the next generation that have developed in hosts fed on caffeine showed a reduced longevity, but no differences in mass and size were found. This study helps at better understanding how plant secondary metabolites, such as caffeine involved in plant–animal interactions, could affect primary consumers, could have knock-on effects on upper trophic levels over generations, and could modify interspecific interactions in multitrophic systems.

Aphids are phloem-feeding insects that cause economic losses to crops globally. Whilst aphid interactions with susceptible plants and partially resistant genotypes have been well characterized, the interactions between aphids and non-host species are not well understood. Unravelling these non-host interactions can identify the mechanisms which contribute to plant resistance. Using contrasting aphid-host plant systems, including the broad host range pest Myzus persicae (host: Arabidopsis; poor-host: barley) and the cereal pest Rhopalosiphum padi (host: barley; non-host: Arabidopsis), we conducted a range of physiological experiments and compared aphid settling and probing behaviour on a host plant vs either a non-host or poor-host. In choice experiments, we observed that around 10% of aphids selected a non-host or poor-host plant species after 24 h. Using the Electrical Penetration Graph technique, we showed that feeding and probing behaviours differ during non-host and poor-host interactions when compared with a host interaction. In the Arabidopsis non-host interaction with the cereal pest R. padi aphids were unable to reach and feed on the phloem, with resistance likely residing in the mesophyll cell layer. In the barley poor-host interaction with M. persicae, resistance is likely phloem-based as phloem ingestion was reduced compared with the host interaction. Overall, our data suggest that plant resistance to aphids in non-host and poor-host interactions with these aphid species likely resides in different plant cell layers. Future work will take into account specific cell layers where resistances are based to dissect the underlying mechanisms and gain a better understanding of how we may improve crop resistance to aphids.

The corn leaf aphid, Rhopalosiphum maidis (Fitch) (Hemiptera: Aphididae), is an important pest of corn, but no corn genotypes resistant to R. maidis are commercially available. Although the ability of silicon to induce plant resistance against some insects is known, the effect of silicon on R. maidis and in corn hybrids with different levels of constitutive resistance is still unknown. This study sought to determine the constitutive resistance of corn hybrids to R. maidis and silicon resistance induction in hybrids with different degrees of constitutive resistance. Field experiments with natural infestations of aphids were conducted in three locations in Brazil (Patos de Minas, Araguari, and Tupaciguara). Greenhouse trials were also used to evaluate the effect of varietal resistance on aphid population growth and identify resistant and susceptible genotypes. Aphid resistance induced by silicon was determined with resistant and susceptible corn hybrids. In the field, the corn hybrids BM8850, AS1625PRO, and DKB310PRO had the greatest proportion of plants infested by R. maidis in all three localities. The hybrids P30F53H, STATUS VIP, BM9288, DAS2B587HX, DKB175PRO, AS1633PRO, and DKB390PRO2 were the least infested in Patos de Minas and Araguari, and P30F53H was the least infested in Tupaciguara. When antibiosis effects were evaluated by aphid population growth, the hybrids AG7088PRO3 and DKB310PRO2 were susceptible, while P30F53YH was resistant. When natural aphid infestation was evaluated, wherein the effects of antibiosis and non-preference could not be discriminated, soil applications of silicon-induced resistance to R. maidis in both susceptible and constitutively resistant corn hybrids.

A careful measure of fitness represents a crucial target in crop pest management and becomes fundamental considering extremely prolific insects. In the present paper, we describe a standardized rearing protocol and a bioinformatics tool to calculate aphid fitness indices and invasiveness starting from life table data. We tested the protocol and the bioinformatic tool using six Myzus persicae (Sulzer) asexual lineages in order to investigate if karyotype rearrangements and ecotype could influence their reproductive performances. The tool showed that different karyotypes do not influence adaptive success and put in evidence a marked invasive potential of the M. persicae lineage 64. The presence of a similar fitness rate of 33H and 7GK asexual lineages (both possessing intra-individual karyotype variations) in respect to the asexual lineage 1 (with a standard karyotype) represents an important demonstration of the potentiality of holocentric chromosomes to reduce the effects of chromosome rearrangements.

The aphid species Sitobion avenae and Rhopalosiphum padi are the most important pests in wheat growing regions of many countries. In this study, we investigated the sublethal effects of imidacloprid on fecundity, longevity, and enzyme activity in both aphid species by comparing 3-h exposure for one or three generations. Our results indicated that 3-h exposure to sublethal doses of imidacloprid for one generation had no discernible effect on the survival, fecundity, longevity, or enzyme activity levels of aphids. However, when pulse exposures to imidacloprid were sustained over three generations, both fecundity and longevity were significantly decreased in both S. avenae and R. padi. Interestingly, the fecundity of R. padi had almost recovered by the F5 generation, but its longevity was still deleteriously affected. These results indicated that R. padi laid eggs in shorter time lags and has a more fast resilience. The change in reproduction behavior may be a phenomenon of R. padi to compensate its early death. If this is stable for the next generation, it means that the next generation is more competitive than unexposed populations, which could be the reason underlying population outbreaks that occur after longer-term exposure to an insecticide. This laboratory-based study highlights the sublethal effects of imidacloprid on the longevity and fecundity of descendants and provides an empirical basis from which to consider management decisions for chemical control in the field.

Rapid, precise and timely identification of invasive pest insects such as aphids is important and a challenge worldwide due to their complex life cycles, parthenogenetic reproduction, sex and colour morphs. In this respect, DNA barcoding employing a 658 bp fragment of 5′ region of the mitochondrial cytochrome oxidase I (CO-I) gene is an effective tool in addressing the above. In the present study, we employed CO-I for discriminating 142 individuals representing 32 species of aphids from India. Sequence analyses revealed that the intraspecific and interspecific distances ranged from zero to 3.8% and 2.31 to 18.9%, respectively. In addition, the study also showed for the first time the prevalence of three cryptic species, namely Brevicoryne brassicae (Linnaeus), Hyperomyzus carduellinus (Theobald) and Brachycaudus helichrysi (Kaltenbach) from India. Our work has clearly demonstrated that DNA barcoding is an efficient and accurate method for identification of aphid species (including cryptic species), an approach that potentially could play an important role in formulating viable pest management strategies, more especially biocontrol.

Clones have been in the forefront of biological interest for many years. Even so, open discussions continue to surround the concept of clonality, which has been recently much debated in the scientific literature, both in terms of philosophical meaning as well as empirical determination. Philosophically, the clone is the horizontally produced lineage from a single fertlized egg (e.g. mammals by division of the fertilized egg and representing a single generation) or vertically produced offspring (e.g. aphids representing different successive generations) from a single asexual stem mother (originally for a particular lineage, following hatching of the overwintering sexual egg in the spring); empirically, the aspect of genetic fidelity is also considered important, so-called clones being assumed to have an identical genome among clone mates. In reality of course, such members of a clonal lineage must differ at various regions of the genome, since mutation is a fundamental property of the DNA itself. Yet few studies have so far set out to show this empirically in eukaryotic organisms, which indulge in periods of asexual reproduction, sometimes, as in aphids, over many generations. In the present study, we have investigated asexual lineages of the grain aphid, Sitobion avenae (F.), a global pest of cereals, over five successive generations employing AFLP-PCR molecular techniques. Our main interest was to see how much variation was present in the early generations and if this variation was transmitted through the asexual lineages. By monitoring AFLP-based polymorphisms, we show that, in this aphid species, of a total of 110 individuals from two lineages tested (termed SA and SB), random mutations (band deletions, more rarely additions) were apparent from the third generation onwards, and although some mutations were found to be transmitted transgenerationally, others were rarely transmitted through the particular lineages they were detected in. Using Arlequin v. 2.0, average gene diversity within the lineages was found to be 0.024 ± 0.013 and 0.031 ± 0.016 for SA and SB, respectively. It was also found from the rearing of the lineages that one lineage, SA, was more fecund than the other lineage, SB, over the five generations (N = 818 vs. N = 358 total stem mothers plus nymphs for the two lineages, respectively).

Walking speeds were calculated for nine clones of the peach potato aphid Myzus persicae collected from three countries along a latitudinal cline of its European distribution from Sweden to Spain (Sweden, UK and Spain), and the effects of collection origin and intra and intergenerational acclimation were investigated. Walking speeds declined with decreasing temperature, with maximum performance at temperatures closest to acclimation temperature (fastest median walking speed of 5.8 cm min–1 was recorded for clone UK 3, collected from the UK, at 25°C after acclimating to 25°C for one generation). Following acclimation at both 20°C and 25°C, walking ceased (as indicated by median walking speeds of 0.0 cm min–1) at temperatures as high as 7.5°C and 12.5°C. However, acclimation at 10°C enabled mobility to occur to temperatures as low as 0°C. There was no relationship between mobility and latitude of collection, suggesting that large scale mixing of aphids may occur across Europe. However, clonal variation was suggested, with clone UK 3 outperforming the majority of other clones across all temperatures at which mobility was maintained following acclimation at 10°C for one and three generations and at 25°C for one generation. The Scandinavian clones consistently outperformed their temperate and Mediterranean counterparts at the majority of temperatures following acclimation for three generations at 25°C.

The molecular detection of predation is a fast growing field, allowing highly specific and sensitive detection of prey DNA within the gut contents or faeces of a predator. Like all molecular methods, this technique is prone to potential sources of error that can result in both false positive and false negative results. Here, we test the hypothesis that the use of suction samplers to collect predators from the field for later molecular analysis of predation will lead to high numbers of false positive results. We show that, contrary to previous published work, the use of suction samplers resulted in previously starved predators testing positive for aphid and collembolan DNA, either as a results of ectopic contamination or active predation in the collecting cup/bag. The contradictory evidence for false positive results, across different sampling protocols, sampling devices and different predator-prey systems, highlights the need for experimentation prior to mass field collections of predators to find techniques that minimise the risk of false positives.

In the interaction between two ecologically-associated species, the population structure of one species may affect the population structure of the other. Here, we examine the population structures of the aphid Metopeurum fuscoviride, a specialist on tansy Tanacetum vulgare, and its specialist primary hymenopterous parasitoid Lysiphlebus hirticornis, both of which are characterized by multivoltine life histories and a classic metapopulation structure. Samples of the aphid host and the parasitoid were collected from eight sites in and around Jena, Germany, where both insect species co-occur, and then were genotyped using suites of polymorphic microsatellite markers. The host aphid was greatly differentiated in terms of its spatial population genetic patterning, while the parasitoid was, in comparison, only moderately differentiated. There was a positive Mantel test correlation between pairwise shared allele distance (DAS) of the host and parasitoid, i.e. if host subpopulation samples were more similar between two particular sites, so were the parasitoid subpopulation samples. We argue that while the differences in the levels of genetic differentiation are due to the differences in the biology of the species, the correlations between host and parasitoid are indicative of dependence of the parasitoid population structure on that of its aphid host. The parasitoid is genetically tracking behind the aphid host, as can be expected in a classic metapopulation structure where host persistence depends on a delay between host and parasitoid colonization of the patch. The results may also have relevance to the Red Queen hypothesis, whereupon in the ‘arms race’ between parasitoid and its host, the latter ‘attempts’ to evolve away from the former.

Polyacrylamide gel electrophoresis of enzymes (carboxylesterases) was used for the first time to monitor rates of parasitism in airborne alate (winged) grain aphid, Sitobion avenae (F.) population samples collected by suction trapping in Hertfordshire, UK. Using previously described electrophoretic ‘keys’, the species of hymenopterous parasitoids present in individual aphids were identified and found to be Aphidius ervi (Haliday) and/or Aphidius rhopalosiphi (De Stephani Perez) (Braconidae). Entomophthoralean fungal infection was also detected using this approach. Aphidiid wasp parasitism was detected from early June to mid-August and fungal infection from late June to late July. The results are discussed in relation to parasitoid population structure and dynamics, especially (i) the fact that winged aphids passively transport the early stages of their braconid parasitoids and fungal pathogens, potentially to newly-founded colonies, which may directly impact on the dual aphid-parasitoid populations genetics; and (ii) the approach used to collect and assay parasitised and fungal infected aphids involving both suction trapping and electrophoretic testing may have potential in assessing the level and efficacy of these biological control agents in integrated pest management (IPM) schemes to combat cereal aphid outbreaks.

Within-crop habitat manipulations have the potential to increase the biological control of pests in horticultural field crops. Wildflower strips have been shown to increase the abundance of natural enemies, but there is little evidence to date of an impact on pest populations. The aim of this study was to determine whether within-crop wildflower strips can increase the natural regulation of pests in horticultural field crops. Aphid numbers in plots of lettuce grown adjacent to wildflower strips were compared with those in plots grown in the absence of wildflowers. The presence of wildflower strips led to a decrease in aphid numbers on adjacent lettuce plants during June and July, but had less impact in August and September. The decrease in aphid numbers was greatest close to the wildflower strips and, the decrease in aphid numbers declined with increasing distance from the wildflower strips, with little effect at a distance of ten metres. The main natural enemies found in the crop were those that dispersed aerially, which is consistent with data from previous studies on cereal crops. Analysis and interpretation of natural enemy numbers was difficult due to low recovery of natural enemies, and the numbers appeared to follow changes in aphid abundance rather than being directly linked to the presence of wildflower strips. Cutting the wildflower strips, to remove floral resources, had no impact on the reduction in aphid numbers achieved during June and July, but decreased the effect of the wildflower strips during August and September. The results suggest that wildflower strips can lead to increased natural regulation of pest aphids in outdoor lettuce crops, but more research is required to determine how this is mediated by natural enemies and how the impact of wildflower strips on natural pest regulation changes during the growing season.

In cyclical parthenogens, clonal diversity is expected to decrease due to selection and drift during the asexual phase per number of asexual generations. The decrease in diversity may be counteracted by immigration of new genotypes. We analysed temporal variation in clonal diversity in colonies of the monophagous tansy aphid, Macrosiphoniella tanacetaria (Kaltenbach), sampled four times over the course of a growing season. In a related field study, we recorded aphid colony sizes and the occurrence of winged dispersers throughout the season. The number of colonies increased from April, when asexual stem mothers hatched from the sexually produced eggs, to the end of June. The proportion of colonies with winged individuals also increased over this period. After a severe reduction in colony sizes in late summer, a second expansion phase occurred in October when sexuals were produced. At the season's end, the only winged forms were males. A linked genetic study showed that the number of microsatellite multilocus genotypes and genetic variability assessed at three polymorphic loci per colony decreased from June to October. Overall, the relatedness of wingless to winged individuals within colonies was lower than average relatedness among wingless individuals, suggesting that winged forms mainly originated in different colonies. The results demonstrate that patterns of genetic diversity within colonies can be explained by the antagonistic forces of clonal selection, migration and genetic drift (largely due to midsummer population bottlenecks). We further suggest that the males emigrate over comparatively longer distances than winged asexual females.

In Europe, aphids contribute significantly to the so-called ‘aerial plankton’ during the spring to autumn months (growing season), although individual flight behaviour has been found, especially from molecular ecological studies, to be species-specific in terms of migratory range (ambit). Many of these species individuals may be assumed to be clonal in origin, that is, derived from a single asexual foundress. We are presently studying two specialist aphid species on Tansy, Tanacetum vulgare L. from samples collected in Jena, Germany – Macrosiphoniella tanacetaria (Kaltenbach) and Metopeurum fuscoviride Stroyan, using microsatellite markers. On plotting the number of sets of different multilocus genotypes or MLGs (i.e. multiple clonal repeats: 1, 2, 3 copies, etc.), against the frequency of their occurrence, a negative exponential relationship was found, with populations of both species consisting mostly of single (i.e. unique) or low number repeats rather than larger multiple copy (clonal) MLG repeats. To test this further, microsatellite data collected from a previous study on M. tanacetaria in Jena in the year 2000 and on samples of the Grain aphid, Sitobion avenae (F.), collected in the UK in 1997/8, the latter both in the field and from 12.2 m high suction traps, were examined in the same way. Again, similar relations were found, with most MLGs occurring as unique or low copy number repeats. The data are briefly discussed in the light of our evidence, as well as that of other similar studies on other aphid species, relating aphid molecular genetic data to aphid life cycle, behaviour and ecology.

The population of peach-potato aphid, Myzus persicae (Sulzer), in Scotland comprises large numbers of a few superclones with much smaller numbers of other clones, and the reason for their differential success has yet to be elucidated. In the current study, the reproduction of lineages derived from these clones was measured by counting the numbers of offspring produced by a one-day-old nymph after 15 days. This was measured on four plant species, including local agricultural hosts and at two different temperatures (14 and 18°C). There were significant differences in clonal lineage reproduction on different hosts and at different temperatures and amongst clonal lineages on the same hosts at the same temperature. Lineages of local insecticide sensitive clones did not have the best reproductive potential; instead, a recently introduced clonal lineage carrying MACE insecticide resistance was the best reproducer. The clonal lineage with the lowest reproductive potential also carried insecticide resistance, but this was kdr. A lineage from a local insecticide-sensitive clone was the least affected by reduced temperature. There was evidence of host plant specialisation in some of the clonal lineages.

Generalist insect predators can significantly impact the dynamics of pest populations; and, using alternative prey, they can rapidly establish in disturbed agroecosystems. However, indirect interactions between prey can occur, leading to either increased or decreased predation on focal prey. The present paper demonstrates how alternative prey can disrupt predation by the hemipteran Orius insidiosus on the soybean aphid Aphis glycines via short-term indirect interactions. We used laboratory microcosms to measure the impact of the predator on the population growth of the aphid in the presence of alternative prey, soybean thrips Neohydatothrips variabilis, and we characterized the foraging behaviour of the predator to assess prey preference. We showed that O. insidiosus predation on aphids was reduced in the presence of thrips and that this positive impact on aphids increased as thrips density increased. Results from the behavioural experiment support the hypothesis of a prey preference toward thrips. When prey-pest ratio is aphid-biased, short-term apparent commensalism between prey occurs in favour of the most abundant prey (aphids) with no switching behaviour appearing in O. insidiosus. These results demonstrate that potential indirect interactions should be taken into account when considering O. insidiosus as a biocontrol agent against the soybean aphid.

The study of intraspecific variation with respect to host plant utilization in polyphagous insects is crucial for understanding evolutionary patterns of insect-plant interactions. Aphis gossypii (Glover) is a cosmopolitan and extremely polyphagous aphid species. If host plant species or families constitute selective regimes to these aphids, genetic differentiation and host associated adaptation may occur. In this study, we describe the genetic structure of A. gossypii collected in six localities in Tunisia on different vegetable crops, on citrus trees and on Hibiscus. The aim was to determine if the aphid populations are structured in relation to the host plants and if such differentiation is consistent among localities. The genetic variability of A. gossypii samples was examined at eight microsatellite loci. We identified only 11 multilocus genotypes among 559 individuals. Significant deviations from Hardy-Weinberg equilibrium, linkage disequilibria and absence of recombinant genotypes, confirmed that A. gossypii reproduces by continuous apomictic parthenogenesis. Genetic differentiation between localities was not significant, whereas a strong differentiation was observed between host plant families (0.175<FST<0.691). The great majority of aphids exhibited one of three predominant multilocus genotypes that were repeatedly and respectively associated to the three plant families, Cucurbitaceae, Solanaceae and Rutaceae, demonstrating host specialization in A. gossypii. These specialized genotypes were simultaneously found with other clones on Hibiscus, suggesting that this perennial host could act as a refuge plant between two vegetable crop seasons.